United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

Proceedings

RMRS-P-67

July 2013

Merging Science and 

Management in a Rapidly 

Changing World:

Biodiversity and Management of the 

Madrean Archipelago III

May 1–5, 2012

Tucson, Arizona
Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane G.; Collins, Loa C., comps. 2013. Merging 

science and management in a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III and 7th Conference on Research and Resource Management in the Southwestern Deserts; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station. 593 p.

Abstract ___________________________________________

The Madrean Archipelago or Sky Islands region of the southwestern United States and northern Mexico is 

recognized for its unique biological diversity, natural beauty, and cultural heritage. This 2012 conference brought 

together scientists, managers, students, and other interested parties from the United States and Mexico to share 

their knowledge and passion about the region and to identify needs and creative solutions for existing and emerging problems. More than 300 people attended the conference including a large and energetic contingent from 

northern Mexico. The conference provided a forum to update the state-of-knowledge that has evolved since the first 

conference in 1994 and the second conference in 2004. It also provided a setting for the formation of new friendships and partnerships. These proceedings contain 80 of the 190 oral and poster presentations and all abstracts 

from the plenary sessions and the concurrent sessions. Abstracts in Spanish are included. Topics include climate 

change in the Sky Island Region, southwestern cienegas, the Northern Jaguar Reserve, amphibian conservation, 

biodiversity of plants and animals, fire effects, grasslands, and human impacts, and tools. The conference hosted 

a symposium about Santa Cruz River Watershed Conservation. A lively open forum at the end of the conference 

generated a list of future scientific and management needs for the Madrean Archipelago and a commitment to 

increase international cooperation. Mexican participants graciously offered to host the next conference, Speakers 

stressed the importance for all interested parties to collaborate--to work side-by-side and constantly inform one 

another about relevant research, timely events, and cross-pollination opportunities throughout the region. 

Keywords: Madrean Archipelago, Sky Islands, southwestern United States, northern Mexico, natural environment, 

fauna, flora, research, management, biodiversity, climate change

You may order additional copies of this publication by sending your 

mailing information in label form through one of the following media. 

Please specify the publication title and number.

Telephone (970) 498-1392

FAX (970) 498-1122

E-mail rschneider@fs.fed.us

Web site http://www.fs.fed.us/rm

Mailing Address Publications Distribution

Rocky Mountain Research Station

240 West Prospect Road

Fort Collins, CO 80526

Cover photo by Erik Enderson.
iii

Acknowledgments

We thank Sky Island Alliance for generously providing the funding to print these proceedings. 

Sky Island Alliance is a grassroots organization dedicated to the protection and restoration of the 

rich natural heritage of native species and habitats in the Sky Island region of the southwestern 

United States and northwestern Mexico. For over 20 years, Sky Island Alliance has worked across 

the region to establish protected areas, connect wildlife pathways, restore healthy landscapes, and 

promote public appreciation of the beauty and richness of this area. Connecting everyday people 

with hands-on science and conservation action, Sky Island Alliance is internationally recognized 

as a leader in conservation, building bridges between communities and across political boundaries 

in order to explore, restore, connect, and protect this special place.

Organizing Partners

Coronado National Forest

Cuenca los Ojos Foundation

Sky Island Alliance

Sonoran Institute

The Nature Conservancy

Universidad de Sonora

University of Arizona

 School of Natural Resources and the Environment

USDA Agricultural Research Service

USDA Forest Service, Rocky Mountain Research Station

US Fish & Wildlife Service

 Desert Landscape Conservation Cooperative

US Geological Survey

Sponsors

Arizona Native Plant Society

Arizona-Sonora Desert Museum

Boyce Thompson Arboretum

Comisión de Ecología y Desarrollo Sustentable 

 del Estado de Sonora

Coronado National Forest

National Fish & Wildlife Foundation 

Nina Mason Pulliam Charitable Trust

Sky Island Alliance

Sonoran Institute

T&E Foundation

The Nature Conservancy

Universidad de Sonora

USDA Agricultural Research Service

US Fish & Wildlife Service

USGS Amphibian Research & Monitoring Initiative
iv
v

Merging Science and Management in a 

Rapidly Changing World:

Biodiversity and Management of the Madrean Archipelago III

 and 

7th Conference on Research and Resource Management in the 

Southwestern Deserts

May 1–5, 2012

Tucson, Arizona

Compilers

Gerald J. Gottfried

Peter F. Ffolliott

Brooke S. Gebow

Lane G. Eskew

Loa C. Collins
vi
vii

Preface

Dale S. Turner

The Nature Conservancy, Tucson, Arizona

Cecil R. Schwalbe

U.S. Geological Survey, Tucson, Arizona

Spanning four states in two countries, the Madrean Archipelago or Sky Island region has been 

a puzzle and an inspiration to researchers and managers for more than a century. Gradients of 

species diversity include a north/south axis for influences from the Sierra Madre Occidental and 

Rocky Mountains, an east/west axis for influences from the Chihuahuan and Sonoran Deserts, 

and vertical gradients from valley grasslands or desert scrub to mountain-top communities. 

Overlying that is a fascinating cultural and political diversity created by an international boundary that bisects the region. 

The first Conference on Biodiversity and Management of the Madrean Archipelago, held in 

1994, focused on defining the region. The assembled experts described different aspects of this 

collection of “sky islands” that form an “archipelago” off the “mainland” of the Sierra Madre. 

From about 100 presentations, we learned that this region is special to people from many disciplines and many walks of life. We recognized the need to collaborate across disciplines and 

across borders.

The 1994 conference served several important functions. It established the state of our knowledge about the region, and led to a proceedings volume (DeBano and others 1995) that became 

a widely cited reference. Beyond that, it created a shared understanding of the similarities and 

differences across this distinct biogeographic area.

At the second conference, in 2004, we went deeper into understanding the region. We focused 

on connections: between mountains and valleys, between people and the landscape. Through 160 

presentations, we looked at big patterns of diversity, in both species and ecological processes 

(Gottfried and others 2005). 

This volume of proceedings resulted from the third conference in this series, held in 2012. 

This time we had 190 presentations, with a stronger representation from Mexico than ever 

before. The presentations, and the papers in this volume, reflect various attempts to adapt to 

a region that is undergoing rapid and accelerating change. Changes along the border between 

Sonora and Arizona have disrupted the movement of animals and damaged the sense of shared 

culture across that border. Technology is evolving, giving us new insights. Species distributions 

are shifting, challenging our best efforts at conservation. And the climate is changing around 

us. 

The 1994 conference had one talk about the anticipated effects of climate change. In 2004 

we had seven. This year we have 21 climate-change talks, and yet barely skim the surface of 

the work that is underway. 

We’ve also had an explosion of information about biodiversity in this region. At previous 

conferences, many people recognized big gaps in our knowledge about the mountains of Sonora. Now there is a great new book summarizing biodiversity of Sonora (Molina and Van 

Devender 2010). And there is the Madrean Archipelago Biodiversity Assessment, a bi-national 

effort to compile everything known about species distributions in the region and to conduct 

new biological inventories. 

In 1994, jaguars were thought to be extinct in Arizona. That changed in 1996, with photographs 

of two different, apparently healthy jaguars in the Peloncillo and Baboquivari Mountains of 

Arizona. The 2012 conference included a half-day symposium to discuss what’s been learned in 
viii

the new jaguar reserve in Sonora, new models for jaguar habitat use, and how that knowledge 

can be applied for jaguar conservation across the Sky Island region.

Clear evidence of the cooperation and collaboration by many organizations and individuals to achieve a common goal of strong biodiversity and sound management of the Madrean 

Archipelago is apparent by the members of the conference organizing committee:

• Cecil Schwalbe, Conference Chair, U.S. Geological Survey

• Dale Turner, Program Chair, The Nature Conservancy

• Acasia Berry, Sky Island Alliance

• Emily Brott, Sonoran Institute

• Alejandro Castellanos, Universidad de Sonora

• Cori Dolan, University of Arizona

• Peter Ffolliott, University of Arizona

• Gerald Gottfried, USDA Forest Service, Rocky Mountain Research Station

• Bill Halvorson, University of Arizona

• Philip Heilman, USDA Agricultural Research Service

• David Hodges, Cuenca los Ojos Foundation

• Larry Jones, USDA Forest Service, Coronado National Forest

• Tom Van Devender, Sky Island Alliance

• Christina Vojta (ret.), U.S. Fish and Wildlife Service

• Claire Zugmeyer, Sonoran Institute.

On behalf of the organizing committee for the Third Conference on Biodiversity and Management of the Madrean Archipelago, we invite you to explore the articles collected in this 

proceedings volume. If you find things of interest, we encourage you to share them broadly, 

and to contact the authors with questions and ideas. Most of these papers are works in progress 

and probably do not represent the final word. It is our hope that they serve, instead, as the 

beginning of many fruitful conversations and additional studies.

References

DeBano, L.F.; Ffolliott, P.F.; Ortega-Rubio, A.; Gottfried, G.J.; Hamre, R.H.; Edminster, C.B., tech. coords. 1995. 

Biodiversity and management of the Madrean Archipelago: the sky islands of southwestern United States and 

northwestern Mexico. Gen. Tech. Rep. RM-GTR-264. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range Experiment Station. 669 p.

Gottfried, G.J.; Gebow, B.S.; Eskew, L.G.; Edminster, C.B., comps. 2005. Connecting mountain islands and desert 

seas: Biodiversity and management of the Madrean Archipelago II. Proceedings RMRS-P-36. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p. 

Molina, F.; Van Devender, T., eds. 2010. Diversidad Biológica de Sonora. Universidad Nacional Autónoma de 

México. 496 p.
ix

A Special Thanks

Bill Halvorson

The organizing committee for the 

Madrean Archipelago III conference offers a special thanks to William L. Halvorson for his significant 

contributions to our knowledge of 

natural resources in the Southwest 

and especially for his dedication 

in helping produce the Madrean 

Archipelago conferences and other 

regional, national, and international 

meetings over the past two decades. 

He has long championed his belief 

that knowledge unshared is knowledge lost.

Bill came to Tucson in 1992 with the National Park Service as Leader of the Cooperative 

Park Studies Unit on the University of Arizona campus. He began organizing conferences 

originally for the National Park Service to bring scientists and managers together to share new 

knowledge of biological and cultural resources on southwestern park lands. In 1998, he was 

awarded both the Rocky Mountain Region and National Natural Resources Research Award 

of the National Park Service. He, along with research scientists in the other Department of the 

Interior agencies, was transferred to the National Biological Survey, which eventually became 

the Biological Resources Discipline within the U.S. Geological Survey. Bill soon expanded 

these biennial conferences to include other agencies and organizations to the extent that he 

was one of the co-chairs of the Madrean Archipelago II conference in 2004, which was also 

the fifth in the series of his biennial meetings on Research and Resource Management in the 

Southwestern Deserts.

Bill Halvorson was one of the organizers of the Society for Ecological Restoration, serving 

as Treasurer for 4 years and serving on the Board of Directors for 13 years. He has also served 

as program chair for a series of international Ecological Restoration Conferences in Florida, 

California, Spain, Australia, and Mexico. He is currently working on the 2013 conference, which 

will be SER’s 25th Anniversary Meeting in the birthplace of ecological restoration, Madison, 

Wisconsin.

Bill has been an important advisor to us in organizing this Madrean Archipelago III conference. We thank him for his contributions to science and management of biological and cultural 

resources in the North American Southwest (including northern Mexico), his wise counsel, 

and his friendship.

Having retired from Federal Service, Bill can be contacted at sustainablesolutions@mindspring.com.
x
xi

Contents

Acknowledgments.....................................................................................................................iii

Preface ......................................................................................................................................vii

Dedication .................................................................................................................................ix

Conference Summary

Biodiversity and Management of the Madrean Archipelago III: Closing Remarks and 

Notes From the Concluding Session.................................................................................................................1

Dale S. Turner

The Nature Conservancy, Tucson, Arizona

Alejandro Castellanos

Universidad de Sonora, Hermosillo, Sonora, Mexico

Plenary Sessions

Papers

National Wildlife Refuge Management on the United States/Mexico Border .................................................... 5

William R. Radke

U.S. Fish and Wildlife Service; San Bernardino and Leslie Canyon National Wildlife Refuges, Douglas, Arizona

Biodiversity in the Madrean Archipelago of Sonora, Mexico............................................................................ 10

Thomas R. Van Devender; Sergio Avila-Villegas; and Melanie Emerson

Sky Island Alliance, Tucson, Arizona

Dale Turner

The Nature Conservancy, Tucson, Arizona

aron D. Flesch

University of Montana, Missoula, Montana

Nicholas S. Deyo

Sky Island Alliance, Tucson, Arizona

Abstracts

Climate Change, Megadroughts, and the North American Monsoon............................................................... 17

Julia Cole

Geosciences and Atmospheric Sciences, University of Arizona, Tucson, Arizona

Floods, Droughts, and Aquatic Biodiversity in the Madrean Archipelago....................................................... 17

David A. Lytle

Department of Zoology, Oregon State University, Corvallis, Oregon

Riparian Vegetation and Disappearing Groundwater ........................................................................................ 18

Juliet Stromberg

School of Life Sciences, Arizona State University, Tempe, Arizona

Conservation Management of the Ajos Bavispe National Forest Preserve and Wildlife Refuge*

J. Mario Cirett-Galan

Ajos Bavispe Preserve, San Pedro River Basin and Sierra San Luis

Pine-Oak Woodlands in Mexico: Challenges and Opportunities in Conserving Sky Islands*

Jose A. Sarakhan-Kermez

Nacional of CONABIO

* Abstract not received

Santa Cruz River Conservation

Conservation Efforts and Possibilities for Increased Collaboration in the 

Santa Cruz River Watershed ............................................................................................................................21

Claire A. Zugmeyer and Emily M. Brott

Sonoran Institute, Tucson, Arizona

Changing Climate in the Sky Island Region

A Conceptual Model of Plant Responses to Climate with Implications for 

Monitoring Ecosystem Change........................................................................................................................27

C. David Bertelsen

Herbarium and the School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion ...........................33

Theresa M. Crimmins

USA National Phenology Network: School of Natural Resources and the Environment, University of Arizona, 

Tucson, Arizona

Michael A. Crimmins

Department of Soil, Water and Environmental Science, University of Arizona, Tucson, Arizona
xii

C. David Bertelsen

Herbarium and School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Are Madrean Ecosystems Approaching Tipping Points? Anticipating Interactions of Landscape 

Disturbance and Climate Change ....................................................................................................................40

Donald A. Falk

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona 

Preliminary Assessment of Changes in a Lizard Assemblage at an Ecotone in 

Southeastern Arizona .......................................................................................................................................48

Lawrence L. C. Jones

USDA Forest Service, Coronado National Forest, Tucson, Arizona

It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change .......................................................53

John L. Koprowski, Sandra L. Doumas, Melissa J. Merrick, Brittany Oleson, Erin E. Posthumus, Timothy G. Jessen, 

and R. Nathan Gwinn

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action .......................60

Louise W. Misztal

Sky Island Alliance, Tucson, Arizona

Gregg Garfin

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Lara Hansen

EcoAdapt, Bainbridge Island, Washington

Southwestern Cienegas

Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona.................71

Elizabeth Makings

School of Life Sciences, Arizona State University, Tempe, Arizona

Paleoenvironmental Framework for Understanding the Development, Stability, and State-Changes 

of Ciénegas in the American Deserts .............................................................................................................77

Thomas A. Minckley

Department of Geography and Program in Ecology, University of Wyoming, Laramie, Wyoming

Andrea Brunelle

Department of Geography, University of Utah, Salt Lake City, Utah

Dale Turner

The Nature Conservancy, Tucson, Arizona

Northern Jaguar Reserve

When Will Female Jaguars Cross the Border? Socio-Demographics of the Northern Jaguar.......................87

Peter Warshall

Northern Jaguar Project, Tucson, Arizona

Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessons ....................................91

Peter Warshall

Northern Jaguar Project, Tucson, Arizona

Aaron D. Flesch,

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Buying Land for Conservation Purposes in Sonora, Mexico.............................................................................97

S. Lucía Perez-Weil, Juan Carlos G. Bravo

Naturalia, Comité para la Conservación de Especies Silvestres, A.C., Northern Jaguar Project, Hermosillo, Sonora, México

Amphibian Conservation

A Comparison of the Herpetofaunas of Ranchos Los Fresnos and El Aribabi in 

Northern Sonora, Mexico................................................................................................................................103

James C. Rorabaugh

Saint David, Arizona

Jeffrey M. Servoss

U.S. Fish and Wildlife Service, Tucson, Arizona

Valerie L. Boyarski

Arizona Game and Fish Department, Phoenix, Arizona

Erin Fernandez, and Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Carlos Robles Elías

Rancho El Aribabi, Imuris, Sonora, Mexico

Kevin E. Bonine

Department of Ecology and Evolutionary Biology and School of Natural Resources and the Environment, University of 

Arizona, Tucson, Arizona
xiii

Comparison of Preliminary Herpetofaunas of the Sierras la Madera (Oposura) and Bacadéhuachi with the 

Mainland Sierra Madre Occidental in Sonora, Mexico................................................................................. 110

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Erik F. Enderson

Drylands Institute, Tucson, Arizona

Dale S. Turner

The Nature Conservancy, Tucson, Arizona

Roberto A. Villa

Tucson Herpetological Society, Tucson, Arizona

Stephen F. Hale

EcoPlan Associates, Inc., Mesa, Arizona

George M. Ferguson

University of Arizona, Tucson, Arizona

Charles Hedgcock

Sky Island Alliance, Tucson, Arizona

Rivers and Streams

Distribution of Riparian Vegetation in Relation to Streamflow in Pima County, Arizona .............................. 119

Julia E. Fonseca

	 Pima County Office of Conservation and Sustainability, Tucson, Arizona

Mike List

Pima County Information Technology, Tucson, Arizona 

Observations on the Seasonal Distribution of Native Fish in a 10-Kilometer Reach of San 

Bernardino Creek, Sonora, Mexico................................................................................................................124

C.O. Minckley

Cuenca los Ojos, Flagstaff, Arizona

Biodiversity and Conservation of the Ciénega de Saracachi area, Sonora, Mexico......................................127

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Martín A. Villa-Andrade; Martín Reyes-Juárez; and Gonzálo Luna-Salazar

Comisión de Ecología Sustentable del Estado de Sonora, Hermosillo, Sonora

Martín Padrés-Contreras, Fernando Padrés

Rancho Agua Fría, Cucurpe, Sonora

Paul S. Martin

University of Arizona, Tucson, Arizona

Biodiversity: Arthropods

Preliminary Assessment of Biogeographic Affinities of Selected Insect Taxa of the 

State of Sonora, Mexico..................................................................................................................................133

Robert W. Jones, Alejandro Obregón-Zuñiga and Sandra Guzman-Rodriguez

Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Querétaro, Qro., México

Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern 

Chihuahuan Desert .........................................................................................................................................138

Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

John S. Ascher

Department of Entomology, American Museum of Natural History, New York, New York

Introduction to the Arizona Sky Island Arthropod Project (ASAP): Systematics, Biogeography, 

Ecology, and Population Genetics of Arthropods of the Madrean Sky Islands ........................................144

Wendy Moore, Wallace M. Meyer, III, Jeffrey A. Eble, and Kimberly Franklin

Department of Entomology, University of Arizona, Tucson, Arizona

John F. Wiens and Richard C. Brusca

Arizona-Sonora Desert Museum, Tucson, Arizona

Preliminary Assessment of the Moth (Lepidoptera: Heterocera) Fauna of Rincon de Guadalupe, Sierra de 

Bacadéhuachi, Sonora, Mexico .....................................................................................................................169

John D. Palting

Department of Entomology, University of Arizona, Tucson, Arizona

Sand Dune of Ruby, Arizona, an Anthropogenically Created Biodiversity Hotspot for Wasps 

and Their Velvet Ant Parasitoids....................................................................................................................172

Justin O. Schmidt

Southwestern Biological Institute, Tucson, Arizona 

Bird List of San Bernardino Ranch in Agua Prieta, Sonora, Mexico...............................................................175

Melinda Cárdenas-García and Mónica C. Olguín-Villa

Universidad de Sonora, Hermosillo, Sonora, Mexico
xiv

Preliminary Assessment of Species Richness and Avian Community Dynamics in the Madrean 

Sky Islands, Arizona........................................................................................................................................180

Jamie S. Sanderlin, William M. Block, Joseph L. Ganey, and Jose M. Iniguez

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Biodiversity: Plants

Biodiversity Effects on Ecosystem Function Due to Land Use: The Case of Buffel Savannas in the Sky 

Islands Seas in the Central Region of Sonora..............................................................................................191

A. E. Castellanos , H. Celaya , C. Hinojo , and A. Ibarra 

DICTUS, Universidad de Sonora, Hermosillo, Sonora

J. R. Romo 

School of Natural Resources and Environment, University of Arizona, Tucson, Arizona

Biogeography and Diversity of Pines in the Madrean Archipelago.................................................................197

George M. Ferguson

University of Arizona, Tucson, Arizona

Aaron D. Flesch

Avian Science Center, Division of Biological Sciences, University of Montana, Missoula, Montana

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Ecosystems and Diversity of the Sierra Madre Occidental..............................................................................204

M. S. González-Elizondo, M. González-Elizondo, L. Ruacho González, I.L. Lopez Enriquez, F.I. Retana Rentería, 

and J.A. Tena Flores

CIIDIR I.P.N. Unidad Durango, Mexico

Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: 

A Diversity to be Explored..............................................................................................................................212

César Hinojo Hinojo, Alejandro E. Castellanos, and Jose M. Llano Sotelo

	 Laboratorio de Ecofisiología Vegetal, Departamento de Investigaciones Científicas y Tecnológicas de la

Universidad de Sonora, Hermosillo, Sonora, Mexico

Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone..........................................218

Robert G. Laport and Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

Chihuahuan Desert Flora of La Calera, Municipio de Agua Prieta, Sonora, Mexico......................................225

Ana Lilia Reina-Guerrero; and Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico ......................................229

Thomas R. Van Devender, Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona

J. Jesús Sánchez-Escalante

	 Departamento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Hermosillo, Sonora, Mexico

Preliminary Flora of the Sierra Bacadéhuachi, Sonora, Mexico ......................................................................236

Thomas R. Van Devender and Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona

George M. Ferguson

University of Arizona, Tucson, Arizona

George Yatskievych

Missouri Botanical Garden, St. Louis, Missouri

Beatriz E. Loyola-Reina

JRM Consultores, Hermosillo, Sonora, Mexico

Gertrudis Yanes-Arvayo, and Maria de la Paz Montañez-Armenta

Universidad de la Sierra, Moctezuma, Sonora, Mexico

John L. Anderson

Bureau of Land Management, Phoenix, Arizona

Stephen F. Hale

EcoPlan Associates, Inc., Mesa, Arizona

Sky Jacobs

Wild Sonora, Tucson, Arizona

Comparison of the Tropical Floras of the Sierra la Madera and the Sierra Madre Occidental, 

Sonora, Mexico ...............................................................................................................................................240

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Gertrudis Yanes-Arvayo

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona
xv

Melissa Valenzuela-Yánez, Maria de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Biodiversity: Reptiles

Biogeographic Perspective of Speciation Among Desert Tortoises in the Genus Gopherus: 

A Preliminary Evaluation ................................................................................................................................243

Taylor Edwards

University of Arizona Genetics Core, University of Arizona, Tucson, Arizona

Mercy Vaughn

Paso Robles, California

Cristina Meléndez Torres

Comisión de Ecología y Desarrollo Sustentable del Estado de Sonora, Sonora, Mexico

Alice E. Karl

Alice E. Karl and Associates, Davis, California

Philip C. Rosen

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Kristin H. Berry

U.S. Geological Survey, Western Ecological Research Center, Riverside, California

Robert W. Murph

Royal Ontario Museum, Toronto, Canada

Biodiversity: Mammals

Richness of Mammals on the San Bernardino Ranch in the Municipality of Agua Prieta, 

Sonora, Mexico................................................................................................................................................248

Mario Erandi Bonillas-Monge and Carlos Manuel Valdez-Coronel

	 Departamento de Investigaciones Científicas y Tecnológicas de la Universidad de Sonora, Sonora, Mexico

Preliminary List of Flying Mammals in the Ajos-Bavispe National Forest Reserve and 

Wildlife Refuge, Sonora ..................................................................................................................................252

Rosa Elena Jiménez

CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

Christ D. Weise 

U.S. Fish and Wildlife Service, Kofa National Wildlife Refuge, Yuma, Arizona

Mario Cirett-Galán, Guadalupe Flores, and Manuel Munguia

CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

E. Isaías Ochoa

Naturalia, A.C., Hermosillo, Sonora

Decade of Wildlife Tracking in the Sky Islands .................................................................................................256

Jessica A. Lamberton-Moreno

Wildlife Linkages Program, Sky Island Alliance, Tucson, Arizona

Sergio Avila-Villegas

Northern Mexico Program, Sky Island Alliance, Tucson, Arizona

Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States.....................263

N. E. Lara-Díaz and C. A. López-González

Universidad Autónoma de Querétaro, Qro., Mexico

H. Coronel-Arellano and A. González-Bernal

Naturalia, A. C. Sonora, Mexico

Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps.........................................269

Don E. Swann and Nic Perkins

Saguaro National Park, Tucson, Arizona

Biodiversity: Aquatic

Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure in the 

Madrean Sky Islands.......................................................................................................................................277

Michael T. Bogan

Oregon State University, Corvallis, Oregon

Oscar Gutierrez-Ruacho and J. Andrés Alvarado-Castro,

CESUES, Hermosillo, Sonora, Mexico

David A. Lytle

Oregon State University, Corvallis, Oregon

Gila Topminnow Interactions With Western Mosquitofish: An Update ...........................................................283

Douglas K. Duncan

	 Arizona Ecological Services Office, U.S. Fish and Wildlife Service, Tucson, Arizona
xvi

Native Aquatic Vertebrates: Conservation and Management in the Río Sonoyta Basin, 

Sonora, Mexico................................................................................................................................................288

C. Minckley

Cuenca los Ojos, Flagstaff, Arizona

Izar Izaguirre Pompa

Reserva de la Biosfera del Pinacate y Gran Desierto de Altar, Puerto Peñasco, Sonora, Mexico

Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Ross Timmons

Arizona Game and Fish Department, Phoenix, Arizona

Dennis Caldwell

Caldwell Design, Tucson, Arizona

Jaime López Méndez

Colegio de Bachilleres del Estado de Sonora, Sonoyta, Sonora, Mexico

Phil Rosen

University of Arizona, Tucson, Arizona

Biodiversity Workshop: Flora and Fauna

Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual 

Flora and Fauna...............................................................................................................................................292

Nicholas S. Deyo and Thomas R. Van Devender

Sky Island Alliance, Tucson Arizona

Alex Smith

School of Natural Resources and the Environment, University of Arizona, Tucson Arizona

Edward Gilbert

SEINet Flora Database, Tucson, Arizona

Restoration

Agricultural Field Reclamation Utilizing Native Grass Crop Production ........................................................303

J. Curé

Polytechnic Campus, Arizona State University, Mesa, Arizona

Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: 

Arizona Reintroduction ..................................................................................................................................310

Sarah L. Hale, and John L. Koprowski

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Holly Hicks

Arizona Game and Fish Department, Phoenix, Arizona

Trajectory and Rate of Desert Vegetation Response Following Cattle Removal ...........................................316

Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

Fire

Soil Erosion and Deposition Before and After Fire in Oak Savannas .............................................................325

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona

Hui Chen, Aaron T. Kauffman, and Cody L. Stropki

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Daniel G. Neary

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires ..................................329

Carly Gibson

USDA Forest Service, Stanislaus National Forest, Sonora, California

Ann Youberg

Arizona Geological Society, Tucson, Arizona

Marc Stamer

USDA Forest Service, San Bernardino National Forest, Fawnskin, California

Ecology and Management of Oak Woodlands and Savannas in the Southwestern Borderlands Region .....337

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona 

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona
xvii

Effects of Prescribed Fires and a Wildfire on Biological Resources of Oak Savannas in the 

Peloncillo Mountains, New Mexico................................................................................................................341

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Daniel G. Neary

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Impacts of Wildfire on Wildlife in Arizona: A Synthesis ...................................................................................345

Shari L. Ketcham and John L. Koprowski

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Burned Saguaro: Will They Live or Die? ...........................................................................................................351

Marcia G. Narog and Bonni M. Corcoran

	 USDA, Forest Service, Pacific Southwest Research Station, Riverside, California

Ruth C. Wilson

Department of Biology, California State University, San Bernardino, California

Post-Wildfire Erosion in the Chiricahua Mountains .........................................................................................357

Ann Youberg

Arizona Geological Survey, Tucson, Arizona

Daniel G. Neary, Karen A. Koestner, and Peter E. Koestner

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Tools for Management

Mapping and Assessing the Environmental Impacts of Border Tactical Infrastructure in the 

Sky Island Region ...........................................................................................................................................365

Caroline Patrick-Birdwell

School of Geography and Development, University of Arizona, Tucson, Arizona

Sergio Avila-Villegas, Jenny Neeley, and Louise Misztal

Sky Island Alliance, Tucson, Arizona

Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management ....................370

Jess Clark

USDA Forest Service, Remote Sensing Applications Center, Salt Lake City, Utah

Marc Stamer

USDA Forest Service, San Bernardino National Forest, Fawnskin, California

Kevin Cooper

USDA Forest Service, Los Padres National Forest, Santa Maria, California

Carolyn Napper

USDA Forest Service, Shasta-Trinity National Forest, Mt. Shasta, California

Terri Hogue and Alicia Kinoshita

University of California Los Angeles, Los Angeles, California

Gila River Basin Native Fishes Conservation Program....................................................................................376

Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Robert W. Clarkson

Bureau of Reclamation, Glendale, Arizona

Mapping Ecological Systems in Southeastern Arizona (NOTE: This paper was originally part of this session

 and belatedly moved to this locatioon, so page numbers are not sequential within these proceedings)......... A-1

Jim Malusa and Donald Falk

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Larry Laing

Environmental Consultant, Gold Canyon, Arizona

Brooke Gebow

The Nature Conservancy, Southeastern Arizona Preserves, Hereford, Arizona

Potential for Extending Major Land Resource Areas into Northern Mexico...................................................381

Roy S. Mann

USDA-NRCS Retired, Range Management

Philip Heilman and Jeffry Stone

USDA Agricultural Research Service, Southwest Watershed Research Center, Tucson, Arizona

Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models.......................389

Melissa J. Merrick and John L. Koprowski

Wildlife and Fisheries Science, School of Natural Resources and the Environment, University of Arizona, Tucson

Craig Wilcox

USDA Forest Service, Coronado National Forest, Safford, Arizona
xviii

Grasslands

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting and Restoring the 

Sky Island Region’s Most Intact Grassland Valley Landscapes.................................................................399

Gitanjali S. Bodner and Peter Warren

The Nature Conservancy in Arizona

David Gori, Karla Sartor, and Steven Bassett 

The Nature Conservancy in New Mexico

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition 

Using State-and-Transition Models in Upper Cienega Creek Watershed ..................................................410

Ron Tiller, Melissa Hughes and Gita Bodner

The Nature Conservancy in Arizona

Human Impacts

Long Distance Commutes by Lesser Long-nosed Bats (Leptonycteris yerbabuenae) to Visit 

Residential Hummingbird Feeders ................................................................................................................427

Debbie C. Buecher

Buecher Biological Consulting, Tucson, Arizona

Ronnie Sidner

Ecological Consulting, Tucson, Arizona

Effects of Roads on Wildlife in Arizona: How Far Have We Traveled?............................................................434

Hsiang Ling Chen and John L. Koprowski 

Wildlife Conservation and Management, School of Natural Resources and the Environment, University of Arizona, 

Biological Sciences East, Tucson, Arizona 

Tools for Research

Wildlife Survey and Monitoring in the Sky Island Region with an Emphasis on 

Neotropical Felids ...........................................................................................................................................441

Sergio Avila-Villegas and Jessica Lamberton-Moreno

Sky Island Alliance, Tucson, Arizona 

Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey in Saguaro 

National Park ...................................................................................................................................................448

Mary Beth Benton

Saguaro National Park, Tucson, Arizona

Siria A. Cerda-Navarro, Katie R. Keck, and Brittany N. McKnight

University of Arizona, Tucson, Arizona

Wildlife Surveys and Monitoring With the Use of Remote Camera Traps in the Greater 

Oak Flat Watershed Near Superior, Arizona .................................................................................................454

Roger Featherstone

Arizona Mining Reform Coalition

Sky Jacobs

Wild Sonora, Tucson, Arizona

Sergio Avila-Villegas

Sky Island Alliance, Tucson, Arizona

Sandra Doumas

University of Arizona, Tucson, Arizona

Poster Papers

Medium and Large Mammals in the Sierra La Madera, Sonora, Mexico .........................................................463

Erick Oswaldo Bermúdez-Enríquez

Universidad de la Sierra, Moctezuma, Sonora

Rosa Elena Jiménez-Maldonado

Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

Gertrudis Yanes-Arvayo, María de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya

 Universidad de la Sierra, Moctezuma, Sonora

Herpetofauna at the Appleton-Whittell Research Ranch..................................................................................466

Roger C. Cogan

Appleton-Whittell Research Ranch of the National Audubon Society, Elgin, Arizona

Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) from a Cave in the 

Huachuca Mountains, Arizona .......................................................................................................................468

Nicholas J. Czaplewski

Oklahoma Museum of Natural History, University of Oklahoma, Norman, Oklahoma

Steve Willsey

Hereford, Arizona
xix

FireScape: A Program for Whole-Mountain Fire Management in the Sky Island Region..............................472

Brooke Gebow

The Nature Conservancy, Southeastern Arizona Preserves, Hereford Arizona; USDA Forest Service, Coronado National 

Forest, Tucson, Arizona

Christopher Stetson

USDA Forest Service, Coronado National Forest, Tucson, Arizona

Donald A. Falk and Corrine Dolan

School of Natural Resources and the Environment, Biological Sciences East, University of Arizona, Tucson, Arizona

Floristic Analysis of Heterogeneous Landscape Patches in a Biological Corridor in the 

El Rodeo-Básora Area near Moctezuma, Sonora, Mexico ..........................................................................474

Ana Lilia Hernández-Rodríguez, María de la Paz Montañez-Armenta, Gertrudis Yanes-Arvayo, and Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora

Association Between Nurse Plants and Saguaros (Carnegiea gigantea) in the Western Sonora..................477

C. Hinojo-Hinojo, C. Trujillo-López, O. Calva-Pérez, O. Galaz-García, A. E. Castellanos-Villegas

	 Departamento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Hermosillo, Mexico

Vegetation Monitoring on Semi-Arid Grasslands Ungrazed by Domestic Livestock ....................................479

Linda Kennedy

National Audubon Society, Appleton-Whittell Research Ranch, Elgin, Arizona

Dan Robinett

Robinett Rangeland Resources LLC, Elgin, Arizona

Mountain Pine Beetle in Southwestern White Pine in the Pinaleño Mountains .............................................482

Ann M. Lynch

USDA Forest Service, Rocky Mountain Research Station, Tucson, Arizona

Christopher D. O’Connor

School of Renewable Natural Resources, University of Arizona, Tucson, Arizona

Population Status of Prairie Dogs (Cynomys ludovicianus) in the 

San Pedro River Basin, Sonora ....................................................................................................................487

Efrén Moreno-Arzate and Carlos A. López González

Universidad Autónoma de Querétaro, Querétaro, Mexico

Gerardo Carreón Arroyo

Naturalia A. C. Hermosillo, Sonora, Mexicoa

Babocomari River Riparian Protection Project .................................................................................................490

Dan Robinett

Robinett Rangeland Resources LLC, Elgin, Arizona

Linda Kennedy

Appleton-Whittell Research Ranch of the National Audubon Society, Elgin, Arizona 

Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for 

Avian Communities.........................................................................................................................................493

Vashti (Tice) Supplee

Arizona Important Bird Area Program, Audubon Arizona, Phoenix, Arizona

Jennie MacFarland 

Arizona Important Bird Area Program, Tucson Audubon Society, Tucson, Arizona

Preliminary Flora of Ojo de Agua Tonibabi, Sierra La Madera, Sonora, Mexico ...........................................500

Melissa Valenzuela-Yánez, Gertrudis Yanes-Arvayo, Maria de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona 

Examining Wildlife Responses to Phenology and Wildfire Using a Landscape-Scale 

Camera Trap Network .....................................................................................................................................503

Miguel L. Villarreal, Leila Gass, Laura Norman, Joel B. Sankey, Cynthia S.A Wallace, and Dennis McMacken

USGS Western Geographic Science Center, Tucson, Arizona

Jack L. Childs

Borderlands Jaguar Detection Project, Amado, Arizona 

Roy Petrakis

University of Arizona, Tucson, Arizona

Mapping Landscape Phenology Preference of Yellow-Billed Cuckoo With AVHRR Data .............................506

Cynthia S. A. Wallace and Miguel Villarreal

Western Geographic Science Center, U.S. Geological Survey, Tucson, Arizona

Charles van Riper, III

Southwest Biological Science Center, Sonoran Desert Research Station, U.S. Geological Survey, Tucson, Arizona.

Abstracts

Madrean Abstracts ............................................................................................................................................... 511

USDA Forest Service Proceedings RMRS-P-67. 2013 1

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Biodiversity and Management of the Madrean Archipelago III: 

Closing Remarks and Notes from the Concluding Session

Dale S. Turner

The Nature Conservancy, Tucson, Arizona

Alejandro Castellanos

Universidad de Sonora, Hermosillo, Sonora, Mexico

	 During the first week of May 2012, the Third Conference on

Biodiversity and Management of the Madrean Archipelago brought

together more than 300 people with an interest in this region. It

included scientists, land managers, activists, and land owners from

both sides of the international border. After three and a half days of

presentations, the participants gathered for a moderated discussion of

major issuesfor biodiversity and management. They were also asked

to suggest ways in which people could build stronger collaborations.

The following is a summaryofthosediscussions,withminor additions

to add context.

International Border

	 We work at an ecotone of ecoregions, policies, and countries

where many things change when you cross the international border.

The borderlands form a mixing zone, different from both countries.

The past decade has brought new challenges to collaboration and

management for biodiversity across the border. The barbed-wire

fence became a steel wall in many places, blocking the movement of

wildlife (Córdova and de la Parra 2007; Flesch and others 2010). New

social barriers were created by anti-immigrant legislation inArizona,

migrant and drug-related problems in Sonora and Chihuahua, and

tougher regulations for legal trans-boundary movement.

	 Despite those influences, useful collaborations continue and new

opportunities exist. Several good examples were presented at the

conference, including riparian restoration on both sides of the border

in San Bernardino National Wildlife Refuge and Rancho San Bernardino, and a bi-national conservation planning workshop for the

SantaCruzRiver.Inadditiontostrongrepresentationatthis conference

from Mexico and the United States, about 17% of the presentations

included authors from both countries. Participants mentioned a large

and growing population of biology students in Mexico, creating new

opportunities for collaboration. It was also mentioned that private

land owners in Mexico are much more open than Americans to having people work on their land, and are more likely to make small

management changes if they know that others care.

Bi-National Coordination and Sharing 

Information

	 The MadreanArchipelago conference series has been a useful way

to share knowledge between those who attend, and the proceedings

from these conferences have been good reference works. However,

land managers, decision makers, and the general public often do not

read such technical reports and may not be aware of their existence.

Conference attendees can make a big difference by sharing the results

with others, both within and beyond their organizations.The information could be even more useful if it were summarized for particular

audiences,with themajor newinformation putinto a regional context.

	 There is also great potential for putting new information into

practice on the ground, by finding ways to demonstrate the results

to land owners and land managers. It is not enough to just tell them

how to do something; showing them a technical report is not helpful.

We need to demonstrate how conservation can work for the land and

for people.

	 For researchers and technical staff, there is a need for consistent

data sets across the region. This includes the need for basic GIS

data such as vegetation, elevation, and streams. In some cases, the

data may be available but not widely known, such as a recent North

American mapping of biotic communities(Brown et al. 2007; www.

peter.unmack.net/biotic). Several presentations at this conference

featured the new regional database of plant and animal records compiled as part of the Madrean Archipelago Biodiversity Assessment

(Van Devender and others, The Madrean Archipelago biodiversity 

assessment, this proceedings), valuable information on biological

diversity for Sonora (Molina-Freaner and Van Devender 2010), and

the ambitious national data sets prepared for Mexico by CONABIO

(www.conabio.gob.mx).

	 TheMadreanArchipelago framework provides an excellent opportunity for developing more innovative ways to increase coordination

amongorganizations,researchers andstudents.Increasinginterest and

student participation in Mexico can help build stronger ties between

research organizations on both sides of the border. Many participants

recognized the need for more cross-border meetings, particularly attendingmeetingsheldinMexico.While thatmightreduceparticipation

from the United States, it would increase Mexican-U.S. collaborative

participation. Some suggested that the next Madrean Archipelago

conference should be held in Sonora. Others noted the growth of

a student-run biology conference held annually at Universidad de

Sonora in Hermosillo, and invited U.S. researchers to participate.

There was also the first announcement of a conference on biology

and management of the Sierra Madre, planned for Durango in 2014.

Participants were also hopeful that improvements in video conferencing could make cross-border meetings easier. Beyond that, small
2 USDA Forest Service Proceedings RMRS-P-67. 2013

Turner and Castellanos Biodiversity and Management of the Madrean Archipelago III: Closing Remarks . . .

workshops ortrainings heldmore frequently could build relationships

and spread knowledge without a lot of work. And the knowledge

transfer needs to go both directions, north and south, because there

are leaders in research and management on both sides of the border.

We consider that an extra effort should be developed in the coming

years to provide ways to build stronger links between institutions, in

particular among students and colleagues.

	 Many participants felt that bad news about the border region was

exaggerated. They mentioned the challenge getting a positive media

story about good collaborative work, and suggested a need to work

harder at sharing positive news, which might include sending stories

to the media, requesting fair coverage, or independently distributing

news through the Internet.

Volunteers

	 Increasing the use of volunteers in research and monitoring is a

trend that needs to continue in this era of tight budgets. Participants

mentioned that an aging U.S. population means a growing pool of

potential volunteers with extensive knowledge and skills, soon to

include many attendees at this conference. Those volunteers can

sometimes make a greater impact if they don’t just rely on managers

toidentifyvolunteerneeds,butinsteadhelpmanagersunderstandwhat

the individual volunteers can provide. Managers can also increase

the number and variety of their volunteers, both young and old, by

making it easier to get help from someone who has time available.

In particular, a web site could be developed to publicize volunteer

research opportunities.

Future Conferences

	 The lastMadreanArchipelago conference had a substantial cultural

resource component, while this conference included only two papers.

Cultural resources provide a tie to the land and can help in the conservation process. Native nations need to be included as well; they also

manage lands and have different perspectives to offer. Preparation

for future conferences should include reaching out to the cultural

resource communities and bringing those voices into the mix.

Conclusions

	 The challenges of merging science and management in a rapidly

changing world cannot be solved in a 4-day conference, but require

strong coordination through time between disciplines and across

borders. The presentations at this event included results from collaborative projects that began at previous Madrean Archipelago

conferences. We look forward to the next conference, and to learning

the new discoveries that began with conversations in the hallways at

this event.

References

Brown, D.E.; Unmack, P.J.; Brennan, T.C. 2007. Digitized map of biotic

communities for plotting and comparing distributions of North American

animals. The Southwestern Naturalist. 52: 610-616.

Córdova,A.; de la Parra, C. (eds.). 2007.Abarrier to ourshared environment:

The border fence between the United States and Mexico. El Colegio de la

Frontera Norte, Tijuana, B.C. México. 206 p.

Flesch, A.D.; Epps, C.W.; Cain, J.W.; Clark, M.; Krausman, P.R.; Morgart

J.R. 2010. Potential effects of the United States-Mexico border fence on

wildlife. Conservation Biology 24: 171-181.

Molina-Freaner, F.; Van Devender, T. 2010. Diversidad Biológica de Sonora.

CONABIO - UNAM, México. 496 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013

3

Plenary Sessions
4 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 5

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

National Wildlife Refuge Management on the United 

States/Mexico Border

William R. Radke

U.S. Fish and Wildlife Service; San Bernardino and Leslie Canyon National Wildlife Refuges, Douglas, Arizona

Abstract—Many conservation strategies have been developed by the U.S. Fish and Wildlife Service in cooperation with others to protect habitat and enhance the recovery of fish and wildlife populations in the 

San Bernardino Valley, which straddles Arizona, United States, and Sonora, Mexico. Habitats along this 

international border have been impacted by illegal activities, frustrating recovery of rare species. In addition, 

potential threats to national security have prompted the United States to aggressively control the country’s 

boundaries, thus creating additional challenges for land managers mandated with protecting the nation’s 

landscapes, natural resources, and associated values. Such challenges are not insurmountable and, with 

focused coordination, resource management and border security can be achieved and can often compliment 

one another. With or without the influence of changes along the international border, an effective species 

recovery strategy must include a coordinated approach that involves assessing the biological requirements 

of selected species through combinations of inventory, monitoring, and research activities; managing and 

protecting existing and historic habitats and populations; assessing potential reintroductions of key species 

into appropriate habitats where feasible; managing exotic plants and animals that threaten the recovery of 

desired conditions; and providing outreach and education relative to the species, their habitats, and the 

ecosystems upon which all fish, wildlife, and humans depend. 

Introduction and Management Context

	 The U.S. Fish and Wildlife Service (Service) manages the 2,369-

acre San Bernardino National Wildlife Refuge (SBNWR) and the

2,765-acreLeslieCanyonNationalWildlifeRefuge (LCNWR)located

in southeast Arizona’s Cochise County adjacent to the international

border with Mexico. Both refuges were established under the authority of the Endangered Species Act of 1973 and the Fish and Wildlife

Act of 1956 in order to “...conserve fish or wildlife which are listed

as endangered species or threatened species...or plants.” The refuge

landscapes support a documented high diversity of species including

at least 493 plants, 332 birds, 67 mammals, 43 reptiles, 13 amphibians, 8 fish, 77 dragonflies and damselflies, 130 butterflies, and over

450 bees.

	 The primary role of SBNWR and LCNWR is to sustain and recover the native fish of the Río Yaqui Basin (USFWS 1994), which

is a large watershed that drains portions of southeastern Arizona and

southwestern New Mexico in the United States, and eastern Sonora

and western Chihuahua in Mexico. This geographic region is well

known for containing enormous biological wealth on both sides of

the border (Brown and Kodric-Brown 1996; Sayre and Knight 2009;

Van Schoik and others 2006), and cooperative efforts between the

Service andenvironmentallysensitive landownersintheUnitedStates

and in Mexico are implementing conservation strategies to protect

and restore land and water sources, promote applicable monitoring

and research, and reintroduce and maintain self‐sustaining fish and

wildlife populations (Austin in press; USFWS 1995). Properties are

being managed in ways that acknowledge that they are all simply

parts of the bigger landscape necessary to perpetuate a healthy mix

of plants, fish, and wildlife within the region. Management and restoration efforts are viewed in the context of the entire landscape with

a view beyond individual ownership boundaries or the international

border. Such a view helps maintain habitat integrity across boundaries, a critical need for wildlife migration and dispersal.

	 Environmental restoration by government agencies and private

landownersis ongoing in thisregion, and specific goals and objectives

ofsuch efforts on both sides ofthe borderin theSanBernardinoValley

are focused primarily on maintenance of open space and sustainable

restoration of ecologicalrelationshipsin such a way that the residents

of the region may continue a productive rural lifestyle. Wetland

restoration throughout the watershed is benefitting populations of all

wetland dependent species by increasing the availability of seasonal

and permanent water, by increasing the overall quality and quantity

of water, and by lessening the potential for erosive scouring of wetlands during extreme flood events. Watershed function of uplands is

also being enhanced, positively impacting the entire landscape along

with the fish, wildlife, and human populations that they support. An

example of these efforts is the ongoing grassland restoration and

stream restoration being implemented on Rancho San Bernardino in

Sonora. This project is allowing native fish like Mexican stonerollers

(Campostoma ornatum) to re-colonize upstream refuge wetlands

without the need for active reintroductions.
6 USDA Forest Service Proceedings RMRS-P-67. 2013

Radke National Wildlife Refuge Management on the United States/Mexico Border

Balancing Border Security And 

Conservation

	 The San Bernardino Valley is attractive to people for reasons other

than its natural biodiversity. SBNWR, LCNWR, and adjacent lands

are isolated wildland areas adjacent to the U.S./Mexico border that

have become arteries for human trafficking and narcotics smuggling

organizations. Such activities are sometimes a hazard to landowners,

resource managers, and area visitors because immigrants and smugglers have become increasingly violent and determined to continue

theirillegal activities.The problemis not just themovement of people

and drugs, but includes related illegal activities (Billington and others 2010) such as vandalism, arson, accidental wildfire, property

damage, burglary, theft, assault, and even murder conducted by both

northbound and southbound people.In response to escalating security

concerns, the U.S. Border Patrol (USBP) is aggressively enforcing

laws to deter and prevent illegal activities and apprehend smugglers.

This effort has promoted nationalsecurity and has also benefitted the

environment by helping reverse the adverse environmental effects of

illegal border activities.

	 Unfortunately, enforcement of illegal activities along the border

may also result in incidental property damage and adverse impacts

to environmental and cultural resources (Cohn 2007; Christen and

Matlack 2009;Duncan and others 2010;Forman andAlexander 1998;

Sayre and Knight 2009; Segee and Neely 2006; Shepard and others

2008; Trombulak and Frissell 2000). Aggressive border security

measures may affect endangered species and their habitats when it

becomes necessary to clear vegetation,whichmay in turn alter natural

waterflow,reducewater absorptionandinfiltration, andimpact aquifer

recharge capacity. Operation of vehicles off established roadways

or across wetlands and flowing streams may discharge petroleum

products or other pollutants, increase siltation within perennial and

seasonal streams or washes, and accelerate erosion and mobilization

of fragile hydric soil types characteristic of the region. Each of these

impacts may also negatively influence recovery of endangered plant

species by increasing the potential for soil particles to become airborne during dry periods, potentially affecting air quality, vegetation

transpiration, and pollination. Terrestrial endangered species may be

impacted through crushing or other directmortality (Glista and others

2009; Lalo 1987; Romin and Bissonette 1996). Aquatic species may

be affected by increasing siltation of streams and other wetlands following major precipitation events. These events may result in higher

than normal levels of erosion and siltation, which potentially affects

oxygen availability, gill function, and reproduction. Illegal activities

and their undesirable impacts on the environment are not new; for at

least the past 100 years the San Bernardino Valley has been used as

a travel route for smugglers and illegal immigrants. However, since

about 2001, numbers of illegal immigrants, smugglers, and USBP

agents have increased dramatically. Therefore, illegal border activities and the enforcement to control those activities remain threats to

resource management and species recovery efforts on SBNWR and

LCNWR.

	 An effective balance can be achieved between the U.S. Department

of Homeland Security (DHS) mission of protecting U.S. borders and

the U.S. Department of Interior (DOI) mission of protecting natural

and cultural resources. SBNWR staff have been active partners with

USBP and Immigration and Customs Enforcement (ICE) in planning

and implementing national security ever since the refuge was established. More recently, increased cooperation between the Service,

USBP, and ICE personnel has additionally enhanced safety, national

security, and natural resource protection. Through such cooperative efforts, many of the damaging impacts of illegal activities and

required enforcement have been avoided, minimized, or mitigated.

This cooperative effort is exemplified by the work of Service law

enforcement officerswho function as environmentally sensitive force

multipliersserving nationalsecurity and landmanagement objectives

in a cost effective manner. At SBNWR and LCNWR, the Service is

assistingwith nationalsecurity efforts and has an effective strategy for

monitoring, preventing, and controlling illegal activities by securing

refugeboundaries, controllingaccess, andhaltingillegalimmigration.

Service lawenforcement officers play an importantrole inmonitoring

human access on Refuge lands by using a system of remote seismic

sensors and cameras, by patrolling for illegal activities and looking

forfootprints, byregularly communicatingwithUSBPagentsthrough

an improved radio system, and through joint patrol work.

	 It has become routine for Fish and Wildlife Officers to work with

USBP to conduct enforcement activities on and adjacent to Refuge

lands and in areas having sensitive naturalresources.The Service and

USBP have been demonstrably proactive in ensuring that national

security and environmental protection are not mutually exclusive

objectives, and such cooperation has been very effective, resulting in

a decrease ofillegal activities occurring on theRefuges. For example,

“unofficial” USBP statistics show an average apprehension rate of

about two individuals per month on SBNWR for the period October

2011–April 2012; down significantly from 2010, when “unofficial”

USBP statistics depicted an average of 142 individuals apprehended

each month on SBNWR (USBP, unpublished data).

	 SBNWRand LCNWRare enclosed by fences, allowing controlled

public access that is limited to daytime use only. Most of the south

SBNWR boundary is protected by a wildlife permeable vehicle barrier to protect sensitive wetlands, and has been completely effective

in preventing vehicle incursions onto the refuge from Mexico since

its construction in 2005. Trail cameras and other tactics are used to

document andmonitorwildlife use ofriparian corridors andwetlands,

and also identify and document illegal human activities occurring

on the refuges. Habitat restoration is being accomplished to benefit

wetland, upland, and riparian-dependent species. Refuge pond construction and stream restoration make human travel through mud and

water difficult, so wetlands tend to be avoided by those engaged in

illegal activities.Grasslandrestorationisbeingaccomplishedthrough

prescribed burning and removal ofinvasivemesquite trees, providing

benefits to resident and migratory wildlife and also removing cover

onceusedbysmugglers,increasingvisibilityacrossthe landscape, and

thereforedeterringillegal activities.Existingadministrative roadways

are maintained to increase refuge management efficiency and also

to promote effective law enforcement, detection, monitoring, and

deterrence of illegal activities. These roadways are being upgraded

with base coarse material to provide all weather use and to minimize

soil particle movement into the air or into adjacent wetlands. Many

roads are remotely monitored by seismically sensitive equipment to

help track human activity timing, and by video surveillance cameras

to help identify refuge users. Refuge staff and heavy equipment

routinely maintain portions of the border road within the Roosevelt

Reservation adjacent to SBNWR to ensure passage by USBP agents

through areas prone to frequent flooding and sediment deposition.

	 ManyUSBPactivitieshave further enhancedRefugeprotectionand

endangered species recovery. For example, USBP operates Mobile

Surveillance System equipment in the San Bernardino Valley to help

interdict illegal activities, and this equipment is located in off-refuge

areasthat maximize the deterrence of illegal activitiesthroughout the

Valley without negatively impacting refuge lands.Additionally, DHS

contractors have surfaced many border roads used by USBP with

material that allows all weather vehicle travel and limits dust and

sediment transfer into the environment. USBP has allowed grassesto
USDA Forest Service Proceedings RMRS-P-67. 2013 7

National Wildlife Refuge Management on the United States/Mexico Border Radke

grow within the 60-foot Roosevelt Reservation rather than attempting to maintain this easement as a corridor that is completely free

of vegetation. The original international border fence was not very

wildlife permeable, consisting of an eight-strand barbed wire barrier

reaching up to 5-feet tall. This barrier altered cross-border movement

ofwildlife,thoughsome specieswere capableofmovingoverorunder

the barbed wire fence, or of utilizing existing breaks in the fence that

allowed easier passage. Construction of vehicle barriersimmediately

adjacent to the existing barbed wire border fence further reduced

wildlife movement. Mule deer (Odocoileus hemionus) that often

seemed willing to jump the barbed wire fence would not attempt to

jump the enhanced obstacle of a barbed wire fence combined with a

vehicle barrier. The removal of the barbed wire fence by USBP, other

agency personnel, landowners, and volunteers has greatly benefitted

cross-border wildlife movement (Sayre and Knight 2009). 

	 Several environmental concerns were favorably addressed by

DHS and their contractors by installing vehicle barriers designed so

that wildlife movement would not be obstructed; investigating and

protecting culturalresourcesfrompotential damage; conducting construction during winter to avoid impactsto nesting birds and to avoid

periods of heavy, erosive rainfall; and reseeding some disturbed areas

to prevent sheet erosion and promote the growth of native grasses. A

substantial bridge was constructed by DHS contractors across Black

Drawtoprovidevehicular access alongtheborderwithintheRoosevelt

Reservation while also minimizing negative impacts to the perennial

streamand associated species.These examplesillustrate that DHS officials are committed toworkingwithSBNWRpersonneltominimize

disturbance to the refuge, to protect the environment, and to protect

endangered species. Not all ofthe foreseeable environmental impacts

related to the congressional mandate of maintaining nationalsecurity

could be minimized or eliminated,so DHS officials collaborated with

the Service to identify solutionsfor potential environmental problems

created by the infrastructure, and DHS committed to funding up to a

total of $50 million through January 2018 to support various environmental projects to be split among California, Arizona, New Mexico,

and Texas. These projects demonstrate some positive approaches

toward reconciling national security and environmental objectives

along the international border.

DHS-Funded Environmental Projects 

	 Work on many DHS-funded environmental projects has been initiated on and adjacent to SBNWR (USFWS 2011). A concrete fish

barrier in Black Draw near the border is being designed and will be

constructed to allow downstream movement of native fish but will

prevent upstream migration of non-native fish species. This is the

best available option for achieving segregation of native fish from

predatory exotic fish occurring downstream in Mexico. A series of

shallow wetland units have been constructed, and will help maximize

biodiversity and optimize the production of self-sustaining populations of native fish and wildlife. The construction and placement of

multiple rock-filled wire cage erosion control gabion structures is

being accomplished in key locations throughout the watershed to

catch water-borne sediments being transported during precipitation

events, improve the landscape’s ability to halt and slow the scouring

erosive impacts of seasonal flood flows, more effectively catch and

hold precipitation runoff to provide water for fish and wildlife, help

restore groundwater recharge in the shallow aquifer, allow increased

soil stabilization, and ultimately help enhance vegetation corridors

for wildlife movement. Construction debris deposited by contractors

in several San Bernardino Valley staging areas during vehicle barrier

construction has been removed and these areas have been seeded with

native grasses to help control the density and spread of non-native

vegetation that could otherwise become invasive and dominate the

landscape. Two separate irrigation wells on and adjacent to SBNWR

were drilled to ensure adequate water quantity and quality capable of

enhancing populations of federally listed species: Yaqui chub (Gila 

purpurea), Yaqui topminnow (Poeciliopsis sonoriensis), Yaqui catfish (Ictalurus pricei), and San Bernardino springsnails (Pyrgulopsis 

bernardina) located in the area on both public and private lands. The

Service andDHSare alsoworking collaborativelywith theUniversity

ofArizona to monitor Río Yaqui fish species and their habitats on and

adjacent to SBNWR to assesstheir population status and evaluate the

potential affectsofsedimentmobilizationintowetlands.Eachofthese

environmental projects, supported financially by DHS, is important

to recovery of listed species in the United States and Mexico.

Rare Species Recovery Strategy

Inventory, Monitoring, and Research

	 With or without the influence of changing human activity or land

use along the international border, effective recovery of rare species

must include coordinated assessments of the biological requirements

of selected species through combinations of inventory, monitoring,

and research. Refuge staff and collaborators are monitoring species

population trends and conducting research to identify management

actionsthatwillsupportspeciesrecoveryinefficient andcost effective

ways. Various refuge monitoring projects are providing information

thatmay be usefulin assessing the effects of various borderimpacts on

wildlife populations, and these projects can each function to produce

data used by border managers in multiple ways by (1) determining

the presence, absence, and relative abundance of species and serving

to provide basic inventories and baselines upon which to compare

against in the future; (2) documenting use of habitat types by various

targetspecies and identifying important wildlife movement corridors

betweenArizona andSonora;(3)documentingthedynamics andlongterm changes in wildlife use of habitats; (4) determining the impacts

of border related activities on habitat use by animal populations; (5)

identifying areas of potential conflict between humans and wildlife;

and (6) evaluating the impacts that illegal human activities have on

plant and animal populations and on animal movement patterns. For

example, to learn more about the groundwater aquifer conditions

and trends relative to climate change or other influences within the

Leslie Creek and San Bernardino Valley watersheds, monitoring

wells have been installed to provide a continuous record of water

levels on both private and Refuge lands. This data has provided

information concerning groundwater and surface water relationships

in the watershed, has substantially increased an understanding of the

hydrology of Leslie Creek, and shows a declining trend in the level of

the aquifer at LCNWR likely related to declining winter precipitation

in the Chiricahua Mountains (Broska 2009a).

Monitoring wells in the San Bernardino Valley are documenting

that groundwater levels have been raised in some locations (Broska

2009b), and the Río San Bernardino now flows continuously and perennially from north of the U.S. border on SBNWR southward across

Rancho San Bernardino to well beyond Mexican Highway 2. To help

determinehydrologicalrelationships amongindividualsprings,ponds,

and wells on SBNWR and the adjacent privately owned Johnson

Historical Museum of the Southwest (Slaughter Ranch), Refuge staff

is collecting samples and analyzing water chemistry. This data will

benefit management of federally listed species while also providing

information for Slaughter Ranch managers and visitors on the safety

of consuming the water.
8 USDA Forest Service Proceedings RMRS-P-67. 2013

Radke National Wildlife Refuge Management on the United States/Mexico Border

	 Partners conduct annual monitoring of native fish populations on

both sides of the border, and on both private and Refuge lands, to

determine and documentfish health,fecundity, and population trends.

Population monitoring of other federally listed species, such as the

Huachuca Water Umbel (Lilaeopsis schaffneriana var. recurva), the

Cochise pincushion cactus(Coryphantha robbinsorum), the Chiricahua leopard frog (Lithobates chiricahuenis), and the San Bernardino

springsnailisaccomplishedperiodicallytohelpdocumenttherecovery

of these species, to identify threats, and to increase our understanding

of ecological associations(Malcom and others 2005; Varela-Romero

and Myers 2010).

	 Long-term monitoring of breeding bird species is being accomplished at both SBNWR and LCNWR to provide data on population

and demographic parameters for targeted species. The information

collected is providing a better understanding of the productivity,

survivorship, and population trends of avian communities on the

refuges.Basic long-termreptile andamphibianpopulationmonitoring

is being conducted on these Refuges to help document species richness, understand ecological trends, investigate population dynamics

and the roles of rare species, and to help support resource management decisions. SBNWR and LCNWR serve a large assemblage

of wildlife species and provide important landscape features in the

form of linear riparian areas. The use of such riparian corridors by

large mammals is not well documented (Bennett 1990; Laurance

and Laurance 1999). Increasing human activities and development

in the San Bernardino Valley might be expected to impact the natural

movement and behavior of large mammals, and wildlife movement

corridors are expected to become even more important, making our

better understanding of these areas crucial to effective management.

Habitat and Population Management and 

Protection

	 Managing and protecting existing and historic habitats and wildlife

populations is being accomplished by the Service and conservation

partners using a variety of effective mechanisms, including the Endangered Species Act Safe Harbor Program, Habitat Conservation

Planning for rare species, Service challenge cost share agreements,

acquisitionof conservationeasements,rights-of-ways,memorandums

of understanding, and other collaborative approaches. Such cooperation is expected to increase in helping recover native plants, fish, and

wildlife asfundingbecomemore limiting.Thesemechanismsprovide

a framework for establishing creative partnerships with the ultimate

goal of reducing conflicts between listed species and planned activities on both public and private lands. For example, the impacts of

climate change for the southwestern United States are anticipated to

include higher temperatures and lower levels of precipitation. These

changes will dramatically impact the ecology of the region. Wildlife

migration routes may change, making it important that landscapes

and the crucial corridors that interconnect private, state, and federal

lands be protected and enhanced, along with the fish and wildlife

populationsthattheysupport. Acoordinatedapproachtomanagement

activities that compliment all the landowners in the San Bernardino

Valley must be progressive and recognize the unique needs of each

partner.

Species Reintroductions

	 When managing the recovery of rare plants and animals, it is often

important to assess potential reintroductions of key species into appropriate habitats, and this is occurring on and adjacent to SBNWR

and LCNWR. The Service has an ongoing program of reintroducing

federally listed threatened or endangered fish including the Yaqui

chub, Yaqui topminnow, beautiful shiner, and Yaqui catfish into suitable habitats on private and public lands in the Río Yaqui watershed.

A major positive step in the recovery of the Chiricahua leopard frog

is being accomplished through an effective partnership between the

Service, Arizona Game and Fish Department, U.S. Forest Service,

and the American Museum of Natural History’s Southwestern Research Station near Portal, Arizona. LCNWR is one of the last sites

in southeast Arizona where Chiricahua leopard frogs occur naturally

in the wild; however, this protected habitat for the frog is rapidly disappearing because increased extreme drought conditions are leaving

the Refuge without flow in Leslie Creek. Tadpoles were therefore

transportedfromLCNWRtoSouthwesternResearchStationfacilities

where they were raised in a controlled environment and their progeny

are then being released back into suitable habitat in the Chiricahua

Mountains. The endemic San Bernardino springsnail currently exists

inmultiple isolatedmetapopulationsin bothArizona andSonora,with

such populationsfurther fragmented by the international border. This

rare speciesisbeinghelpedthroughrestorationof appropriatewetland

habitat on Slaughter Ranch and SBNWR. Once adequate habitat isin

place,springsnailsfrom adjacent metapopulations will be restored to

areas they once populated without human intervention. 

Non-Native Species Management 

	 Introduction of non-native plants and wildlife into environments

where they did not occur naturally can have devastating effects on

native populations (Bazzaz 1986; Courtenay and Stauffer 1984;

D’AntonioandVitousek1992;McLaughlin2002;RosenandSchwalbe

1996; Rosen and Schwalbe 1998; Scott 1992). This is the case with

the Asian tapeworm (Bothriocephalus acheilognathi) and bullfrog

(L. catesbeiana), both of which are now established in the San Bernardino Valley. The Asian tapeworm is adversely affecting recovery

of rare cyprinid fish species (Kline and others 2007). Populations of

the Chiricahua leopard frog, lowland leopard frog (L. yavapaiensis),

and Mexican garter snake (Thamnophis eques) have all experienced

severe decline or have even been eliminated in the area as a result of

bullfrog depredation (Rosen and Schwalbe 2002).

	 Continuing efforts are made by trainedRefuge staff and volunteers

to control the spread of non-native species from Refuge lands, however, total control will never be achieved given the complexity of the

landscape and the widespread establishment and abundance of these

species in the area. Fortunately, the vast majority of plant species in

the area are natives, leaving the ecosystem relatively free from exotic

plant invaders. Nonnative plantsthat are a current concern to Refuge

managers include Russian thistle (Salsola iberica), Johnson grass

(Sorghum halapense),Lehmann’slovegrass(Eragrostis lehmanniana),

Malta starthistle (Centaurea melitensis), giant reed (Arundo donax),

saltcedar(Tamarix ramosissima),andbufflegrass(Pennisetum ciliare).

The existence of all these species requires regular monitoring and

control to reduce their spread and their resulting negative impacts to

the recovery of rare native species. In some cases, if nonnative species were to establish themselves on the Refuges, restoration work

being done would be severely compromised. Given the difficulty

and expense of controlling non-native plants and animals once they

have established themselves in an environment, efforts must focus

on preventing the initial introduction of such species.
USDA Forest Service Proceedings RMRS-P-67. 2013 9

National Wildlife Refuge Management on the United States/Mexico Border Radke

Outreach and Education

	 Under the National Wildlife Refuge System Improvement Act of

1997, national wildlife refuges offer several “priority public uses,”

includinghunting,fishing,wildlifeobservation,wildlifephotography,

environmental education, and interpretation when such activities are

compatible with the purpose for which the refuge was established.

Arizona’s Cochise County is a major international tourist destination

dueprimarilytotheregion’sinterestinghumanhistoryandalsobecause

of the region’s unique and abundant natural history. SBNWR and

LCNWR are two of the few refuges administered by the Service that

were created specifically to protect rare native fish. Protection of this

border region and restoration of associated fish and wildlife habitats

and populations in both the United States and in Mexico by private

landowners, conservation organizations, and government agencies

including DHS will help ensure that the ecological balance that has

been effective for centuries will continue to provide the conditions

necessary for all fish, wildlife, and humans dependent upon the Río

Yaqui Basin.

Acknowledgments

	 I thank reviewers Jonathan Andrew and Larisa Ford for providing

suggestions that improved this manuscript. The findings and conclusions in this paper are those of the author and do not necessarily

represent the views of the U.S. Fish and Wildlife Service.

References

Austin,V.In press.Cross border fish and wildlife recovery and habitatrestoration in the San Bernardino Valley. In The border wall between Mexico and 

the United States. Venues, mechanisms, and stakeholders for a constructive 

dialogue, ed. A. Cordova. El Colegio de la Frontera Norte.

Bazzaz, F.A. 1986. Life history of colonizing plants: Some demographic,

genetic, and physiological traits. In Ecology of biological invasions on 

North America and Hawaii, ed. H.AMooney and J.A. Drake, 96-110. New

York: Springer-Verlag.

Bennett, A.F. 1990. Habitat corridors: Their role in wildlife management and

conservation. Department of Conservation and Environment, Melbourne.

Billington, C.C., Gimblett, R., Krausman, P.R. 2010. Implications of illegal

border crossing and drug trafficking on the management of public lands.

In Southwestern desert resources, ed. W. Halvorson, C. Schwalbe, and C.

van Riper III, 109-122. University of Arizona Press, Tucson.

Broska, J. 2009a. Water resources of the Leslie Canyon National Wildlife

Refuge,Arizona.ResearchPaper.Albuquerque,NewMexico:U.S.Department of the Interior, Fish and Wildlife Service. 28 pp.

Broska, J. 2009b. Water resources of the San Bernardino National Wildlife

Refuge,SanBernardinoValley,Arizona.ResearchPaper.Albuquerque,New

Mexico: U.S. Department of the Interior, Fish and Wildlife Service. 12 pp.

Brown,J.H.,Kodrick-Brown,A.1996.Biodiversityintheborderlands.Natural

History 105:58-61.

Cohn,J.P. 2007.The environmentalimpacts of a borderfence.BioScience, 57:96.

Courtenay,W.R., Stauffer,J.R., eds. 1984. Distribution, biology, and management of exotic fishes. Baltimore: Johns Hopkins University Press.

Christen,D.C.,Matlack,G.R. 2009.The habitat and conduitfunctions ofroads

inthespreadofthreeinvasiveplantspecies.BiologicalInvasions11:453-465.

D’Antonio, C.M., Vitousek, P.M. 1992. Biological invasions by exotic

grasses, the grass/fire cycle, and global change. Ann. Rev. Ecological

Systems 23:63-87.

Duncan, D.K., Fernandez, E., McCasland, C. 2010. The U.S.-Mexico border

and endangered species. In Southwestern desert resources, ed. W. Halvorson, C. Schwalbe, and C. van Riper III, 123-130. University of Arizona

Press, Tucson.

Forman,R.T.T.,Alexander,L.E.1998.Roads andtheirmajor ecological effects.

Annual Review of Ecology and Systematics 29:207-231.

Glista, D.J., DeVault, T.L., DeWoody, J.A. 2009. A review of mitigation

measures for reducing wildlife mortality on roadways. Landscape and

Urban Planning 91:1-7.

Kline,J.,Archdeacon,T.,Iles,A.,Bonar, S.A. 2007. Factorsinfluencing distribution of introducedAsian tapeworm and effects on selected southwestern

fishes (Yaqui topminnow and Yaqui chub). Heritage Program, Arizona

Game and Fish Department, Phoenix. 55pp.

Lalo, J. 1987. The problem of roadkill. American Forests 50:50-52.

Laurance, S.G., Laurance, W.F. 1999. Tropical wildlife corridors: use of

linear rainforest remnants by arboreal mammals. Biological Conservation

91: 231-239.

Malcom,J.W., Radke, W.R., Lang, B.K. 2005. Habitat associations of the San

Bernardino Springsnail, Pyrgulopsis bernardina (Hydrobiidae).Journal of

Freshwater Ecology 20:71-77.

McLaughlin, S.P. 2002. Exotic plant species in the western United States. In

Invasive exotic species in the Sonoran region, ed. B. Tellman. 47-62. Tucson: University ofArizona Press and TheArizona-Sonora Desert Museum.

Romin, L.A., Bissonette,J.A. 1996. Deer-vehicle collisions:status ofstate monitoring activities and mitigation efforts. Wildlife Society Bulletin 24:276-283.

Rosen, P.C., Schwalbe, C.R. 1996. Bullfrog impacts on sensitive wetland

herpetofauna of the San Bernardino National Wildlife Refuge. Heritage

Program, Arizona Game and Fish Department, Phoenix. 47 pp.

Rosen, P.C., Schwalbe, C.R. 1998. Status of native and introduced species of

aquatic herpetofauna at SanBernardinoNationalWildlifeRefuge.Heritage

Program, Arizona Game and Fish Department, Phoenix. 52 pp.

Rosen, P.C., Schwalbe, C.R. 2002. Widespread effects of introduced species

on reptiles and amphibiansin the Sonoran Desert region. In Invasive exotic 

species in the Sonoran region, ed.B.Tellman. 220-240.Tucson: University

of Arizona Press and The Arizona-Sonora Desert Museum.

Scott,N.J.Jr.1992.RanidfrogsurveyoftheGrayRanchwithrecommendations

for management of frog habitats, August 1990-September 1991. Report to

Gray Ranch, Animas, New Mexico.

Sayre, N.F., Knight, R.L. 2009. Potential effects of United States-Mexico

border hardening on ecological and human communities in the Malpai

borderlands. Conservation Biology 24(1):345-348.

Segee, B.P., Neely, J.L. 2006. On the line: the impacts of immigration policy

on wildlife and habitat in the Arizona borderlands. Defenders of Wildlife,

Washington, D.C.

Shepard,D.B.,Kuhns,A.R.,Dreslik,M.J.,Phillips,C.A.2008.Roadsasbarriersto

animalmovementinfragmentedlandscapes.AnimalConservation11:288-296.

Trombulak, S.C., Frissell, C.A. 2000. Review of ecological effects of roads

on terrestrial and aquatic communities. Conservation Biology 14:18-30.

U.S. Border Patrol. 2012. Public Lands Weekly Activity Report ending April

21, 2012. Unpublished data on file at: U.S. Border Patrol Tucson Sector

Headquarters, Tucson, AZ, 8 p.

U.S. Fish andWildlife Service. 1994.Yaqui Fishes Recovery Plan. USDI Fish

and Wildlife Service, Albuquerque, New Mexico. 48 pp.

U.S. Fish and Wildlife Service. 1995. San Bernardino and Leslie Canyon

National Wildlife Refuges Comprehensive Management Plan 1995-2015.

USDI Fish and Wildlife Service, Albuquerque, New Mexico. 113 pp.

U.S. Fish and Wildlife Service. 2011.EnvironmentalAssessment for San Bernardino National Wildlife Refuge projects associated with mitigation for

Department of Homeland Security tactical infrastructure, Cochise County,

Arizona.USDIFishandWildlifeService,Albuquerque,NewMexico.53pp.

Van Schoik, D.R., Lelea, E., Cunningham, J. 2006. Sovereignty, borders, and

transboundary biodiversity: Turning a potential tragedy into a true partnership an introduction to the theory and practice of conservation biology in the

security bisected U.S.-Mexico border region. In The U.S.-Mexican Border 

Environment: transboundary ecosystem management, ed. K. Hoffman.

1-68. San Diego: San Diego State University Press.

Varela-Romero, A., Myers, T.L. 2010. Genetic evaluation of springsnails

in the San Bernardino River watershed of Sonora, Mexico, in relation

to Pyrgulopsis bernardina. Heritage Program, Arizona Game and Fish

Department, Phoenix. 13pp.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
10 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Biodiversity in the Madrean Archipelago of 

Sonora, Mexico 

Thomas R. Van Devender, Sergio Avila-Villegas, and Melanie Emerson

Sky Island Alliance, Tucson, Arizona

Dale Turner

The Nature Conservancy, Tucson, Arizona

Aaron D. Flesch 

University of Montana, Missoula, Montana

Nicholas S. Deyo

Sky Island Alliance, Tucson, Arizona

Introduction

	 Flowery rhetoric often gives birth to new termsthat convey images

and concepts, lead to inspiration and initiative. On the 1892-1894

expeditiontoresurveytheUnitedStates-Mexicoboundary,Lieutenant

David Dubose Gaillard described theArizona-Sonora borderlands as

“bare, jagged mountains rising out of the plains like islands from the 

sea” (Mearns 1907; Hunt and Anderson 2002). Later Galliard was

the lead engineer on the Panama Canal construction project.

	 In 1951, Weldon Heald, a resident of the Chiricahua Mountains,

coined the term ‘Sky Islands’for the ranges in southeastern Arizona

(Heald 1951). Frederick H. Gehlbach’s 1981 book, Mountain Islands 

and Desert Seas: A Natural History of the US-Mexican Borderlands,

provided an overview of the natural history of the Sky Islands in

the southwestern United States. The ‘desert seas’ in the valleys are

desert grassland and Chihuahuan desertscrub in the north, Sonoran

desertscrub to the west, and foothillsthornscrub in the south. Variants

of the term include Sky Island region (McLaughlin 1995; Gottfried

and others 2005; Skroch 2008),sky-island ranges(Felger andWilson

1995,Fishbeinandothers1995),andskyislandbioregion(Fishbeinand

others 1995; Skroch 2008).Today we mostly use ‘Sky Island Region’

with capital letters.The best name in Spanish isIslas Serranas, rather

than the literal translation Islas del Cielo, because cielo means both

‘sky’ and ‘heaven.’ The analogy to oceanic islands (Warshall 1995)

is limited because Sky Islands differ from true insular areas in high

species diversity, low local and regional endemism, and low percentages of non-native species (McLaughlin 1995). While any isolated

area is a potential area for speciation in small populations, there are

relatively few species restricted to the Sky Island Mountains.

	 The term ‘Madrean’ comes from the Sierra Madre. The Mexican

Plateau is a vast area of grasslands and desertscrub between the

Rocky Mountains in New Mexico and ca. 1,300 km to the south the

Trans-Mexican Volcanic Belt in south-central Mexico.The Plateau is

open to incursions of frigid Arctic air from the north, and the Sierra

Madres Oriental and Occidental create a double rain shadow and

the Chihuahuan Desert. Madrean is a general term used to describe

things related to the Sierra Madres. In a biogeographical analysis of

theherpetofaunaofSaguaroNationalMonument,UniversityofArizona

herpetologist and ecologist Charles H. Lowe was probably the first to

use the term ‘Madrean Archipelago’to describe the Sky Island ranges

between the Sierra Madre Occidental in Sonora andChihuahua and the

Mogollon Rim of central Arizona (Lowe, 1992). Warshall (1995) and

McLaughlin (1995) expanded and defined the area and concept.

Biodiversity

	 In 2007, Conservation International named the Madrean Pine-oak

Woodlands as a global biodiversity hotspot. This was a very large

area that included the Sierra Madre Oriental in eastern Mexico, the

Sierra Madre Occidental (SMO) in western Mexico, and the Sky Island ranges north of them into Texas, New Mexico, andArizona. The

Sierra Madre Occidental extends in western Mexico from Zacatecas

and Jalisco north to Chihuahua and Sonora. The southwestern United

States is famous for its diversity of animals and plants, and in many

groups diversity increasessouthward in Sonora into the Sierra Madre

Occidental and the New World tropics.

	 Species richness is enhanced in the Sky Island Region for many

reasons. The western half of North America is mountainous, with

topographically diverse habitats and vegetation zoned along elevational-climatic gradients. Biotic communities are concentrated in

small geographic areas compared to the eastern United States where

vegetation mostly changes along a north-south latitudinal gradient.

	 The Madrean Archipelago is a convergence zone for five biotic

provinces(fig. 1). TheRockyMountains andColoradoPlateausto the

north have temperate climates.In themid-continent, the grasslands of

theGreatPlains extendfromCanada southtoNewMexico,Texas, and

the Mexican Plateau. In the south, a mosaic with desert grassland in

the valleys and Chihuahuan desertscrub on adjacent limestone slopes

extendswestward into southeasternArizona and northeasternSonora.

On the western edge of the Sky Island Region, Sonoran desertscrub

surrounds Sky Island ranges. In the south, more tropical oak woodlands and pine-oak forests are in the Sierra Madre Occidental and

Sky Island mountaintops. In the lowlands, foothillsthornscrub (FTS)

and tropical deciduous forest are present. The transition between the

NewWorld tropics and the northern temperate zone is at about 29°N in
USDA Forest Service Proceedings RMRS-P-67. 2013 11

Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

east-central Sonora.The northern limits of FTS in Sonora are at about

30°11’N east of Sinoquipe in the Río Sonora Valley and 30°26’N at

Presa Angostura on the Río Bavispe at the southern end of the Sierra

El Tigre. FTS does not reach Arizona, but the distributions of quite

a few FTS species cross the border into southern Arizona in desert

grassland or oak woodland.

Sky Island Bioblitzes

	 Today the term ‘bioblitz’ is popular, meaning an intensive effort

in a short period to document the diversity of animals and plants in

an area. The first bioblitz in the southwestern United States was the

1848-1855 survey of the new boundary between the United States

and Mexico after the Treaty of Guadalupe Hidalgo of 1848 ended

the Mexican-American War. The border between El Paso, Texas, and

theColoradoRiverinArizona wassurveyed in 1855-1856,following

the Gadsden Purchase in 1853. Besides surveying and marking the

border with monuments, these expeditions made extensive animal

and plant collections, often by army physicians. Botanists John M.

Bigelow, Charles C. Parry, Arthur C. V. Schott, Edmund K. Smith,

George Thurber, and Charles Wright (fig. 2) and zoologist John H.

Clark collected between the Santa Rita Copper Mines(= Silver City),

New Mexico and Santa Cruz, Sonora in the modern Arizona-Sonora

borderlands. On the second United States-Mexico Boundary Survey

in 1892-1894, Edgar A. Mearns (fig. 3) collected 30,000 animal and

plantspecimens.Themammals were published in the book Mammals 

of the Mexican Boundary of the United States (Mearns 1907). In

1890, NorwegianCarl S. Lumholtz led an anthropological expedition

through northeastern Sonora. He published a popular travelogue in

1902. Sweden’s Carl V. Hartman was a botanist in charge of general

biological collections (fig. 4).

Río Mayo-Yécora Baseline

In the 1930s, Howard Scott Gentry was a recent graduate from the

University of California at Berkeley. In search of employment, he

went to the NewWorld tropicsin the Álamos area in southern Sonora,

in hopes of selling biological specimens to museum collections. His

plant collections were published in the bookRío Mayo Plants(Gentry

1942).In the 1970s, 80s, and 90s, Paul S. Martin led expeditionsfrom

the University of Arizona, which retraced Gentry’s travels, explored

new areas, and resulted in a revision of the Río Mayo flora (Martin

andothers1998).Thislarge area includestheSierraMadreOccidental

in eastern Sonora and western Chihuahua.

	 Studies of the flora and fauna of the Yécora area in the SMO in

eastern Sonora provide baseline information needed to compare

biotas in the Madrean Archipelago with the mainland Sierra Madre

Occidental, putting the biodiversity of Sky Island ranges in regional

context. Mexican Federal Highway 16 (MEX 16), one of the few

highways that crosses the SMO, provides a west-to-east elevational

transect from the Río Yaqui (180 m) east to the Chihuahua border,

42 km east of Yécora (fig. 5).

Figure 2—(A) The U.S.-Mexico Boundary and Gadsden Purchase 

Surveys of 1848-1857 went through Guadalupe Canyon (photo by T. 

R. Van Devender). (B) Arthur C.V. Schott and (C) Charles Wright were 

expedition botanists.

B

C

A

Figure 3—(A) The second U.S.-Mexico Boundary Survey in 1892-

1894 was led by Edgar A. Mearns. (B) The Arizona-Sonora Boundary 

is between the Sierra San José and the Huachuca Mountains (photo 

by T. R. Van Devender).

Figure 1—Map of the biotic convergence in the Madrean Archipelago 

(map by Nick Deyo).

B

A
12 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Avila-Villegas, Emerson, and others Biodiversity in the Madrean Archipelago of Sonora, Mexico 

de Yécora region on 36 fieldtrips from 1995 to 2008. This flora with

1,776 taxa in 3,300 km2 is very diverse, ca. 30% more diverse than

the Huachuca Mountains in southeastern Arizona (Reina-G. and Van

Devender 2005). There are 2,125 observations of 301 species of birds

fromtheMunicipio deYécora in theMadreanArchipelagoBiodiversity

Assessment (MABA) database (http://www.Madrean.org).

Borderlands Exploration

Higher Sky Island mountain rangessuch asthe Sierras de losAjos,

Elenita, Marquita, and Tigre have extensive pine-oak forest and

mixed-conifer forests on the highest peaks (fig. 6; Marshall 1957).

The biotas of these areas are temperate mixtures of Sierra Madrean

and Rocky Mountain species.

	 Inthe1930s and1940s,theUniversityofMichiganhadaveryactive

biological exploration program in Mexico. In 1935, Barry Campbell

collected amphibians and reptilesin the Sierra elTigre, Sonora. From

1938-1941, Stephen S. White led three botanical expeditions to explore the Río Bavispe region of northeastern Sonora (fig. 7). The Río

Bavispe flora with 1200 species (currently 995 taxa after revision) in

549 genera in 114 familiesfrom a wide range of habitatsin the region

was the first modern flora in the Sonoran borderlands (White 1948).

	 Ornithologist and intrepid explorer Joe T. Marshall studied the

birds and dominant plants of pine-oak woodland in mountain ranges

in southeastern Arizona, northeastern Sonora, and in the northern

Sierra Madre Occidental in Chihuahua in the summers of 1951-1953

Figure 4—The Lumholtz Expedition. (A) Carl S. Lumholtz. 

(B) Botanist Carl V. Hartman. (C) Sketch of the church in 

Bacadéhuachi in Lumholtz (1902).

C

A

B

Figure 5—Sunset in the Sierra Madre Occidental west of Yécora, 

Sonora (photo by Erik F. Enderson).

	 Between 1953 and 2005, herpetologist-ecologist Charles H. Lowe

collected amphibians and reptiles in the Yécora area for the University of Arizona Herpetological Collection on 33 trips involving 26

people. From 2004 to 2008, Erik F. Enderson and Robert L. Bezy

documentedtheYécora areaherpetofauna throughphotographyon40

field days. Van Devender and Ana L. Reina-Guerrero at the ArizonaSonora Desert Museum (ASDM) studied the flora of the Municipio
USDA Forest Service Proceedings RMRS-P-67. 2013 13

Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

(Marshall, 1957). In Sonora, he worked in the SierrasAconchi,Azul,

“Cananea” (=Elenita-Mariquita),Oposura (=LaMadera), andPinitos.

	 Beginning in 1963, Raymond M. Turner, J. Rodney Hastings and

later Tony L. Burgess made numerous trips to Sonora and Baja,

California, to document the distributions of 414 species of plants in

the Sonoran Desert. Their observations, collections, and ecological

summaries were included in the book Sonoran Desert Plants. An 

Ecological Atlas (Turner and others 1995).

	 Beginning in 1964, Stephen M. Russell and Gale Monson went

to Sonora to observe birds on 150 trips and 692 field days. Their

observations of 525 species were published in the book The Birds 

of Sonora (Russell and Monson 1998). Beginning in 2000, Aaron D.

FleschexpandedontheirfieldobservationsinnorthernSonora (Flesch

2008).

	 In the 1980s and 1990s,theUnitedStatesFish andWildlifeService,

Área Natural Protegida (ANP) Ajos-Bavispe (a Comisión Nacional

de Áreas Naturales Protegidas [CONANP) reserve), University of

Arizona, and The Nature Conservancy Plant Collections organized

expeditions to the Sierra de los Ajos just east of Cananea: Frank W.

Reichenbacherin1982,PaulS.Martinin1983and1991,PeterWarren

and Esther Saucedo in 1989, and Richard S. Felger, Mark Fishbein,

George M. Ferguson, Dale S. Turner, James Malusa, Florentino

Garza-S., and Martín Haro in October 1992 and July 1993 (Fishbein

and others 1995).Turner, Garza-S., and Haro returned on a zoological

expedition in 1998 along with Cecil Schwalbe, Taylor Edwards, Carl

Olson, Yar Petryszyn, Dave Prival, and others (Alberti and others

1998).

	 With a few exceptions, documentation of animals and plantsin the

Arizona-Sonora borderlands was mostly neglected after the 1848-

1857 and 1892-1894 boundary surveys. Even common, widespread

species had not been collected or observed. Beginning in 2001, Van

Devender andReina-Guerrero (2005) began collecting and observing

plants in La Frontera, the area in northern Sonora within 100 km of

the Arizona border, as part of several inventory projects. 

	 Leonardo Varela-Espinosa and Alberto Búrquez-Montijo at the

Universidad Nacional Autónoma de México in Hermosillo studied

the flora of the Sierra San Javier on seven tripsfrom July 1996 to May

1997(Varela-Espinosa2005).TheSierraSanJavieristhesouthernmost

Sky Island with tropical deciduous forest below oak woodland.

Since 1994, Herbario Universidad de Sonora (USON) has grown into

a major regional collection. In 2003-2004, Curator J. Jesús SánchezEscalante, Manuel Espericueta-Betancourt, and Reyna A. CastilloGámez fromUSON, andVanDevender andReina-GuerreroatASDM

studied the flora of the Sierra de Mazatán (Sánchez-Escalante and

Figure 7—Stephen S. White and José Vera-Santos in the Sierra El Tigre 

on 13 October 1941. 

Figure 6—(A) View from the Sierra de los Ajos (photo by Dale S. Turner). 

(B) Mixed-conifer forest in Sierra El Tigre (photo by George M. Ferguson).

A

B
14 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Avila-Villegas, Emerson, and others Biodiversity in the Madrean Archipelago of Sonora, Mexico 

others 2005). Thisrange isthe southwestern-most Sky Island located

70 km east of Hermosillo in central Sonora. Between March 2009

and September 2011, Sánchez-Escalante, Denise Z. Ávila-Jiménez,

David A. Delgado-Zamora, and Liliana Armenta-Cota from USON,

and Van Devender andReina-Guerrero at Sky IslandAlliance studied

the flora of Rancho El Aribabi in the Sierra Azul on seven field trips.

El Aribabi Conservation Ranch in the Sierra Azul was designated an

Área Natural Protegida Privada by CONANP in March 2011.

	 In July-August 2005, biologists from the University of Arizona

and the Sonoran Desert Research Station inventoried the fauna and

flora along the Río Áros, a tributary of the Río Yaqui in east-central

Sonora (O’Brien and others 2006). Participants included Charles

O’Brien, Aaron D. Flesch, Eric Wallace, Michael D. Bogan, Samia

Carrillo-Percástegui, Sky Jacobs, and Charles van Riper III.

	 InApril-May 2008, Sky IslandAlliance organized an expedition to

the Sierra El Tigre to observe plants and animals (Avila-Villegas and

Jacobs 2008). Participants were SergioAvila-Villegas, Paul Condon,

SkyJacobs,MattSkroch,RobertVilla,RodolfoVilla, andlandowners.

Madrean Archipelago Conferences

	 As recently as two decades ago, few people knew of the Madrean

Archipelago as a distinct region and the evidence for its importance

was scattered among many disciplines in two countries. In September 1994, a conference entitled Biodiversity and Management of the 

Madrean archipelago: The Sky Islands of Southwestern United States 

and Northwestern Mexico was organized by the Rocky Mountain

Forest and Range Experiment Station, U.S. Forest Service, along

with Sky Island Alliance (DeBano and others 1995). There were

69 presentations and 20 posters. The second Madrean Archipelago

Conference in May 2004 was entitled Connecting Mountain Islands 

and Desert Seas: Biodiversity and Management of the Madrean 

Archipelago. There were 93 presentations and 14 posters. The lack

of information from the Sky Island Region in Mexico was noted in

both conferences.

	 InMay 2012,the thirdMadreanArchipelagoconference assembled

the current state of our knowledge about the unique natural and cultural resources of the Madrean region and continued the discussion

of management practices useful for maintaining those resources with

24 sponsoring organizations.It brought togetherresearchers, partners

in resource stewardship, land managers, educators and students,

government officials, consultants, and the interested public from

both sides of the border to examine the Madrean Archipelago of the

southwestern United States and northwestern Mexico. There were

151 presentations and posters, including substantial advances in the

knowledge of animal and plant distributions in Mexico.

Madrean Archipelago Bodiversity 

Assessment

	 One ofthe conclusions of both oftheMadrean conferenceswasthat

there was an urgent need forinformation from the Mexican portion of

the Sky Island Region. Sky Island Alliance was a principal organizer

for both of these conferences. In spring 2009, the Northern Mexico

Conservation Program at SIA initiated the Madrean Archipelago

Biodiversity Assessment (MABA) project, in part funded by a grant

from the Veolia Environment Foundation. The concept of documenting all plant and animal species in the Mexican Sky Island Region

for use in conservation, land management, research, and education

was proposed by former SIA Executive Director Matt Skroch, Dale

Turner, and Aaron D. Flesch. 

	 The online MABA database (http://www.madrean.org) was created to make observations and images available to the public. The

FLORA half of the database is directly linked to the Southwestern

Environmental Information Network (SEINet) database, which has

data for over 2 million herbarium specimens. The FAUNA half of

the database was a new database. The MABA database differs from

SEInet in that many records are observations rather than specimens,

and that observations are often illustrated with color images. Historical records are from museum collections, literature, field notes and

other databases. New observations are made on fieldtrips to remote

high diversity Sky Island ranges. Study areas were selected based on

the need for biological observations, conservation initiatives, and opportunities to collaborate with landowners, land managers, and local

researchers.

	 While observations are made on all MABA trips, expeditions to

high priority areas with 25-45 participants are especially productive.

These are unprecedented groups of animal and plant specialists,

landowners, agency biologists, university professors and students,

journalists,photographers, andvolunteers(fig.8).In2009-2011,there

were five MABA Expeditions. In September 2009, 17 participants

went to the Sierra San Luis on Cuenca Los Ojos Foundation property.

In March 2010, 25 participants went to the Sierra El Tigre on private

rancheswithintheANPAjos-Bavispe.InAugust2010,39participants,

including professors and students from the nearby Universidad de

la Sierra (UNISIERRA), went to the Sierra La Madera on private

ranches, partially within the ANPAjos-Bavispe. In April-May 2011,

40 participants went to the Ciénega de Saracachi and the Sierra San

Antonio.TheCiénega is an important wetland natural area onRancho

Agua Fría that was proposed as an Área Natural Protegida Estatal

by the Comisión de Ecología y Desarrollo Sustentable del Estado de

Sonora (CEDES)in 2010.The Sierra SanAntonio is privately owned.

In August 2011, 45 participants went to the Sierra Bacadéhuachi,

which ismostly privately owned.Rincón deGuadalupe, owned by the

CatholicDiocese ofHermosillo, has excellent potentialfor protection

as a natural area.

	 TheMABAdatabase isthe primary source of biologicalrecordsfor

the state of Sonora. FLORA contains 112,623 Sonoran observations,

including 20,606 fromMABA, 20,331 fromthe book Sonoran Desert 

Plants. An Ecological Atlas (Turner and others 1995), and 13,285

from the Comisión para el Uso y Conocimiento de la Biodiversidad

(CONABIO). FAUNA contains 128,843 Sonoran records, including

60,856 from MABA, 39,022 from the book The Birds of Sonora (Russell and Monson, 1998), 33,993 from the eBird database, and 27,404

from CONABIO. Species numbers in the MABA database provide 

preliminary diversity estimates for the Madrean Archipelago in Sonora. There are ca. 2880 plant taxa in northeastern Sonora. Insects are 

very diverse, but only 1380 taxa are documented in the Sky Islands 

Region. Vertebrates are better represented: fish (39 species), amphibians and reptiles (104 species), birds (358 species), and mammals (76 

species). Surprisingly, diversity appears to decrease southward, but it 

probably reflects the north-northwest to south-southeast orientation 

of the ranges, and decreasing area and fewer biological inventories 

to the south.

Conclusions

The MABA project has pulled together information from biological expeditions in the Madrean Archipelago from the last 160 years.

The knowledge of plant and animal distributions has increased

dramatically since 1950, reflecting fieldwork by American botanists,

herpetologists, ornithologists, and mammalogists, the development

of biology programs at the Universidades de Sonora and de la Sierra,
USDA Forest Service Proceedings RMRS-P-67. 2013 15

Biodiversity in the Madrean Archipelago of Sonora, Mexico Van Devender, Avila-Villegas, Emerson, and others

the CONABIO national biodiversity database, and the Sonoran state

Área Naturales Protegidas program by CEDES. The 2010 book Diversidad Biológica de Sonora, edited by Francisco Molina-Freaner

and Van Devender, summarized the state of knowledge of plants,

animals, and vegetation in Sonora. These accomplishments are only

thebeginning—theopportunitiesfor additionalnaturalhistorystudies

in the Madrean Archipelago are enormous!

Acknowledgments

	 For their support and participation on MABA Expeditions, we

thank Mario Cirett and Rosa Elena Jiménez-Maldonado of ANP

Ajos-Bavispe;GertrudisYanes-Arvayo,HugoSilva-Kurumiya,María

de la Paz Montañez-A., and their students of UNISIERRA; J. Jesús

Sánchez-Escalante of UNISON; Martín Villa-Andrade, Gonzálo

Luna-Salazar, Martín Reyes-Juárez, and Rogelio Molina-Freaner;

and Enrique Yescas of SonoraEs magazine. For their field observations and great company, we thank Ana Lilia Reina-Guerrero, John

Palting, George Ferguson, Chip Hedgcock, Sky Jacobs, Carl Tomoff,

Steve Hale, Robert Villa, and Trevor Hare. We thank cocineros del 

campo Omar Gutiérrez-Castro and Roberto Torres-García for tortillas y frijoles. Marc Trinks and Caroline Patrick-Birdwell helped

with MABA trip logistics. We thank Melanie Emerson and the SIA

Board of Directors for their support of the MABA project. Bioblitz

FathersPaulMartin,SteveRussell,WendellMinckley,CharlesLowe,

Raymond Turner, Howard Gentry, Paul Martin, Stephen White, Joe

Figure 8—MABA group at Rincón de Guadalupe in the Sierra Bacadéhuachi in August 2011 (photo by Chris Marzonie).

Marshall, Edgar Mearns, John Lemmon, John Bartlett, and William

H. Emory walked the biodiversity path long before us.

References

Alberti, Barbara; Edwards, Taylor; Fisher, Jessa; [and others]. 1998. Biotic

and geologic reconnaissance and inventory of Sierra losAjos. 01-06August

1998 report to Reserva Forestal Ajos-Bavispe.

Avila-Villegas, Sergio; and Jacobs, Sky. 2008. Trip report. Sierra El Tigre,

Sonora,Mexico.SkyIslandAlliance.Online:http://www.skyislandalliance.

org/misc/Sierra_El_Tigre_Report.pdf

DeBano, Leonard F.; Ffolliott, Peter F.; Ortega‐Rubio, Alfredo; [and others],

coords. 1995. Biodiversity and management of the Madrean Archipelago:

The Sky Islands of southwestern United States and northwestern Mexico.

Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture, Forest

Service, Rocky Mountain Forest and Range Experiment Station.

Felger, Richard S., and Wilson, Michael F. 1995. Northern Sierra Madre Occidental and its Apachian outliers. Pp. 36-59 in DeBano, Leonard F.; Ffolliott,PeterF.;Ortega‐Rubio,Alfredo;[and others], coords.Biodiversity and

management oftheMadreanArchipelago:The Sky Islands ofsouthwestern

United States and northwestern Mexico. Gen. Tech. Rep. RM-GTR-264.

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest

and Range Experiment Station.

Fishbein, Mark, Felger, Richard S.; and Garza-S., Florentino 1995. Another

jewel in the crown: A report on the flora of the Sierra de los Ajos, Sonora,

Mexico.Pp.126-134in DeBano,LeonardF.;Ffolliott,PeterF.;Ortega‐Rubio,

Alfredo;[andothers], coords.BiodiversityandmanagementoftheMadrean

Archipelago:TheSkyIslandsofsouthwesternUnitedStatesandnorthwestern
16 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Avila-Villegas, Emerson, and others Biodiversity in the Madrean Archipelago of Sonora, Mexico 

Mexico. Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture,

Forest Service, Rocky Mountain Forest and Range Experiment Station.

Flesch, Aaron D. 2008. Distribution and status of breeding birds in northern

Sonora, Mexico. Studies in Avian Biology, 37: 23-45.

Gehlbach, Frederick. H. 1981. Mountain islands and desert seas: A natural

history of the US‐Mexican borderlands. Texas A & M University Press,

College Station, TX.

Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster,

Carleton B., comps. 2005. Connecting mountain islands and desert seas:

Biodiversity andmanagement oftheMadreanArchipelago II.Proc.RMRSP-36. Fort Collins, CO: U.S. Department of Agriculture, Forest Service,

Rocky Mountain Research Station.

Heald, Weldon F. 1951. Sky Islands of Arizona. Natural History, 60: 56-63,

95-96.

Hunt,Robert; andW.Anderson,Walter.2002.Arapidbiological andecological

inventory and assessment of the Cajón Bonito watershed, Sonora, Mexico.

Part 1: Natural history. Desert Plants, 18: 3-20.

Lowe, C. H. 1964. Arizona landscapes and habitats. Pp. 3-132 in The Vertebrates of Arizona, Tucson: University of Arizona.

Lowe, Charles H. 1992. On the biogeography of the herpetofauna at Saguaro

National Monument. Pp. 91-104 in Stone, C. P.; and Bellantoni, Elizabeth

S., eds. Proceedings of the Symposium on Research in Saguaro National

Monument, Tucson, AZ.

Lumholtz, Carl S. 1902. Unknown Mexico:ARecord of Five Years’Exploration among the tribes of the Western Sierra Madre; in the Tierra Caliente

of Tepic and Jalisco; and among the Tarascos of Michoacán. New York:

Charles Scribner’s Sons.

Marshall,JoeT.,Jr. 1957. Birds of the pine‐oak woodland in southernArizona

and adjacent Mexico. Pacific Coast Avifauna, 32: 1‐125.

McLaughlin, Stephen P. 1995. An overview of the flora of the Sky Islands,

southeastern Arizona: Diversity, affinities, and insularity. Pp. 60-70 in DeBano, Leonard F.; Ffolliott, Peter F.; Ortega‐Rubio, Alfredo; [and others],

coords. Biodiversity and management of the Madrean Archipelago: The

Sky Islands of southwestern United States and northwestern Mexico. Gen.

Tech. Rep. RM-GTR-264. U.S. Department ofAgriculture, Forest Service,

Rocky Mountain Forest and Range Experiment Station.

Mearns,EdgarA.1907.MammalsoftheMexicanBoundaryoftheUnitedStates.

Part 1. Smithsonian Institution. Bulletin 56. Government Printing Office.

Molina-Freaner, Francisco;. and Van Devender, Thomas R., eds. 2010.

Diversidad Biológica de Sonora, Universidad Nacional Autónoma de

México, Hermosillo.

O’Brien, Charles; Flesch, Aaron D.; Wallace, Eric; [and others]. 2006. Biological inventory of the Río Áros, Sonora, Mexico:Ariver unknown. Final

Report to T&E, Inc.

Reina-G.,AnaL.; andVanDevender,ThomasR. 2005.Floristic comparison of

anArizona ‘sky island’ and the Sierra Madre Occidental in eastern Sonora:

The Huachuca Mountains and the Yécora area. Pp. 154-157 in Gottfried,

Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster, Carleton

B., comps. Connecting mountain islands and desert seas: Biodiversity and

management of the MadreanArchipelago II. Proc. RMRS-P-36. Fort Collins, CO: U.S. Department ofAgriculture, Forest Service, Rocky Mountain

Research Station.

Russell, Stephen M.; and Monson, Gale. 1998. The Birds of Sonora. Tucson:

University of Arizona.

Sánchez-Escalante, J. Jesús; Espericueta-Betancourt, Manuel; and CastilloGámez, Reyna A. 2005. A preliminary floristic inventory in the Sierra de

Mazatán, Municipios de Ures and Mazatán, Sonora, México. Pp118-126

in Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster,

Carleton B., comps. Connecting mountain islands and desert seas: Biodiversity andmanagement oftheMadreanArchipelago II.Proc.RMRS-P-36.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station.

Skroch, Matthew. 2008. Sky Islands of North America. A globally unique

and threatened inland archipelago. Terrain.org Blog, http://www.terrain.

org/articles/21/skroch.htm

Turner, Raymond M.; Bowers,Janice E.; and Burgess, Tony L. 1995. Sonoran

desert plants. An ecological atlas. Tucson: University of Arizona Press.

Varela-Espinosa, L. 2005. Flora y vegetación del la región de San Javier,

Sonora, México. Tesis de Licenciatura, Facultad de Ciencias, Universidad

Nacional Autónoma de México. México D.F.

Warshall, Peter. 1995. The Madrean Sky Island Archipelago: a planetary

overview.Pp.6-18 inDeBano,LeonardF.;Ffolliott,PeterF.;Ortega‐Rubio,

Alfredo;[andothers], coords.BiodiversityandmanagementoftheMadrean

Archipelago:TheSkyIslandsofsouthwesternUnitedStatesandnorthwestern

Mexico. Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture,

Forest Service, Rocky Mountain Forest and Range Experiment Station.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 17

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane G.; Collins, Loa C., comps. 2013. Merging science and management in a rapidly 

changing world: Biodiversity and management of the Madrean Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station. 

Plenary Abstracts

COLE, JULIA. Geosciences and Atmospheric Sciences, University of Arizona, 1040 E 4th St., Tucson, AZ 85721, jecole@email.arizona.edu.

Climate change, megadroughts, and the North American monsoon. Climate change in theSouthwesttranslatesto changesinwater availability,

and records of past hydroclimate help us delineate the natural range of those changes. Past droughts are apparent in tree-ring based records and

emerge from longer paleoclimate records as well. But do these droughts reflect changes in winter or summer moisture? This question is more

than academic: as we move into a warmer world, we can expect generally drier winters and springs. But projections of future climate disagree

with respect to changes in the North American monsoon, and global models do not yet simulate this system well. Here we use paleoclimate

records to explore how summer rainfall has responded during past climate regimes. In this talk I will describe new results from cave-based

climate records that demonstrate how, over the past 7000 years, the monsoon has clearly weakened. This result is consistent with other monsoon

systems globally and with climate model simulations. I will explore implications for future hydroclimate regimes in our Southwest region.

El cambio climático, megasequías y el monzón norteamericano. El cambio climático en el Suroeste Norteamericano induce cambios en la

disponibilidad de agua, y los registros ambientales hidrológicos nos ayudan a delinear el rango natural de esos cambios. Las sequías anteriores

son evidentes en los registros de anillos de árboles (dendroclimatología) y también emergen de registros de paleo-climas más largos. Pero estas

sequías, ¿reflejan cambios en la humedad de invierno o verano? Esta pregunta es más que académica: conforme el mundo se vuelve más cálido podemos esperar inviernos y primaveras más secas. Pero las proyecciones del clima del futuro no coinciden en los cambios en el monzón

norteamericano y los modelos mundiales todavía no simulan bien este sistema. Aquí usamos registros paleo climáticos para explorar como

las lluvias de verano han respondido durante regímenes climáticos anteriores. En esta charla describiré nuevos resultados basados en registros

climáticos de cuevas para mostrar como los últimos 7000 años, el monzón claramente se ha debilitado. Este resultado es consistente con otros

sistemas monzónicos mundiales y con modelos climáticos. Exploraré implicaciones para regímenes hidroclimáticos futuros en nuestra región

del Suroeste Norteamericano..

LYTLE, DAVID A. Department of Zoology, Oregon State University, Corvallis, OR 97331, lytleda@oregonstate.edu.

Floods, droughts, and aquatic biodiversity in the Madrean Archipelago. The Madrean Sky Island region harbors a rich variety of aquatic

habitats, ranging from mountain streamsto lowland rivers and ciénegas. Compared with other regions, MSI aquatic habitats are unusual because

they are patchily distributed acrossthe landscape and strongly influenced by seasonal flooding and drought. Thisfragmented spatial arrangement

coupled with strong disturbance dynamics has shaped both the evolution and ecology of MSI aquatic organisms. Many aquatic insects, fish, and

riparian plants are now known to possess adaptations for surviving extreme events such as flash flooding or prolonged drought. Additionally,

population genetic studies have revealed marked evolutionary divergence of aquatic populations at fine spatial scales. In one case, populations

of an aquatic invertebrate predator have evolved unique behaviors to escape flash floods, and these behavioral adaptations differ in response

to local canyon-scale flood regimes. Disturbance dynamics and spatial isolation also facilitate high aquatic invertebrate biodiversity within a

surprisingly small amount of habitat, with largely distinct communities coexisting within meters of each other. This fine-scale partitioning of

genetic and species diversity is a double edged sword.Although changes in hydrologic regime due to climate change, groundwater pumping and

other factors have eliminated unique populations and communities, the diversity and adaptability of these organisms could provide resilience

to ecological change.

Inundaciones, sequías, y biodiversidad acuática en el Archipiélago Madrense. La región de las Islas Serranas Madrenses alberga una gran

variedad de hábitats acuáticos, desde los arroyos de montañas a los ríos y ciénegas de la tierra baja. Comparados con otras regiones, los hábitats

acuáticos de la región Madrense son diferentes por estar fragmentados y bastante influenciados por inundaciones y sequías de estación. Esta

fragmentación espacial combinada con fuertes dinámicas de disturbio ha influenciado la evolución y ecología de los organismos acuáticos de

la región Madrense. Ahora se sabe que muchos insectos acuáticos, peces y plantas ribereñas están adaptados para sobrevivir eventos extremos,

como son las crecidas o sequías prolongadas. Además, estudios genéticos poblacionales han revelado una marcada divergencia evolutiva de las

poblaciones acuáticas en escalas espaciales pequeñas. En un caso, las poblaciones de un depredador invertebrado acuático han evolucionado

comportamientos únicos para escapar de las crecidas, y estas adaptaciones conductuales difieren en repuesta a los regimenes locales de inundaciones a escala del cañón. Las dinámicas de perturbación y aislamiento espacial también facilitan una alta biodiversidad de invertebrados

acuáticos en un hábitat sorpreseivamente pequeño, con comunidades muy distintas coexistiendo separadas por unos cuantos metros. Esta división en una escala pequeña de diversidad genética y de especies es una espada de doble filo. Aunque los cambios en el régimen hidrológico

debido al cambio climático, extracción de agua subterránea y otros factores han eliminado poblaciones y comunidades únicas, la diversidad y

adaptabilidad de estos organismos podría aumentar la capacidad de recuperación del cambio ecológico.
18 USDA Forest Service Proceedings RMRS-P-67. 2013

Plenary Abstracts/ Resúmenes

STROMBERG, J. C.1, M. S. White1, G. L. Katz2. 1 Arizona State University, School of Life Sciences, Tempe AZ 85287-4501, jstrom@asu.edu; 2

Appalachian State University, Dept. of Geography and Planning, Boone NC 28608.

Riparian vegetation and disappearing groundwater. Over the past century in the southwestern United States, rapid economic growth, expanding urban centers and agriculture have driven steep increases in freshwater demands, which have been met through groundwater pumping,

surface flow diversions, and dams - all of which alter water availability and flow patterns in rivers. These shifting baseline conditions, combined

with episodic drought, have led both to the drying of river reaches that were once perennial as well as to the wetting of previously dry river segments. Drawing from case studies of rivers in southern Arizona including Cienega Creek, Garden Canyon, the San Pedro River, and the Santa

Cruz River, we explore several scenarios: 1) the biological changes that occur in groundwater-dependent ecosystems as water tables deepen

and no-flow periods increase, 2) the patterns and rates of vegetation change that occur when diversion and pumping pressures are reduced; and

3) the response of riparian vegetation to discharge of municipal effluent into stream channels.

Vegetación ribereña y desaparición de aguas subterráneas. Desde el siglo pasado en el suroeste de Estados Unidos, el rápido crecimiento

económico, la expansión de los centros urbanos y la agricultura han aumentado la demanda de agua dulce, que se ha resuelto a través de bombeo

del acuífero, desviaciones de corrientes en la superficie y presas, todo lo cual altera la disponibilidad de agua y los patrones de las corrientes de

ríos. El cambio de estas condiciones básicas, combinado con la sequía, han llevado tanto al secado de tramos de ríos que antes eran perennes,

y también ha causado que tramos anteriormente secos tengan agua. A partir de estudios de caso de ríos en el sur de Arizona, incluyendo los

ríos Ciénega Creek, Garden Canyon, San Pedro, y Santa Cruz, exploramos varios escenarios: 1) los cambios biológicos que ocurren en los

ecosistemas que dependen de aguas subterráneas, conforme baja el nivel freático y aumentan los períodos sin corriente, 2) los patrones e índices

de cambio de vegetación que ocurren cuando se reduce el estrés por la desviación y el bombeo, y 3) la respuesta de la vegetación ribereña a la

descarga de aguas residuales en los cauces de arroyos..

The content of these abstracts reflect the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 19

Santa Cruz River Conservation
20 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 21

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Conservation Efforts and Possibilities for Increased 

Collaboration in the Santa Cruz River Watershed 

Claire A. Zugmeyer and Emily M. Brott

Sonoran Institute, Tucson, Arizona

Abstract—Attendees of the annual Santa Cruz River Researchers’ Day meetings have identified a need to 

expand collaboration, partnership, and sharing of lessons learned across the watershed. To help guide this 

interest, Sonoran Institute organized a symposium on 2 May 2012 entitled “Santa Cruz River Conservation.” 

The symposium had simultaneous Spanish/English translation and featured (1) presentations of work being 

done within the watershed in Mexico, (2) a binational mapping session to inventory current knowledge of 

conservation efforts in the watershed, and (3) a discussion and invitation to consider forming an interim 

watershed-wide steering committee. Featured themes included: the importance of strong community engagement and relationship building for conservation success, highly valued areas are often located in highly 

threatened areas, and increased watershed-wide collaboration could provide numerous benefits and help 

ensure greater collective impact. Participants were actively engaged and many will volunteer to form an 

Interim Santa Cruz River Steering Committee to begin brainstorming if and how to form a larger watershedwide collaboration.

Introduction

	 Rivers andriparianareasinsemi-aridregionssupport anabundance

of life. The Santa Cruz River of southeastern Arizona and northwest

Mexico supports tremendous diversity with nearly 600 species of

plants and animals documented from a small stretch of the river at

the Tumacácori National Historical Park (Powell and others 2005).

The river has also supported human communities for thousands of

years with evidence of human settlement dating to 9500-9000 B.C.

(Logan 2002). In the past 60 years however, increasing demands for

freshwater have led to significant groundwater pumping, dropping

of water tables, and drying of many reaches (Glennon 2002), thus

threatening the river’s ability to sustain abundant biodiversity and

provide ecosystem services that benefit human communities.

	 Numerous conservation efforts are underway throughout the Santa

Cruz Watershed to promote watershed health and protect and restore

these important riparian areas. In order to share results of research

and lessons learned, an annual Santa Cruz River Researchers’ Day

was initiated in 2009. While initially focused solely on research and

monitoring in the Upper Santa Cruz River, in Santa Cruz County,

Arizona, partner interest in the event has grown and attendees have

identified thedesire toexpandcollaboration, partnerships, andsharing

of lessons learned throughout the watershed, both binationally and

across subwatersheds.

	 Tohelpguide thisgrowinginterestinwatershed-wide collaboration,

the Sonoran Institute organized a featured symposium on Santa Cruz

River Conservation on May 2, 2012. This symposium consisted of

three sessionsfeaturing (1) presentations of projects conducted in the

Mexico portion of the watershed, (2) an inventory of conservation

efforts throughout the watershed, and (3) a discussion and invitation

to consider forming a watershed-wide steering committee. To reduce

language barriers, simultaneous translation of English and Spanish

was available throughout the day.

Santa Cruz River Through Mexico

	 Most Mexican partners have been unable to attend the annual

Santa Cruz River Researchers’ Day due to expense and challenges

associated with the long distance travel. To begin increasing binational collaborations and sharing oflessonslearned, Sonoran Institute

invited researchers, managers, and individuals working on diverse

conservation efforts within theMexican part ofthe watershed to share

their work during the first session of the symposium. Presentations

included monitoring and restoration within the rural areas along the

main stem of the river and effortsto improve watershed health within

the city of Nogales, Sonora.

Rural Monitoring and Restoration 

	 Upon leaving the headwaters in the San Rafael Valley in Arizona,

the Santa Cruz River flows 25 miles (40.2 km) through several small

ejidosorruralcommunities(Arizona-SonoraDesertMuseum2003).In

addition to being the primarywaterresource forthese smalltowns,the

Santa Cruz River comprises a dominant portion of the water portfolio

for the cities of Nogales, Sonora, and Nogales, Arizona, or “Ambos

Nogales” (Sprouse 2003).Thusthe riverisrecognized as an important

resource, while simultaneously facing numerous concerns including
22 USDA Forest Service Proceedings RMRS-P-67. 2013

Zugmeyer and Brott Conservation Efforts and Possibilities for Increased Collaboration in the Santa Cruz River Watershed 

overgrazingofrangelands,increasedgroundwaterpumping,recurrent

drought, and mining, among others(Solis and others 2012). To better

understand changing river conditions, the University of Sonora led

severalstudies over 15 yearsthat examined surface and groundwater

quality,surface water flows, and riparian vegetation (Solis and others

2012).

	 Water samples from the river and from wells that supply drinking

water were periodically tested between 1996 and 2009 for many

parameters including nutrients, agrochemicals, heavy metals, and

microbial indicators (Solis and others 2012, 2011). Though many

parameters met water quality standards, total coliforms, fecal coliforms, and E. coli were present in both surface and groundwater,

with levels in the river exceeding the standards for many uses such

as drinking water, recreation, and agriculture (Solis and others 2012).

Microbial contamination of wells may be responsible for diarrhea in

children, which represented 6.3% of illnesses along the Santa Cruz

River in San Lázaro, Sonora (Solis and others 2012). Composition

and abundance of riparian vegetation along the river were studied in

2000 (Solis and others 2012, 2002). The study documented 335 plant

native species and 38 non-native species, with Cottonwood (Populus 

fremontii) and Gooddings’willow (Salix gooddingii) asthe dominant

species.

	 In addition to research, environmental education programs taught

local students about water quality monitoring among other topics

(Solis and others 2012). More funding is needed to continue these

workshops, aswell asto continuemonitoring efforts along thisstretch

ofthe river.Continuedsharingoflessonslearnedwillincrease research

and conservation success.

	 Restoration efforts near San Lázaro, in the southern most portion

of the Santa Cruz River watershed, have consisted ofslowly building

collaborations with local community members that eventually led to

the installation of over 200 simple rock gabion structures in the mid2000s (Murrieta and Hare 2012). These gabions slow water, build up

soil, and create a “sponge” that will capture and slowly release the

run-off, helping to restore degraded grazing lands(Murrieta and Hare

2012). Community involvement and celebration of all achievements,

even the small ones, was emphasized as a critical element to restoration success. Sky Island Alliance is leading the preliminary planning

phases of a new restoration project that will involve a collaborative

effort with 4 to 5 local ranchers. The project would install fencing

around riparian areas and establish pole plantings of cottonwood and

willow to help mitigate erosion (Murrieta and Hare 2012). The fencing would exclude cattle and cross the river in some areas, requiring

repair following large flood events.

Urban Watershed Health Improvements

	 Equally important for river health is managing and mitigating

the quality of stormwater runoff generated in urban areas within the

watershed.The impermeable surfaces associatedwith roads and other

surfaces in urban areas quickly shed flows downstream and reduce

infiltration of rainfall into the surrounding soil (Shipek and others

2012). Surface runoff in urban cities often carries pollutants that

then flow directly into local rivers, impacting water quality. Rainwater harvesting and installation of other green infrastructure can

reduce contamination,improve stormwater quality, and increase local

infiltration (Shipek and others 2012). Local demonstration projects,

completed with the community,simultaneously promote community

awareness, train local leaders, and improve local stormwater quality.

	 Watershed Management Group hasled the development ofseveral

demonstration sites in the Santa Cruz Watershed, including one at

Instituto Tecnolόgico de Nogales (Shipek and others 2012). The

Instituto Tecnolόgico de Nogales campus is large and composed of

many steep surfacesthat quickly drain stormwater into major washes

that eventually lead to the Santa Cruz River. Active and passive

rainwater harvesting systems were installed to better utilize rainfall,

while reducing and slowing stormwater runoff (Shipek and others

2012). Rainwater from the roof of a campus building was actively

directed into a large cistern for landscape irrigation. Earthworks and

bermsslowed stormwater runoff allowing water to infiltrate and contaminants to be filtered out locally rather than flowing downstream.

Together these rainwater harvesting features can capture 2.75 inches

(6.99 cm) of rainfall in a 100-year storm (Shipek and others 2012).

Community involvement is vital with these projects. The bulk of the

volunteers participating in the construction were from local school

groups, mainly students from the Instituto Tecnolόgico and students

from schools involved in a grassroots effort called Asociación de

Reforestación de Ambos Nogales (ARAN). In addition, partnership

withComisiόnEstatal deAgua, the state water agency, involves other

levels of the community. Next steps for this project include development of evaluation and monitoring protocols as well as community

workshops on maintenance of rainwater harvesting features.

	 Inadequate wastewater infrastructure can also lead to contamination ofstormwater runoff. U.S. and Mexican economic policies have

caused industrialization and rapid urban growth along the border,

often with insufficient time and resources to build adequate public

infrastructure (Lipnick and others 2012). Latrines and septic systems

canoverflowduringflood events anddirectlycontaminatewashes and

rivers through stormwater runoff, as well as cause health problems

on both sides of the border. Limited water resources and insufficient

wastewater infrastructure suggest the use of household composting

toilets in Nogales, Sonora (Lipnick and others 2012). Composting

toilets use little or no water to treat human waste and provide benefits

including pollution control, production of compost, and reduced

spending on water.

	 Building on a 2002 effort in Colonia Bella Vista, a collaborative

multiphase project began in 2008 to install composting toilets within

Colonias del Sol in Nogales, Sonora. University ofArizona’s Bureau

of Applied Research in Anthropology (BARA), has led this current

effort with partnership from municipal officials and other community

groups(Lipnickandothers2012).Communityoutreachandeducation

is essential to the success of the project. Evaluations of the initial

phase of toilet construction found that toilets were used properly

(frequent use of toilet, sufficient sawdust) when people received

outreach; those receiving inadequate outreach had toilets that were

poorly maintained (e.g. smelly, not enough sawdust; Lipnick and

others 2012). Community perception of composting toilets varies.

Somemembers viewcomposting toilets positively,while othersthink

they are unpleasant. Several community members will connect to

wastewaterinfrastructure and convert to flush toiletswhen that option

is available, while others enjoy their composting toilet and have the

secondary benefit of compost for their plants. Ideally, with continued

outreach and education, a greater percentage of the community will

choose composting toilets(Lipnick and others 2012).Currentdemand

for composting toilets exceeds project capacity, with few resourcesto

build new toilets and continue the monitoring of toilets from earlier

project phases.

Community Participation Brings Success

	 A common theme identified, whether working in rural ranching

communities or in densely populated urban settings of Mexico, was

the importance of collaboration with local communities. Establishing

relationships with community members by taking time to gauge their
USDA Forest Service Proceedings RMRS-P-67. 2013 23

Conservation Efforts and Possibilities for Increased Collaboration in the Santa Cruz River Watershed Zugmeyer and Brott

interest, identify local problems, and develop collaborative solutions

generated greater project success; however, finding adequate time

and resources to build relationships is often a challenge. While

achievinggreater‘buy-in,’more importantlycommunityinvolvement

helps develop local skills and leadership that will be vital for long

term success. Though a synthesis discussion primarily focused on

the Mexican portion of the watershed, participants agreed that these

ideas would apply within the U.S. portion of the watershed.

Binational Mapping

	 Mapping and inventory of current conservation effortsis an important tool and firststep to help increase collaboration both binationally

andacrosssubwatersheds.Thebinationalmappingsessionhadseveral

goals: 1) review draft maps of known conservation efforts, and 2)

map and describe other conservation efforts, high value areas, and

threatened areas. The mapping exercise focused on the upper half

of the Santa Cruz River watershed and was further subdivided into

four regions (fig. 1). The draft maps highlighted areas in the United

States known to be either permanently protected for conservation or

managed for conservation (e.g.Arizona State Trust Land that is managed for conservation use). As land ownership differs in Mexico, the

existence of permanently protected areasis uncertain and could not be

mapped. The draft maps also included general polygons identifying

monitoring and restoration efforts. Each polygon was marked with a

numberthat corresponded to a general description ofthe conservation

effort and the lead organizations involved. Participants were asked

to comment on existing polygons and descriptions, as well as draw

additional polygons associatedwith other conservation efforts. While

primarily monitoring and restoration projects were identified in the

draft map, the inventory of conservation efforts included the following: land protection, monitoring, research, restoration, coalition or

partnership, or some other effort.

	 The 90-minute mapping session was an effective starting point to

what will likely be a larger mapping effort in the future. There was

good representation from all foursubregions and numerous additions

were made to the maps. Results of the mapping will be reviewed and

eventually made available, with the idea that the datasets will be an

active resource that is periodically updated.

	 Initial review of the mapping datasheets highlighted common

themes among the different regions. High value areas included one

or more of the following: perennial water, important ecosystems(i.e.

grasslands, riparian), good wildlife habitat, and undeveloped areas

or protection from development. Threatened areas often overlapped

with areas of high value if there were known activities that may

impact the high value area, such as loss of perennial flow to ground

water pumping or mining activities. Other threats common to two or

more of the sub-regional maps included mining, grazing, non-native

species, and development.

Action Planning

The lastsessionofthedaywas a facilitateddiscussionaboutincreasingbinational andsubwatershedcollaborationandthegrowingideaof

forming a larger watershed entity for this purpose. Participants were

asked to identify 1) the benefits and values associated with increased

collaboration at the watershed level, and 2) the initial actions or first

steps that would need to be taken.

	 Benefits to increased collaboration were numerous including:

diversity of partnerships and perspectives; sharing of resources and

lessonslearned; leveraging ofskills and expertise; greater knowledge

of funding opportunities; continuity of conservation efforts; and

greater efficiency of work so that duplication is avoided and projects

are coordinated. In addition, a watershed-wide collaboration could

provide greater recognition and visibility than single projects could

on their own. A watershed-wide collaboration comprised of diverse

groups and perspectives would demonstrate community buy-in for

conservation efforts,which is vital to the region’slong termconservation success.

	 Energy was high with the interest clear and excitement palpable

during the brainstorm session of possible actions for a watershedwide collaboration to complete. The mission of a watershed-wide

collaboration would need to be clearly defined and would likely be a

first step should such an endeavor be undertaken. Defining the structure would also be important, as increased collaboration could range

from a general alliance to a formal group.Awatershed-wide strategic

evaluation could examine long-term watershed health given current

threats and issues faced, as well as identify the long-term stakeholders—the ‘hubs’ and ‘nodes’of conservation effortsthat will maintain

the momentum. A strategic evaluation could additionally identify

priorities, data gaps, and opportunities. Goals and benchmarks of

success could be developed in order to measure long-term progress

and collective success in the watershed. Other actions could include

an annual celebration that could be based on existing events, not

necessarily new ones. Community involvement is very important;

utilizing existing events would likely be more successful than creating new ones.

	 Interest in a watershed-wide collaboration primarily stems from

discussions at the annual SantaCruzRiverResearchers’Day. Though

initiallyfocusedontheUpperSantaCruzRiver,thiseventhasexpanded

Figure 1—Orientation map depicting the locations of the four subregional maps (A-D) that were used to inventory conservation efforts 

within the upper half (hatched area) of the Santa Cruz River Watershed 

(grey area), Arizona and Sonora (Santa Cruz River = solid line, major 

tributaries= dashed line).
24 USDA Forest Service Proceedings RMRS-P-67. 2013

Zugmeyer and Brott Conservation Efforts and Possibilities for Increased Collaboration in the Santa Cruz River Watershed 

to include other regions and could perhaps contribute to a watershedwide collaboration. Participants were asked if they were interested in

forming an Interim Santa Cruz River Steering Committee to continue

discussions of how to increase collaborations and brainstorm if and

howawatershed-wide collaborationmightfunction.Sixteenattendees

responded that they may be interested in participating in an interim

committee; an additional 19 attendees ofthis year’sResearchers’Day

also expressed interest. Initial follow-up meetings and the formation

of the interim committee will occur sometime in the summer or fall

of 2012.

Conclusion

	 Organizing a symposiumon SantaCruzRiverConservation within

the MadreanArchipelago III conference provided numerous benefits

including (1) elevating the importance of the conservation efforts

in one of the Archipelago’s binational watersheds, (2) providing an

opportunity to highlight and discuss Santa Cruz River issues with

an audience that may not participate in the annual Researchers’ Day

meeting, and (3)facilitatingmany logistical detailssuch assimultaneoustranslation, venue,food, and audio-visual equipment. Therewere

over 37 attendees of the symposium throughout the three sessions.

More than 60% of attendees, including five from Mexico, did not

attend this year’s annual meeting on 29 March 2012. In evaluations

completed at the end of the day, participants listed the collaboration

discussions and opportunities to network and meet others working

throughoutthewatershed as an important highlight ofthe symposium.

Hearing presentations of work in Mexico was a big interest and much

appreciated.

	 In preparation for this event, cost of attendance (single day registration $110 U.S. or $1,517 Mexican peso) was often given as a

reason for not being able to participate. While there were numerous

benefits to having the symposium within the Madrean Conference,

future meetings will need to be free or much reduced in cost to

maximize participation. Additionally, while travel stipends helped

defray travel costs of some participants coming from Mexico, last

minute schedule conflicts and presumably the cost of travel time

amidst busy schedules, prevented several Mexican partners from

attending. Fortunately, respective U.S. partners were able to present

these binational conservation efforts, but participation by all partners

is desired. Facilitating binational participation will continue to be a

challenge to increasing watershed level collaboration. As presence

of Mexican partners is still highly desired for future events, hosting

meetings that alternate between the U.S. and Mexican venues, are

closer to the border, or use video conference technology (i.e. Skype,

GoToMeeting, orGoogle+)may need to be considered—suggestions

that were also made during the final wrap-up session of the entire

conference.

	 The Santa Cruz River Conservation symposium provided an initial inventory of conservation efforts and numerous ideas of how to

increasecollaborativeefforts.Mostimportantly,thesymposiumhelped

build new and existing relationships that will advance the collective

impact of conservation efforts watershed-wide.

Acknowledgments

	 We thank Carola A. Hirsch and Ivette Marian Arnold for simultaneous translations during the symposium; Tahnee Roberston for

facilitating our session; and Evan Canfield, Carrie Presnall, Jason

Welborn, and ScottWilborfor volunteering to compile notes and lead

group discussions. We thank John L. Koprowski andAmy L. McCoy

for reviews of the manuscript.

References

Arizona-Sonora Desert Museum. 2003. The Santa Cruz River. Sonorensis

23(1):1-29.

Glennon, Robert. 2002. Water follies: Groundwater pumping and the fate of

America’s fresh waters. Washington, DC: Island Press. 314 p.

Lipnick, Carolyn; Robles, Pedro; Trujillo, Francisco. 2012. Composting toilets in the Santa Cruz watershed: Reducing contamination and improving

soils. Featured symposium presentation at the Madrean Archipelago III

Conference; 2 May 2012.

Logan, Michael F. 2002. The lessening stream: An environmental history of

the Santa Cruz River. Tucson: University of Arizona Press. 311 p.

Murrieta,Joaquin; Hare,Trevor.Riverrestorationandcommunityengagement

in San Lázaro, Sonora. Featured symposium presentation at the Madrean

Archipelago III Conference; 2 May 2012.

Powell, Brian F.; Albrecht, E. W.; Halvorson, W. L.; [and others]. 2005. Vascular plant and vertebrate inventory of Tumacáori National Historic Park.

USGS Open-File Report 2005-1142. U. S. Geological Survey, Southwest

Biological Science Center, Sonoran Desert Research Station, University

of Arizona. Tucson, AZ. 140 pp.

Shipek,Catlow;Garibaldi,JuanCarlos;Flores,Francisco.Stormwaterharvesting at the Instituto Tecnolόgico de Nogales, Sonora. Featured symposium

presentation at the Madrean Archipelago III Conference; 2 May 2012.

Solis Garza, Gilberto; Lόpez Estudillo, R.; Briggs, M.; Jenkins, P. 2002.

Composiciόn y abundancia de la vegetaciόn del Río Santa Cruz, Sonora,

Mexico. Biotecnia 3(3):74-79.

Solis Garza, Gilberto; Arturo Isreal Villalba Atondo; Gerardina Nubes Ortíz;

José María del CastilloAlarcόn; FlorAdriana MerazAcosta. 2011. Físicoquímica del agua superficial y sediment en el Río Santa Cruz, Sonora,

Mexico. Biotecnia 13(1):3-9. Online: http://www.biotecnia.uson.mx/revistas/articulos/9-ART%20FISICO%20QUIMICA%20DEL%20AGUA.

pdf [May 9, 2012]

Solis-Garza, Gilberto; Agustin Gomez-Alvarez; Arturo Villalba-Atondo.

Monitoring and conservation efforts along the Santa Cruz River in Sonora,

Mexico. Featured symposium presentation at the MadreanArchipelago III

Conference; 2 May 2012.

Sprouse,TerryW. 2003.Equitablemanagement of Mexican effluent inAmbos

Nogales. Journal of the Southwest 45(4):595-610.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 25

Changing Climate in the Sky Island Region
26 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 27

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

A Conceptual Model of Plant Responses to Climate with 

Implications for Monitoring Ecosystem Change

C. David Bertelsen

Herbarium and the School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—Climate change is affecting natural systems on a global scale and is particularly rapid in the 

Southwest. It is important to identify impacts of a changing climate before ecosystems become unstable. 

Recognizing plant responses to climate change requires knowledge of both species present and plant 

responses to variable climatic conditions. A conceptual model derived from observations made during a 

28-year phenological study is presented and implications for monitoring ecosystem change are discussed.

Introduction

	 Climate change impacts are affecting natural systems world-wide

(McCarty 2001; Parmesan and Yohe 2003). The rate of climate

change in the American Southwest is more rapid than elsewhere on

the continent, with the possible exception of the Arctic (Overpeck

and Udall 2010). Increasing temperatures and decreasing precipitation in the Southwest are predicted by most climate models (Karl

and others 2009; Weiss and Overpeck 2005; Sheppard and others

2002). Species currently present have had thousands of yearsto adapt

to climate variability (Van Devender 1995, 2000), but the margin

of continued success will become increasingly smaller as climatic

change intensifies, particularly in non-mountainous biomes such as

deserts and grasslands (Loarie and others 2009; Munson and others

2012). Resource managers are increasingly focusing on climate

change in planning and management activities (USDI 2008; Heller

and Zavaleta 2009; Mawdsley and others 2009).Asthe rate of change

accelerates, it becomes important to identify when an ecosystem

becomes unstable because mitigation efforts become more difficult

and expensive as ecosystems near the point of collapse (CCSP2009).

Adequate monitoring is essential not only to detect changes but to

identify appropriate adaptive management actions and to measure

their effectiveness (Hobbs 2009; West and others 2009).

Plant Responses to Climate Change

	 Plants respond to climate change by moving, adapting, or dying

(Peterson and others 2005).Movement ismore likely than adaptation,

but human uses may limit movement by reduction and fragmentation

of habitat, as well as reduction in population sizes (Parmesan and

others 2000). Climate variability per se may also limit movement

(Early and Sax 2011). Adaptation is usually limited to the existing

genetic variability within a species; genetic shifts may mitigate local

climate effects,butitisunlikelysuch shiftswillmitigate impacts atthe

species level (Parmesan 2006; Huntley 2005; Thomas 2005). Dying

or extinction is likely for species with low capacity for adaptation or

dispersal (Walther and others 2002).

	 Expected responses of plants to climate change include (1) phenological changes, e.g., earlier onset of flowering,shortening orlengthening of growing seasons; (2) geographic range changes, e.g., range

shifts, extensions, and contractions to higher elevations or latitudes;

(3) population and reproductive biology changes, e.g., changes in

abundance, reproductive success; and (4) community and ecosystem 

changes, e.g., changesincomposition,habitats,productivity,structure

(CCSP2009;McCarty 2001; Parmesan 2005;Root andHughes 2005;

Walther 2010). Plants respond to climate change at the species, not

community,level, andcontinuedchangewilllikelyresultinecosystem

destabilization and community shifts (Huntley 2005; Walther and

others 2002). Ecosystems disintegrate and at the same time reassemble into new configurations at the species level (Lovejoy 2005).

Predictions of what new configurations will develop are uncertain at

best (Hobbs and others 2009) because the “rules” for assemblage of

plant communities are not well understood (Gotzenberger and others

2012). Species’ interrelationships are largely unknown, and the loss

of a single species may have cascading effects through and within

trophic levels (Voigt and others 2003; Brooker 2006). Reduction in

biodiversity may lead to loss of resilience, often in the form of shifts

in dominant life forms, and this usually results in drastic transformationto an alternative and unpredictable state (Scheffer 2010).Scheffer

and others (2001:596) conclude that “...efforts to reduce the risk of

unwanted state shifts should address the gradual changes that affect

resilience rather than merely control disturbance.”

Measuring Change in a Variable 

Climate 

	 I have inventoried plant taxa in flower along a 5-mile route that

climbs4158feetthroughsixvegetative associationsintheFingerRock

Canyon drainage in the Santa Catalina Mountains of Pima County,

Arizona, since 1984 (fig. 1). The 1100-ac study area is about 0.6%

of the area of the range but includes over 40% of the known plant

taxa (Verrier, unpublished paper). During 1368 hikesin the drainage,
28 USDA Forest Service Proceedings RMRS-P-67. 2013

Bertelsen A Conceptual Model of Plant Responses to Climate with Implications for Monitoring Ecosystem Change

Figure 1—The study area, showing the route to Mount Kimball (white line) and the six vegetative associations. DS = desert scrub, RS = riparian scrub, SG = scrub grassland, OW = oak woodland, OPW 

= oak-pine woodland, and PF = pine forest. (Source: Jeff Belmat and Theresa Crimmins, based on data 

provided by the author.)

averaging once a week, I have recorded over 140,000 observations

of 601 plant taxa by the mile segment on which they were seen.

	 Perhapsthe mostsalient characteristic of the data shown in figures

2 and 3 istemporal and spatial variability that cuts acrossspecies and

plant life forms, e.g., annual forbs, herbaceous perennials, shrubs,

succulents, and trees. Speciesrespond individualistically to the same

climatic conditions. Presence or vegetative growth does not guarantee reproduction, and reproductive success can fluctuate widely.

The distribution of most species in the flora is not only different

from year to year but difficult to predict. In my experience, a basic

tenet of much ecosystem monitoring—that key areas, study plots, or

transectsrepresent the broader community overtime—assumesmore

homogeneity in composition, distribution, and frequency than exist

in natural systems. Legendre and Fortin (1989:107) state “in nature,

living beings are distributed neither uniformly nor at random. Rather,

they are aggregated in patches, or they form gradients or other kinds

of spatial structures.” No matter what the vegetative association,

microhabitats within it differ significantly from the larger area, and

a significant portion of the biodiversity of an area is found in them.

Small differencesin temperature or precipitation, or changesin aspect

and slope, may change soil moisture and evapotranspiration, and

therefore favor different species.

	 All of the specific responses to climate change listed above have

occurredinmystudyarea (Crimmins andothers2008,2009).Although

my data do not span a period of time sufficient to attribute these

changes to climate change, the study area has experienced a severe

drought accompanied by abnormally high temperatures since 1999

(Woodhouse and others 2010; Overpeck and Udall 2010) and plant

responsesto extreme climate events are likely indicative ofresponses

to climate change (Parmesan and others 2000) . In 2002, I began to

notice significant mortality of dominant species in several vegetative associations, namely saguaros, white oaks, alligator junipers,

and ponderosa pines. I began to look for other changes and quickly

learned that a major indicator of change was what species were no

longer present. Significant change, which I found is a combination

of many changes at the species level, is not easy to see until a certain

threshold is reached, even if expected species are well known and an

area is visited regularly.

	 To recognize that plants are responding to climate change and to

measure those responses,validreferencepointswithwhichtocompare

current conditions are prerequisite. First, knowledge of the area and

local floras is essential (USDI 2009; USFWS 2010). Although not a

substitute for comprehensive speciesinventories, historical data may

be useful if interpreted in light of climate (Joyce and others 2008).

Accurate mapping of vegetative associations can be useful in establishing parameters but without knowing what is or recently has been

present, it is not possible to determine when or whatspeciesincrease,

decline, or disappear, and these are the species most likely to be the

“early responders” to climate change.Adecline in biodiversity is one

ofthemost likely consequences of climate change (Bellard and others

2012). Biodiversity is a major component of ecosystem resiliency

and, as Maestre and others (2012:214) state, “is crucial to buffer

negative effects of climate change and desertification in drylands.”

Resiliencemaybereducedbygradualanddifficult-to-detectchangesin
USDA Forest Service Proceedings RMRS-P-67. 2013 29

A Conceptual Model of Plant Responses to Climate with Implications for Monitoring Ecosystem Change Bertelsen

environmental conditions to the point that natural fluctuations result

incatastrophic ecosystemshifts,typicallywithout earlywarningsigns

(Scheffer 2010). Early identification of responses to climate change

must be made at the species level where such responses first appear.

Focus at thislevelmay also facilitate better understanding of plant-toplant interactions(e.g., competition, facilitation, and adaptation) that

can “mediate the impacts of environmental change” (Brooker 2006).

	 Second, a measure of “natural” or “expected” climate variability

is required. If the response of individual species or groups of species

to various climate change scenarios is to be determined, knowledge

of how they respond to climate variability at the species level is

needed (Parmesan 2005; Peterson and others 2005). Precipitation and

temperature are perhaps the most important abiotic drivers of plant

reproduction(Crimmins andothers2008,2010,2011).Nomatterwhat

climate change brings, climatewill undoubtedly continue to be highly

variable (fig. 4). The impacts of climate change can be difficult to

discern even when looking at long-term data because there isso much

year-to-year variability (Parmesan 2005).Assessments of abundance

or frequency in particular are meaningless without knowledge of the

Figure 2—Number of species in flower by year and mile in the Finger Rock Canyon drainage, 1984-

2010. Elevations are as follows: Mile 1 = 3100-3540 ft; Mile 2 = 3540-4500 ft; Mile 3 = 4500-5480 ft; 

Mile 4 = 5480-6360 ft; Mile 5 = 6360-7258 ft. (Data is incomplete for 2004-2005.)

Figure 3—Number of annual forbs and herbaceous perennials in flower by month and year in the Finger Rock Canyon drainage, 1984-2010. 

(Data is incomplete for 2004-2005.)
30 USDA Forest Service Proceedings RMRS-P-67. 2013

Bertelsen A Conceptual Model of Plant Responses to Climate with Implications for Monitoring Ecosystem Change

climatic context in which assessments are made and knowledge of

expected species variability.

Conceptual Model

	 A conceptual model of plant responses to climatic conditions is

shown in figure 5. It is simplistic in that it does not consider biotic

factorssuch as genetic variability or plant-to-plant interactions. Each

ofthe foursolid circles(A-D)represents an assemblage ofspeciesthat

include species represented by the circles within them. The number

of species in each assemblage will vary not only with vegetative associations but also over time. Different assemblages can be expected

for any given year, season, or even month. The relative size of these

assemblages, however, isthought to be consistent across association.

Most biodiversity is found in the visible portions of the three largest

circles, labeled a, b, and c.

	 Circle A represents the total flora of a vegetative association.

Climate variability and climate change may result in highly variable

biodiversity as species respond at variable rates (Wather and others

2002). The visible portion of the circle, a, represents species infrequently seen for a variety of reasons, e.g., species requiring highly

specific climatic conditions and species at the extreme periphery of

their ranges. Peripheral populations are important because they may

include distinct traits that facilitate adaptation to climate change

(Angert and others 2011; Lesica and Allendorf 1995). In the Finger

Rock Canyon drainage, 4% of the taxa have been seen flowering only

1 year and 15% 5 or fewer years.

	 Circle D includes species that flower every year, i.e., those most

well adapted to the local climate regime. These species comprise

less than 25% of the total flora in my study area. They are generally

common and are frequently the primary focus of monitoring efforts,

in my experience. Common species may be the easiest to monitor,

but they are common because they are well-adapted to local climatic

conditions (Cole 2010). Thus they are least likely to indicate adverse

effects of climate change before the system nears or reaches a tipping

point.

	 Climatic variability,specifically the timing and magnitude of temperature and precipitation events, is most apparent in the responses

of species represented by circles B and C. The species in these assemblages vary from year to year but are fairly consistent. Circle B

includes species that usually reproduce during climatically optimal

conditions, as much as 85% of total species. The visible area of the

circle, b, represents the most climate-sensitive species. With climate

change, it seems highly likely that the first indications of system instability will be seen here and that the most rapid change will occur

in this group. Circle C, not more than 60% of the total flora, includes

species that are usually reproductive (although populations may be

small) during adverse conditions and have a high tolerance to climate

variability.

Figure 5—Conceptural Model of Plant Responses to Climate. A = total 

species in a vegetative association or biotic community. a = seldom seen 

species. B = species normally reproductive under optimal conditions. b 

= highly climate sensitive species. C = species normally reproductive 

under adverse conditions. c = species adapted to adverse conditions. 

D = species reproductive every year (species most well-adapted to local climatic variability). E = species that should be monitored to detect 

early ecosystem changes.

Figure 4—Monthly precipitation (mm) and average monthly temperature (°C) for the Finger Rock Canyon drainage, 1984-2010. (Source: Michael 

Crimmins based on PRISM [Parameter-elevation Regressions on Independent Slopes Model] climate mapping system data.)
USDA Forest Service Proceedings RMRS-P-67. 2013 31

A Conceptual Model of Plant Responses to Climate with Implications for Monitoring Ecosystem Change Bertelsen

Monitoring for Climate Change

	 CircleErepresentsspecies on which ecosystemmonitoring should

focus in order to detect instability before it crosses a threshold of no

return. Over time, monitoring of a wide range of species can lead not

only to a far better understanding of the species in the ecosystem and

expected climate variability within species but also to identification

of changes occurring in the system in response to climate change.

The conceptual model has a number of implications for ecosystem

monitoring as it pertains to climate change:

	 1. Comprehensive flora inventories need to be developed to determine which species need protection and to provide a baseline against

which change can be measured. These inventories can be developed

during the process of thorough, on-going monitoring.

	 2. The goal should be to monitor all species. It is not possible to

predict accurately which of the species will be most affected by a

changing climate because of their individual responses. Monitoring

of life forms, which draw moisture from different soil depths is also

essential because changes in life form composition can indicate

structural instability (House and others 2003; McCluney and others

2011; Schenk and Jackson 2002).

	 3. Monitoring of ecosystem change needsto occur more than once

a year. Different species grow, reproduce, and die at different times

of the year. Species flowering in spring may respond to different climatic factors than those flowering in summer (Crimmins and others

2010, 2011). At any point in time a different mix of species will be

present. Growth in spring, both above and below the ground, may be

important to the reproductive success ofspecies flowering in summer.

At a minimum, monitoring should be completed in both spring and

summer, preferably in the peak of the growing season. Some climate

change models predict a sharp decline in winter precipitation (Karl

and others 2009), and this has major ramifications for spring growth

and reproduction in arid and semi-arid biomes (Crimmins and others

2008; 2010).

	 4. More extensive and intensive monitoring during climatically

optimal and adverse years can lead to identification of climate sensitive species and better understanding of climate variability at the

species level. Past and present monitoring data should be interpreted

in terms of “normal,” “optimal,” and “adverse” conditions. We need

to look for change and trends.

	 5. Microhabitats including topographical characteristics such as

slope and aspect should be represented in areas monitored because

of the biodiversity found in such areas.

	 Perhaps the greatest limitation on current monitoring practices is

inadequate resources—including time, personnel, and knowledge—

and a lack of understanding, at the funding level, of the importance of

monitoring.Itwould be idealto followCraig’s(2010)first principle to

“monitor and study everything all the time,” but thisis hardly possible

in the “real world” of declining budgets. Even if significantly more

funding were available, difficult choices will have to be made as to

where and when adequate monitoring is to be done. With a focus on

specieswith known responsesto climatic factors, existingmonitoring

protocols may be useful in detecting long-term change, but a new

paradigm is needed if we are to see change before ecosystems reach

the point of collapse. There are no easy, simple, or inexpensive answerstothemonitoringconundrumbecause ecosystems are extremely

complex and in constant flux.

References

Angert AL, Crozier LG, Rissler LJ, [and others]. (2011). Do species’ traits

predictrecentshifts at expanding range edges?EcologyLetters14:677-689.

Bellard C, Bertelsmeier C, Leadley P, Thuiller W, Courchamp F [and others].

(2012). Impacts of climate change on the future of biodiversity. Ecology

Letters 15:365-377.

BrookerRW.(2006). Plant-plant interactions and environmental change. New

Phytologist 171:271-284.

CCSP. (2009). Thresholds of climate change in ecosystems. A report by the

U.S.ClimateChangeScienceProgramandSubcommitteeonGlobalChange

Research. U.S. Geological Survey, Reston, VA.

Cole KL.(2009).Vegetation response to early Holocene warming as an analog

for current and future changes. Conservation Biology 24:29-37.

Craig RK. (2010). “Stationarity is Dead”—Long live transformation: Five

principles for climate change adaptation law. Harvard Environmental Law

Review 34: 9-73.

Crimmins T, Crimmins MA, Bertelsen D, BalmatJ. (2008). Relationships betweenalphadiversityof plantspeciesinbloomand climaticvariables across

an elevationgradient.InternationalJournalofBiometeorology52:353-366.

Crimmins T, Crimmins MA, Bertelsen D. (2009). Flowering range changes

across an elevation gradient in response to warming summer temperatures.

Global Change Biology 15:1141-1152.

Crimmins T, Crimmins MA, Bertelsen CD. (2010). Complex responses to

climate drivers in onset of spring flowering across a semi-arid elevation

gradient. Journal of Ecology 98:1042-1051.

CrimminsT,CrimminsMA,BertelsenD.(2011).Onsetofsummerfloweringin

a“SkyIsland”isdrivenbymonsoonmoisture.NewPhytologist191:468-479.

Crowley TJ. (2000). Causes of climate change over the past 1000 years.

Science 289: 270-277.

EarlyR,SaxDF.(2011).Analysis of climate pathsreveals potentiallimitations

on species range shifts. Ecology Letters 14: 1125-1133.

Gotzenberger L, de Bello F, Brathen KA, [and others]. (2012). Ecological

assembly rulesin plant communities—approaches, patterns, and prospects.

Biological Reviews 87:111-127.

Heller NE, Zavaleta ES. (2009). Biodiversity management in the face of

climate change: a review of 22 years of recommendations. Biological

Conservation 142:14-22.

Hobbs RJ, Higgs E, Harris JA. (2009). Novel ecosystems: implications for

conservation and restoration.TrendsinEcology andEvolution 24:599-605.

House JI, Archer S, Breshears DD, Scholes RJ, [and others]. (2003). Conundrumsinmixedwoody-herbaceous plantsystems.Journal ofBiogeography

30: 1763-1777.

Huntley B. (2005). North temperate responses. In Lovejoy TE, Hannah L,

eds. Climate Change and Biodiversity. Yale University Press, New Haven.

Pp 109-124.

KarlTR,MelilloJM,PetersonTC, eds.(2009).GlobalClimateChange Impacts

in the United States. Cambridge University Press, New York.

Joyce LA, Blate GM, Littell JS, McNulty SG, et al., (2008) National forests.

In: Julius SH, West JM (eds) Preliminary review of adaptation options for

climate-sensitive ecosystems andresources.U.S.EnvironmentalProtection

Agency, Washington, DC.

Legendre P, Fortin M-J. (1989). Spatial pattern and ecological analysis.

Vegetation 80: 107-138.

Lesica P,Allendorf FW. (1995). When are peripheral populations valuable for

conservation? Conservation Biology 9:753-769.

LoarieSR,DuffyPB,HamiltonH,[andothers].(2009).Thevelocity of climate

change. Nature 462: 1052-1055.

Lovejoy TE. (2005). Conservation with a changing climate. In Lovejoy TE,

Hannah L, eds. Climate Change and Biodiversity. Yale University Press,

New Haven. Pp 325-328.

Maestre FT, Quero JL, Gotelli NJ, [and others]. (2012). Plantspeciesrichness

and ecosystem multifuntionality in global drylands. Science 335: 214-218.
32 USDA Forest Service Proceedings RMRS-P-67. 2013

Bertelsen A Conceptual Model of Plant Responses to Climate with Implications for Monitoring Ecosystem Change

Mawdsley JR, O’Malley R, Ojima DS. (2009). A review of climate-change

adaptation strategies for wildlife management and biodiversity conservation. Conservation Biology 23:1080-1089.

McCarty JP. (2001). Ecological consequences of recent climate change.

Conservation Biology 15:320-331.

McCluney KE, Belnap J, Collins SL, Gonzalez AL, [and others]. (2011).

Shifting speciesinteractionsin terrestrial dryland ecosystems under altered

water availabilityandclimate change.BiologicalReviews.Publishedonline

17 November 2011.

Munson SM, Webb RH, Belkap J, [and others]. (2012). Forecasting climate

change impacts to plant community composition in the Sonoran Desert

region. Global Change Biology 18:1083-1095.

Overpeck J, Udall B. (2010). Dry times ahead. Science 328: 1642-1643.

Parmesan C. (2006). Ecological and evolutionary responses to recent climate

change.AnnualReview of Ecology, Evolution, and Systemics 37:637-669.

Parmesan C. (2005). Biotic response: range and abundance changes. In Lovejoy TE, Hannah L, eds. Climate Change and Biodiversity. Yale University

Press, New Haven. Pp 41-55.

ParmesanC,YoheG.(2003).Aglobally coherentfingerprint of climate change

impacts across natural systems. Nature 421:37-42.

Parmesan C, Root TL, Willig MR. (2000). Impacts of extreme weather and

climate on terrestrial biota. Bulletin of the American Meteorological

Society 81:443-450.

Peterson AT, Tian H, Martinez-Meyer E, [and others]. (2005). Modeling

distributional shifts of individual species and biomes. In Lovejoy TE,

Hannah L, eds. Climate Change and Biodiversity. Yale University Press,

New Haven. Pp. 211-228.

Root TL, Hughes L. (2005). Present and future phonological changes in wild

plants and animals. In Lovejoy TE, Hannah L, eds. Climate Change and

Biodiversity. Yale University Press, New Haven. Pp. 61-69.

Scheffer M. (2010).Alternative states in ecosystems. In Terborgh J, Estes JA.

Trophic cascades: Predators, prey, and the changing dynamics of nature.

Island Press, Washington, D.C. Pp. 287-298.

Scheffer M, Carpenter S, Foley JA, [and others]. (2001). Catastrophic shifts

in ecosystems. Nature 413:591-596.

SchenkHJ,JacksonRB.(2002).Rootingdepths,lateralrootspreads andbelowground/above-ground allometries of plants in water-limited ecosystems.

Journal of Ecology 90:480-494.

Sheppard PR, Comrie AC, Packin GD, [and others]. (2002). The climate of

the US Southwest. Climate Research 21:219-238.

Thomas CD. (2005). Recent evolutionary effects of climate change. In Lovejoy TE, Hannah L, eds. Climate Change and Biodiversity. Yale University

Press, New Haven. Pp. 75-88.

USDI. (2008). An analysis of climate change impacts and options relevant

to the Department of the Interior’s managed lands and waters. Subcommittee on Land and Water Management Task Force on Climate Change.

Washington, DC: U.S. Department of the Interior.

USFWS. (2010). Strategic Plan for Inventories and Monitoring on National

Wildlife Refuges: Adapting to Environmental Change. Washington, D.C.:

U.S. Department of the Interior, Fish &Wildlife Service, NationalWildlife

Refuge System.

VanDevenderTR.(1995).Desert grassland history: changing climates, evolution, biogeography, and community dynamics. In McClaran MP and Van

Devender TR, eds. The desert grassland. Tucson, University of Arizona

Press. Pp. 68-99.

Van Devender TR. (2000). The deep history of the Sonoran Desert. In Phillips SF, Comus PW, eds. A natural history of the Sonoran Desert. Tucson,

Arizona-Sonora Desert Museum. Pp. 61-69.

Verrier J. (2008). Flora of the Santa Catalina Mountains, Pima and Pinal

Counties, Arizona. Unpublished paper dated May 17, 2008.

Voigt W, Perner J, DavisAJ, [and others]. (2003). Trophic levels are differentially sensitive to climate change. Ecology 84: 2444-2453.

Walther G-R (2010). Community and ecosystem responses to recent climate

change. Philosophical Transactions of the Royal Society B, Biological

Sciences 365: 2019-2024.

Walther G-R, Post E, Convey P, [and others]. (2002). Ecological responses

to recent climate change. Nature 416: 389-395.

Weiss JL, Overpeck JT. (2005). Is the Sonoran Desert losing its cool? Global

Change Biology 11: 2065-2077.

West JM, Julius SH, Kareiva P, [and others]. (2009). U.S. natural resources

and climate change: concepts and approaches for management adaptation.

Environmental Management 44: 1001-1021.

Woodhouse CA, Meko DM, MacDonald GM, [and others]. (2010). A 1,200-

year perspective of 21st century drought in southwestern North America.

PNAS 107:21283-21288.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 33

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Temporal Patterns in Species Flowering in Sky Islands of 

the Sonoran Desert Ecoregion

Theresa M. Crimmins

USA National Phenology Network; School of Natural Resources and the Environment, University of Arizona, 

Tucson, Arizona

Michael A. Crimmins

Department of Soil, Water and Environmental Science, University of Arizona, Tucson, Arizona

C. David Bertelsen

Herbarium and School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—Highly variable moisture conditions in the Sonoran Desert play a significant role in shaping the 

composition and phenology of plants in this water-limited region. The flowering patterns of plants of the 

Finger Rock trail, located in the Santa Catalina Mountains of southern Arizona, have been very carefully 

documented on approximately a weekly basis for nearly three decades. These observations, made along 

a gradient encompassing more than 4,000 ft in elevation, have revealed that flower presence, timing, and 

duration vary dramatically from year to year for many species in this unique environment. 

We developed a daily probability of flowering estimate for each species from the empirical records of 

presence or absence of flowering and implemented a clustering algorithm to elucidate recurrent temporal 

flowering patterns among groups of species. Several patterns emerged, and these patterns are consistent with 

plant functional types. First, across the elevation gradient, over half of all species that have been documented 

exhibit very low (<10%) probabilities of flowering in any given year, a pattern that exemplifies the high degree of variability and opportunism in this water-limited environment. In addition, there are small groups of 

species that are consistently seen in flower nearly every year and at nearly the same day of year, regardless 

of the preceding or concurrent climate conditions. The reliability with which these species in these groups 

flower, both from year to year and at the same time every year, increases with increasing elevation, and is 

also higher in summer than in the spring season. Clusters comprised mainly of annual species tended to 

show lower flowering probabilities and therefore greater variability in flowering; the opposite was generally 

true for perennial and woody species. The baseline patterns established in this analysis enhance our current 

understanding of seasonal flowering patterns in this region and serve as a critical step toward understanding 

how the system may change under future climate conditions.

Introduction

	 The Sonoran Desert is characterized by two distinct periods of

flowering, one in spring and one in summer.However, the assemblage

of species seen in flower in a particular season varies widely from

yearto year(Crimmins and others 2008). Similarly, though the coarse

patterns of distinct spring and flowering seasons are consistent from

year to year, the timing of species’ flowering onset and the length of

time individual species remain in flower can vary considerably.

	 In dry environments, precipitation is the dominant limiting factor, shaping species composition and productivity (Noy-Meir 1973;

McClaran and Van Devender 1995) and playing a critical role in cueing phenological events (Beatley 1974; Bowers and Dimmitt 1994;

Crimmins and others 2010; Peñuelas and others 2004). However, the

relationship between precipitation and plant phenology has generally

been established either at a coarse level—for entire plant communities at the seasonal scale (e.g., Crimmins and others 2008)—or for

small groups of species (e.g., Aronson and others 1992; Bowers and

Dimmitt 1994; Kemp 1983; Kimball and others 2009). Therefore,

we can make predictions regarding the flowering response in a given

season at a coarse level, such as whether a spring season is likely to

be a “good” wildflower year, characterized by abundant and diverse

wildflower populations (e.g., Bowers 2005). On the other hand,

we mostly lack the ability to make finer-scale predictions such as

whether particular species are likely to be seen in flower, when they

may initiate flowering, and what the duration of their flowering over

the course of the season might be.

	 Datasets that capture the response of an entire biological community are rare, and repeated measurements of an entire system

are even scarcer. Datasets that offer repeated measurements of an

entire community are rich and valuable resources that can enhance

our empirical understanding of species’ responses within the larger

ecosystem. In this study, we utilized a long-term record of flowering
34 USDA Forest Service Proceedings RMRS-P-67. 2013

Crimmins, Crimmins, and Bertelsen Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion

plant phenology of allspecies observed (encompassing over 600 taxa

on a nearly weekly interval), which enabled us to explore in detail

both the year-to-year patterns in flowering for individual species and

each species’inter-annual variability in flowering onset and duration.

	 We use a dataset spanning 27 years to investigate variability and

timing of flowering patterns within and spanning the distinct spring

andsummerseasons characteristicoftheSonoranDesert.Specifically,

we asked fromyearto year,(1) howconsistently do individualspecies

flower; (2) how much does flowering onset and duration vary among

species; and (3) are there patterns in flowering onset, duration, and

inter-annual variability that are relatively consistent across species?

	 In this study, we evaluated temporal flowering across a 1200-m

elevation gradient and explored the consistency of patterns across

a moisture gradient encompassing xeric desert scrub to mesic pine

forest communities. The findings of this evaluation can guide temporal sampling regimes for long-term monitoring to optimize the

likelihood of observing particular species in flower and can also be

used to identify peaksin flowering diversity. In addition, these results

provide information on resource availability and habitat condition,

which is valuable for wildlife species. Finally, this information can

also serve as an important baseline from which to begin evaluations

of how the system may change under future climate conditions.

Methods

	 The data examined in this study include 136,036 observations of

plantsin flower for 601 species made by David Bertelsen. These data

are the results of 1,243 round-trip (16 km) hikes of the route to Mt.

Kimball, and 25 partial trips of 4.8-12.1 km made over a 27-year

period (1984–2010), excluding 2004 and 2005, when the observer

was unable to complete regular hikes(4.4 ± 0.01 hikes month-1 [mean

± SE]). The 8-km hiking route to Mt. Kimball is located on the south

slope of the Santa Catalina Mountains, approximately 7 km north of

the city limits of Tucson,Arizona, USA, and followsthe Finger Rock

and Pima Canyon trails. The observer recorded species observed in

bloom within approximately 9.1 m of eitherside ofthe trail. Observations were recorded along five trail segments approximately 1 mile

(1.6 km) in length both on the ascent and descent; on the descent

attention was given to verifying records and adding any taxa not

already recorded. Trail mile 1 refers to the lowest elevation segment;

trail mile 5 refers to the highest elevation; further details regarding

the study site and data collection methods are provided in Crimmins

and others (2011). Each species was assigned to one of the following

functional types: annual/biennial forbs, herbaceous perennials, and

woody plants. Functional type groupings were selected based on

the different water use strategies employed by plants of semi-arid

environments (Burgess 1995; Ehleringer and others 1991).

	 To evaluate consistency of flowering on each calendar day from

year to year, we calculated a “probability ofspecies flowering” (PSF)

for each day in the entire record (>9,800 days) for each species. This

was done by first interpolating individualspecies flowering presence/

absence observations collected at an irregular interval to a consistent

daily matrix of 9,800 days. Changesin the flowering status(presence

= 1 or absence = 0) were linearly interpolated between consecutive

hikes with a mid-point value of 0.5. All days in the record with a

value higher than 0.5 were converted to flowering days (presence =

1)for each species.The full data matrix of daily flowering occurrence

wasthen used to calculate the empirical probability of occurrence for

each day of the year. If a species was observed in multiple trail miles,

observations specific to each mile were treated independently. The

resultant species-mile PSFs provide us with the likelihood of a species being observed in flower on each day of the year in a particular

trail mile.

	 Several potential probability of flowering curves could arise from

long time-series of phenological status observations. Examples of

these and the structure of the input data responsible for these patterns appear in fig. 1. These models represent the extreme cases; we

expect most curves would fall somewhere between one or more of

these extremes. Individual species PSF plots provide information

on several elements: the length of time, or duration, that a species

is typically seen in flower, the year-to-year consistency in the days

of year a species is seen in flower, and the probability of a species

being observed in flower in a given year. The height and width of the

peaksresult from a combination of these features. Species associated

with tall, narrow peaks (fig. 1a) are characterized by a short flowering duration, a high year-to-year consistency, and are seen in flower

nearly every year. Species that exhibit a short flowering duration and

high consistency but are seen infrequently have narrow, short peaks

(fig. 1b). Species that flower in most years for a long duration with

high consistency are represented by tall, wide peaks (fig. 1c). These

lengthy,short peaks can arise bymultiple pathways. First,speciesthat

exhibit a long flowering duration with high consistency but flower in

few years produce this pattern (fig. 1d). Similarly, short peaks that

stretch over a longer period can result from species that flower for

short durations and exhibit low consistency, that is, their flowering

periods from year to year are highly variable (fig. 1e).

	 To collapse the individual 1,588 individual species-mile PSF

curves into common flowering patterns, individual species’ curves

were clustered within each mile using a K-means algorithm using

Matlab (R2011b). The algorithm uses five iterations of the K-means

classifier and retains the clustering with the smallest sum squared

error. To assess the optimal number of clusters for the input dataset,

an ensemble of clusters ranging from 2 to 20 are generated and then

compared using the Davies-Bouldin Index (DBI). A lower DBI indicates greater within-clustersimilarity and greaterseparation between

clusters. The smallest number of clusters with the lowest DBI value

indicates the simplest efficient grouping of the input data. Seven

clusters were retained for each trail mile based on these criteria.

Results

	 For each of the five trail miles we generated seven clusters (fig. 2),

whichrepresentpatternsrepeatedacrossindividualspecies’PSFplots.

We identified 10 cluster patterns (labeled A-J), eight of which were

consistent across multiple trail miles (fig. 2). Only cluster I (mile 1)

and cluster J (mile 5) were unique to a single mile.

	 A very clear pattern is represented in cluster A, which is present

in each of the five miles. Species associated with this cluster exhibit

a very low probability of being seen in flower, and may flower over

a very broad range of dates. This cluster encompasses greater than

half of the species observed in each of the miles (table 1). The low

probabilities associated with species in this cluster result from the

comparatively small number of years observed in flower(average 5.7

to 6.9 years) (table 1). Accordingly, cluster A is comprised of many

speciesthatflowerinfrequently and unpredictably.These speciesmay

best be represented by model 1b (fig. 1).

	 Several other patterns were consistent across multiple trail miles.

All miles were characterized by at least one spring-flowering cluster

(clusters C or F), at least one summer-flowering cluster (clusters D,

E, G, or J), and a bimodal flowering cluster (cluster B; fig. 2). Miles

1 and 2 exhibited multiple distinct spring peaks. Miles 2, 3, and 4

also exhibited a mid-year flowering cluster (cluster H), comprised

primarily ofsucculents(table 1). Cluster peaks were generally higher
USDA Forest Service Proceedings RMRS-P-67. 2013 35

Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion Crimmins, Crimmins, and Bertelsen

Figure 1—Conceptual model of potential probability of species flowering (PSF) models and the input data structure associated 

with these models. Potential PSF models appear in the left column; input data matrices appear in the right column. In the input 

data matrices, cells represent individual days of the year across a series of years. Observations of a species in flower are represented by cells shaded gray; observations of no flower are represented by unshaded cells. (a) a species that flowers in most 

years, for a short duration and with high consistency in year-to-year timing; (b) a species that flowers infrequently, with a short 

flowering duration, and with high year-to-year consistency in timing; (c) a species that flowers in most years, with high year-toyear consistency in timing, and for a long duration; (d) a species that flowers infrequently, with high year-to-year consistency in 

timing, and for a long duration; (e) a species that flowers for a short duration and with low year-to-year consistency in timing.
36 USDA Forest Service Proceedings RMRS-P-67. 2013

Crimmins, Crimmins, and Bertelsen Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion

Figure 2—Probability of species occurrence plots for 5 trail miles. (a) mile 1; (b) mile 2; (c) mile 3; (d) mile 4; (e) mile 5. Dashed lines 

indicate that the cluster is shared between multiple miles; solid lines indicate that cluster is unique to that trail mile. Thicker lines denote 

clusters that represent ≥10% of species in that mile.
USDA Forest Service Proceedings RMRS-P-67. 2013 37

Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion Crimmins, Crimmins, and Bertelsen

and narrowerin the summer, particularly in the higher elevationmiles

(4 and 5). Membership of several of the clusters in each mile was

quite small (5% or less of the species observed in that mile; fig. 2

and table 1). The distinct flowering patterns exhibited by the species

in these clusters are what drive these clusters to exist.

	 Clusters tended to be dominated by one plant functional type or

comprised of a consistent mix of functional types across the elevation gradient (table 1). Cluster B, the bimodal flowering pattern, was

comprised of between 73% and 100% herbaceous perennial species

(table1).ClusterC,the springfloweringclusterseenacross all5miles,

was a mix of annual and perennialspecies and exhibited a probability

of flowering around 60% in miles 1-3; in contrast, the earliest spring

flowering cluster exhibited lower probability of flowering, between

40-50% (miles 1 and 2; Cluster F), and was comprised of >75%

annual species (table 1). Cluster A, seen in all miles, was a mix of

annual and herbaceous perennial species.

	 For clusters identified in multiple miles, spring cluster peaks

generally occurred later at higher miles (fig. 2). The peak for cluster

B occurred as a plateau between April 13 and April 27 in mile 1 and

peaked around May 19 in mile 5. Summer peaks generally did not
38 USDA Forest Service Proceedings RMRS-P-67. 2013

Crimmins, Crimmins, and Bertelsen Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion

show a lag with increasing elevation. For example, cluster B peaked

around September 2 in mile 1 andAugust 30 in mile 5 in the summer.

Similarly, the peak for cluster D was approximatelyAugust 30 across

all trail miles.

Discussion

	 The largemembershipinclusterAcharacterizesthegreatvariability

in which species flower from year to year in the Sonoran Desert and

Sky Islands ecosystems; this analysis reveals that greater than half

of the species in each trail mile exhibit a smaller than 10% chance

of flowering in any given year or on any particular day. The species

in this cluster appear to exemplify the opportunistic nature of many

desert plantsthat are known or expected to depend heavily on seasonal

moisture and particular antecedent climate conditions to trigger and

complete germination, growth, and flowering (Beatley 1974; Bowers

1987; Bowers and Dimmitt 1994; Kemp 1983). This cluster is comprised of a mix of annual plants, herbaceous perennials, and woody

species, which employ different strategies for dealing with variable

moisture conditions. As the species in this cluster exhibit very low

probability of flowering throughout the entire year and from year

to year, these species probably represent a wide range of various

antecedent and interacting conditions that cue and sustain flowering.

	 Aside from the dominant pattern of pronounced year-to-year variability captured in cluster A, there are small “core” groups of species

that are consistently seen in flower, similar to the model in figure 1a,

regardless of the preceding or concurrent climate conditions. The

flowering reliability of these “core” groupsincreases with increasing

elevation, and is also higher in summer than in the spring season. The

higher probabilities observed in flowering in summer-season species

can be explained by these species’reliance on the summer monsoon

to initiate flowering. Compared to winter and spring precipitation,

the onset of the summer monsoon is rather regular in this region, and

many species’flowering cues are closely tied to this seasonal rainfall

(Crimmins and others 2011).

	 The results of this analysis demonstrate clear patterns among

functional types that are nested within the broad bimodal flowering

patterns characteristic of the Sonoran Desert. At lower elevations,

the spring season is characterized by multiple overlapping waves

of flowering. The first of these waves is characterized primarily by

annual species, though the chances of these species flowering are

around 50%. At higher elevations, the spring season is characterized

by a mix of life forms and a very small number of species flowering.

	 At low elevations, the probability ofspecies flowering increases as

spring progresses, as doesthe relative proportion ofspeciesin flower

that are perennial. Similarly, there are patterns within the summer

flowering season that vary by functional types composition, though

the timing of these peaks exhibits less temporal variability than the

spring season.

	 Our results show a progressively later spring flowering peak with

increasingelevation, consistentwithstudiesinhighelevationenvironments, which have demonstrated a lag in the onset of flowering with

increasingelevationwhere temperature isthemostinfluential variable

(Rusch 1993; Ziello and others 2009). Indeed, spring flowering of

species in this study area has been shown to be heavily influenced by

temperaturevariables(Crimmins andothers2010).In contrast,we did

not see a lag in peak flowering dates with increasing elevation in the

summer season. This pattern may be a result of the water limitation

in this system. Crimmins and others (2011) showed that across this

elevation gradient, the onset of summer flowering is driven by the

commencement of the summer monsoon rains and that, especially

in dry years, onset of summer flowering occurs rather synchronously

across the plant communities occupying the five trail miles.

	 The results of thisstudy have value for informing long-term monitoring. Recognizing the low probability of flowering for greater than

half of the species present in this system in a particular year could

significantly benefit monitoring by informing sampling time frames

and/or species selection. Further, if monitoring goals necessitate

selection of either consistently or infrequently flowering species, this

clustering approach could provide guidance. 

	 The findings of this analysisserve to increase our understanding of

the seasonal and year-to-year patterns in plant flowering across this

elevationgradient,establishingacriticalbaselineforcomparisonunder

future climate conditions. Recent studies in temperate environments

have shown varying sensitivity to climate, with earlier-flowering

species advancing their flowering dates at a more rapid pace than

later-flowering species(Miller-Rushing and Inouye 2009; Sherry and

others 2007). Future research in this unique, water-limited system,

where species are adapted to high climate variability, should explore

whether this pattern can be expected to manifest here as well. Implications of shifts include opening niches for potential invasion by

non-native species and de-coupling of trophic interactions.

Acknowledgments

	 We sincerely appreciate the helpful reviews of this manuscript that

Jherime Kellermann and Kathy Gerst provided.

References

Aronson, J.; Kigel, J.; Shmida, A.; Klein, J. 1992. Adaptive phenology of

desert and Mediterranean populations of annual plants grown with and

without water stress. Oecologia 89: 17-26.

Beatley,Janice C. 1974. Phenological events and their environmental triggers

in Mojave desert ecosystems. Ecology 55: 856-863.

Bowers, Janice E. 2005. El Niño and displays of spring-flowering annuals in

the Mojave and Sonoran deserts. Journal of the Torrey Botanical Society

132(1): 38-49.

Bowers, Janice E.; Dimmitt, M.A. 1994. Flowering phenology of six woody

plants in the northern Sonoran Desert. Bulletin of the Torrey Botanical

Club 121(3): 215-229.

Bowers, MichaelA. 1987. Precipitation and the relative abundances of desert

winter annuals: a 6-year study in the northern Mohave Desert. Journal of

Arid Environments 12: 141-149.

Burgess,TonyL.1995.Desertgrassland,mixedshrubsavanna,shrubsteppe,or

semidesertscrub? The dilemma of coexisting growth forms. In: McClaran,

Mitchell, P.; Van Devender, T.R., eds. The desert grassland. Tucson: The

University of Arizona Press. 346 p.

Crimmins, Theresa M.; Crimmins, M.A.; Bertelsen, D.; Balmat, J. 2008.

Relationships between alpha diversity of plant species in bloom and

climatic variables across an elevation gradient. International Journal of

Biometeorology 52: 353-366.

Crimmins, Theresa M.; Crimmins, M.A.; Bertelsen, C.D. 2010. Complex

responsesto climate driversin onset ofspring flowering across a semi-arid

elevation gradient. Journal of Ecology 98: 1042-1051.

Crimmins, Theresa M.; Crimmins, M.A.; Bertelsen, C.D. 2011. Onset of

summer flowering in a ‘Sky Island’ is driven by monsoon moisture. New

Phytologist 191: 468-479.

Ehleringer, James R.; Phillips, S.L.; Schuster, W.S.F.; Sandquist, D.R. 1991.

Differential utilization of summer rains by desert plants. Oecologia 88:

430-434.

Kemp, Paul R. 1983. Phenological patterns of Chihuahuan Desert plants in

relation to the timing of water availability.Journal of Ecology 71:427-436.

Kimball,Sarah;Angert,A.L.;Huxman,T.E.;Venable,L.E.2009.Contemporary

climate change in the Sonoran Desert favors cold-adapted species. Global

Change Biology 16:1555-1565.
USDA Forest Service Proceedings RMRS-P-67. 2013 39

Temporal Patterns in Species Flowering in Sky Islands of the Sonoran Desert Ecoregion Crimmins, Crimmins, and Bertelsen

McClaran,MichellP.;VanDevender,T.H. 1995.The desert grassland.Tucson:

The University of Arizona Press. 346 p.

Miller-Rushing, Abraham J.; Inouye, D.W. 2009. Variation in the impact of

climate change on flowering phenology and abundance: An examination

of two pairs of closely related wildflower species. American Journal of

Botany 96: 1821-1829.

Noy-Meir, Imanuel. 1973. Desert ecosystems: environment and producers.

Annual Review of Ecology and Systematics 4: 25-51.

Peñuelas,Josep;Filella,I.;Zhang,X.;Llorens,L.;Ogaya,R.;Lloret,F.;Comas,

P.; Estiarte, M.; Terradas, J. 2004. Complex spatiotemporal phenological

shifts as a response to rainfall changes. New Phytologist 161: 837-846.

Rusch,VeronicaE. 1993.Altitudinal variation in the phenology ofNothofagus 

pumilo in Argentina. Revista Chilena de Historia Natural 66: 131-141.

Sherry,RebeccaA.;Zhou,X.;Gu, S.;Arnone,J.A.III; Schimel,D.S.;Verburg,

P.S.; Wallace, L.L.; Luo, Y. 2007. Divergence of reproductive phenology

under climate warming. Proceedings of the NationalAcademy of Sciences

of the USA. 104: 198-202.

Ziello, Chaira; Estrella, N.; Kostova, M.; Koch, E.; Menzel, A. 2009. Influence of altitude on phenology ofselected plantspeciesin theAlpine region

(1971-2000). Climate Research 39: 227-234.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
40 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Are Madrean Ecosystems Approaching Tipping Points? 

Anticipating Interactions of Landscape Disturbance and 

Climate Change

Donald A. Falk

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona 

Abstract—Contemporary climate change is driving transitions in many Madrean ecosystems, but the time 

scale of these changes is accelerated greatly by severe landscape disturbances such as wildfires and insect 

outbreaks. Landscape-scale disturbance events such as wildfires interact with prior disturbance patterns 

and landscape structure to catalyze abrupt transitions to novel ecosystem configurations. These reorganized 

landscapes are characterized by new disturbance regimes and potentially dramatic departures in species 

diversity and distributions, community composition, carbon storage, landscape mesoclimate, and soil and 

hydrologic processes. Post-transition ecosystems can be highly resilient in their altered state, and resistant 

to return to pre-disturbance conditions in the current and near-term climate regime, possibly representing 

alternative metastable states. Severe landscape disturbance interacts with, and may be driven by, climate 

variation to govern ecosystem dynamics and large-scale ecosystem change. We present a conceptual model 

organized around six key attributes that may drive ecosystems to rapid change and novel trajectories.

Background

	 Many studies predict changes in species distributions in response

to changing climate.At localscales, community composition reflects

changes in the suitability of existing habitat for species persistence.

Both modeling and empirical studies suggest that such changes due

to climate alone are likely to be expressed at multi-annual to decadal

time scales (Allen and Breshears 1998, summarized in Parmesan

2006). Recent meta-analyses of range limit studies (Parmesan and

Yohe 2003; Chen and others 2011) found that latitudinal and upper

elevational limits in the Northern Hemisphere have moved 6.1-16.9

km northward and 6.1-11.0 m upslope per decade. The mechanisms

of climate impacts on ecological communities varies, including shifts

in species distributions (Gonzalez and others 2010; Rehfeldt 2006),

disease (Pounds and others 2005), and phenology (Crimmins and

Bertelson 2009).

	 In contrast,severe large-scale disturbances can reorganize ecosystems on much shorter time scales of days to months (Overpeck and

others 1990). High severity fires can cause extensive mortality, alter

soils and hillslope geomorphology, and trigger cascading ecosystem

changes, especially in vegetation types for which severe fires are

atypical (fig. 1). Landscape fires can cause rapid reorganization of

ecosystem mass and energy (McKenzie and others 2011), with the

consequence that post-fire recovery can extend over years or even

decades.

	 Post-fire ecologicaltransitions have been observed inmultiple sites

in Southwestern North America. Iniguez (2009) observed evidence

of a 19th century type conversion near Rincon Peak in the Rincon

Mountains (fig. 2) from ponderosa pine to an evergreen oak community, which appearsto have occurred following a high-severity fire in

1867.Savage andMast(2005)reviewedpost-fire recoverytrajectories

at 10 ponderosa pine sites in the Southwest that experienced crown

fires from 1948-1977. Some sites returned to pine dominance, albeit

at higher density than previously, while other areas converted to oak

shrubfields, chaparral, or non-forested grasslands.Areas burned in the

1,858-ha 1977 Radio Fire on Mt. Elden outside of Flagstaff, Arizona

(Passavoy and Fulé 2006), which burned largely in ponderosa pine,

remain dominated by shrubs 45 years post-fire. Similar transitions

have been observed in the Santa Catalina Mountains following the

2002-2003 Bullock and Aspen fires.

	 We posit that it is the combination of climate change and severe

disturbance that is most likely to trigger abrupt ecosystem transitions

into novel configurations, rather than either factor acting separately.

These new configurations may be resilient in their new state, and

resistant to return to pre-disturbance conditions due to synergistic

influences of disturbance and climate (Kitzberger and others 2011).In

additiontoecological effects,these transitions alsoincludepotentially

persistent alterations to geomorphic, soil, hydrological, and biogeochemicalsystems(Scheffer and others 2001). Such abrupt transitions

are predicted to become more common under conditions of altered

future climate and amplified disturbance regimes (Flannigan and

others 2000; Westerling and others 2006; Zinck and others 2011).

	 Abruptecologicalchangeisgenerallydefinedasathresholdresponse

in key biotic and abiotic ecosystemcomponents(Anderson and others

2008). Such changes may be expressed in shifts in dominant vegetation physiognomy (for example, from forest to shrub communities),

species composition, or community structure.Rapid post-disturbance

transitions are especially likely if established individuals of dominant
USDA Forest Service Proceedings RMRS-P-67. 2013 41

Are Madrean Ecosystems Approaching Tipping Points? Falk

Figure 1—2011 Horseshoe 2 Fire, Chiricahua Mountains.

Figure 2—Remnant southwestern white pine (Pinus strobiformis) with 

multiple fire scars in a netleaf oak (Q. rugosa) shrub field originating after stand-replacing fire in 1867, Rincon Peak, Arizona (photo: 

J.M. Iniguez).

species do not survive a disturbance event. Altered soil, hydrologic,

andbiogeochemical conditions canreinforce shiftsinvegetationtypes

and disturbance regimes,such as a transition from a low-severity forest surface fire regime to a high-severity chaparral shrubland regime

(Mayer 2011). Climate provides the envelope within which these

dynamics occur; disturbance provides the trigger for abrupt system

reorganization.

Theoretical Foundation

	 The theoretical basis for understanding type conversion behavior

derives from alternative stable state theory. This family of models

describes the potential of ecological communities to exist in more

than one state over relatively long ecological time. Strictly speaking,

these ecological configurations are more properly metastable states,

i.e., displaying quasi-equilibrial behavior over limited domains of

time and under a range of external conditions(e.g., long successional

sequences in a given climate). Henceforth, we will refer to alternative metastable state (AMS) theory, which provides a framework

for ecosystem behavior under both historical and current conditions

(Elmqvist and others 2003).

	 Bydefinition,metastable statesmaintainagivenconfiguration(e.g.,

a plant community type) within limited degrees of environmental

variation. For example, Madrean ponderosa pine stands or encinal

woodlands retain their primary characteristics (species composition,

physiognomy, biomass distribution) despite interannual variation

in rainfall and temperature, or episodic low-severity fires or insect

outbreaks, due to the persistence of established individuals and ongoing reproduction. The community can be displaced or reorganized

over long (millennial) time by climate variation, which modifies the

envelope ofsuitability,such as has occurred in the Madrean bioregion

throughout the late Pleistocene and Holocene (Holmgren and others 2006; Jackson 2006; Van Devender and others 1984). Transient

events—unusually severe fires, droughts, or insect outbreaks—have

the potential to trigger rapid community reorganization outside of the

envelope of metastability, especially if there is widespread mortality

and/or alteration ofthe biophysical template (Folke and others 2004).

Thustransient events, in combination with altered climate,maymake

rapid ecosystem transitions more likely.

	 Transitions between metastable states are distinguished from ongoing successional change by the rapidity with which state changes

occur. This occurs because the boundaries between metastable states

are characterizedbypositive-feedbackinteractionsthat createunstable

equilbria. Following a major perturbation, the system reorganizes

along one of several possible pathways. One potential outcome is a

return to the pre-disruption state, but depending on a variety of factors (see below), the system can instead develop along an alternative

trajectory. Positive feedback relationships then tend to reinforce a

given trajectory once it has been initiated, because species present in

the post-disturbance community provide a source of propagates for

ongoing recruitment as well as competitors to other species. These

same feedbacks create the emergent property of resilience in a given

condition once it is established. Unstable equilbria are often asymmetric: a change in one direction is not necessarily equally as likely

as a reciprocal change in the other, due to mechanisms that tend to

reinforce a given configuration. For example, once dry pine forest

has been converted to interior chaparral, the resulting new higherintensity fire regime may tend to favor resprouting chaparral shrub

species, and select against a return to slower-growing arboreal life

forms with heat-sensitive canopies;similar dynamics are observed in

semi-arid ecosystemsinvaded by cheatgrass(Bromustectorum), which

facilitates fire regimes that exclude previously dominant species.

	 Post-disturbancepathways are sensitive toinitial conditions,which

are a primary factor in the assembly rulesthat govern how communitiesreorganize (Palmer and others 1997;Temperton and others 2004).

For example, the presence of survivors and early post-disturbance

colonizers can determine establishment order and thus the capture
42 USDA Forest Service Proceedings RMRS-P-67. 2013

Falk Are Madrean Ecosystems Approaching Tipping Points? 

of space and resources. This early dominance can be reinforced by

growth, reproduction, and competition, resulting in a new metastable

state. It is also important to recognize that in some cases, what appear

to be alternative metastable states may simply represent slower successionaldynamicsoperatingonlongertime schedulesthanordinarily

recognized.

A Framework for Anticipating Threshold 

Change

	 We propose a model based on six primary factors that appear to

determinepost-fire ecosystemtrajectoriesunder current andnear-term

climate in Madrean ecosystems. In this model, the post-disturbance

trajectory of an ecosystem is influenced by (1) the type and severity

of disturbance, (2) persistence or mortality of dominantspeciesin the

pre-disturbance community,(3) alteration ofthe biophysical template

(soils, hydrology, topography), (4) landscape structure that governs

multiple spatial ecologicalprocessessuchasdispersal,(5)the local and

regionalspeciespoolsof availablepost-disturbance colonizers, and(6)

respective lifehistoryattributes, especiallydispersal,recruitment, and

competition, and local and regional climate that determines, among

other factors, the availability of suitable conditions for recruitment.

Interactions among these mechanismsinteract to create the emergent

properties often associated with macrosystem behavior, including

resistance and resilience. These variables can be used to simulate

threshold responses in modeling environments (Keane and others

2011).

Type and Severity of Disturbance—A wide range of ecological

and Earth system processes are classified as “disturbance,” including wildfire, insect outbreaks, diseases, floods, hurricanes, tsunamis,

landslides, earthquakes,severedroughts andpluvials,freezingevents,

and numerous others. Some of these (insect outbreaks, disease, and

fire) are clearly of biotic origin and can, thus, be considered ecological interactions, which operate at least in part in a density-dependent

fashion (i.e., effects are a function of density and size of members of

a community). Other disturbances(e.g., floods, hurricanes, tsunamis,

landslides, earthquakes, severe droughts and pluvials, and freezing

events) are expressions ofEarth systemprocessesinwhich ecological

effects tend to be density-independent. Surface and mixed-severity

fires in the Madrean ecoregion are keystone ecosystem processes

that regulate biomass, nutrient cycling, and community structure in

a wide range of community types (Swetnam and others 2001).

	 The introductionofnewspeciesintoa communitycanalsointroduce

noveltypesofdisturbance;for example,the rapidspreadofnon-native

African grasses such as buffelgrass (Pennisetum ciliare) into upland

communitiesin the Sonoran Desert hasintroduced a novel fire regime

to which most resident species are poorly adapted (McDonald 2009,

Stevens and Falk 2009). As discussed below, these shifts in the fire

regime canmove an ecosysteminto a new self-reinforcingmetastable

state which is highly resistant to return to the pre-invasion condition.

Mortality and Persistence—Severe disturbancessuch as wildfires

leaverelativelyfewsurvivingestablishedindividuals,openingupniche

and physical space for community-level turnover. Indeed, mortality

is one of the two primary ways in which severity is indexed; for

example, the U.S. Rapid Assessment of Vegetation Condition after

Wildfire(RAVG)program(http://www.fs.fed.us/postfirevegcondition/

index.shtml) usesmulti-spectralreflectance criteria to assess post-fire

burn severity, by classifying basal area loss by vegetation type (fig.

3). The extent of mortality among the pre-fire community is a key

determinant of post-fire trajectories, because established individuals

provide communities with physical structure, seed sources, nurse

plant microenvironments, sources of leaf litter, habitat resources for

wildlife, in situ geochemical poolssuch as carbon, nitrogen, and other

key nutrients, shade, microclimate regulation, and many other functions. The legacy of persisting individuals also represents the initial

conditions for community reassembly, and thus, their presence or

absence has a major impact on development of the post-disturbance

community.

Soil, Hydrologic, and Geomorphic Effects—Low-severity fires

appear to have few adverse effects on soils and geomorphology and

can contribute to topsoil development through mineralization of

nitrogen into plant-available forms, reduction of litter accumulation

that makes mineral soil more accessible to seedling roots, and accelerated decomposition of coarse woody debris. In contrast, severe

fires can alter soils and hydrology rapidly and substantially through

a variety of mechanismsincluding soil hydrophobicity, volatilization

of nitrogen and other key nutrients, loss of soil organic carbon, and

loss of litter and duff layers that assist with slope stabilization (Ice

and others 2004; Neary and others 2008; fig. 4).

	 Thesedirectfirst-orderfire effects(i.e.,immediate effectsthatoccur

as a direct consequence of combustion) are compounded by a wide

range of second-order fire effects including reduced water percolation, increased overland water flow, sheet erosion, sediment fluxes

into stream channels, debris flows, bank destabilization, accelerated

topsoil loss, and hillslope failures (DeBano and others 1998). For

example, extensive post-fire debris flows have been documented

in the Huachuca and Chiricahua Mountains (Youberg and others,

this volume) in the monsoon season following severe fires in 2011.

Recovery times for stream channels and riparian communities after

severe debris flows can be on the order of decades to centuries.

	 Where these effects are slight to moderate in severity and extent,

many communities will recover after fire to a state similar to the prefire condition. However, when soil and hydrogeomorphic impacts are

severe and widespread, an immediate return to the pre-fire condition

may be precluded because the biophysical template is no longer suitable. Extensive soil loss also eliminatesthe soilseed bank, which is a

key component of post-disturbance biotic recovery. Hart and others

(2005) found that severe fires can also alter the soil microbial communities (both fungal and bacterial) through extensive belowground

mortality. These changes can be linked to shifts in plant community

composition and changes in key ecosystem processes such as C sequestration, nutrient fluxes,soil physicalstructure, and plant-microbe

mutualisms.

Landscape Structure and Dispersal Processes—The spatial

properties of post-fire environments are important determinants of

post-fire trajectories. Large fires characteristically leave a legacy of

complex landscape mosaics of burn severity across vegetation types.

Roughly 4 to 35%ofthe areawithin the perimeters oflarge recentfires

in the western United States is classified as high burn severity using

delta Normalized Burn Ratio (dNBR), and Normalized Difference

Vegetation Index (NDVI) indices (www.mtbs.gov). Moderate severity typically accounts for 20 to 45%, and low severity or unburned

areas 33 to 61% (Kotliar and others 2003; Lentile and others 2007).

Less is known about the severity patch structure of historical fire

regimes, although tree-ring studies in high fire frequency systems

such as ponderosa and Jeffrey pine (P. jeffreyi) forests indicate that

the extentofhigh-severityeffectswasgenerallylimited(Swetnamand

others 2001; Falk and others 2011). At higher elevations, however,

patches were generally larger contiguous areas of high severity in

wet mixed-conifer and spruce-fir forests(Margolis and others 2007).

	 In addition to the proportion ofthe landscape in each severity class,

the size of contiguous patches is an important landscape variable
USDA Forest Service Proceedings RMRS-P-67. 2013 43

Are Madrean Ecosystems Approaching Tipping Points? Falk Figure 3—Burn severity images from 2011 Horseshoe 2 fire, Chiricahua Mountains, Arizona. Left: Vegetation burn severity, Rapid Assessment of Vegetation Condition after Wildfire (RAVG), USFS Remote Sensing Applications Center (RSAC, (http://www.fs.fed.us/postfirevegcondition/index.shtml). Right: soil burn severity,), accessed through Monitoring Trends in Burn Severity (MTBS, www.mtbs.gov ).
44 USDA Forest Service Proceedings RMRS-P-67. 2013

Falk Are Madrean Ecosystems Approaching Tipping Points? 

(Haire and McGarigal 2010). In contrast to historical fire regimes,

contemporary fires often create large patches of high-severity fire

effects in the order of 1,000-10,000 ha in size in what were formerly

low-severity/high-frequency systemssuch as ponderosa pine and dry

mixed conifer forests where such patches are believed to have been

historically of 10-100 ha (fig. 5). The resulting altered landscape

structure such as anomalously large high-severity patches following

extreme disturbance events may restrict seed dispersal of previously

dominant species while favoring other species with rapid long-range

dispersal. For example, Bonnet and others (2005) found that most

post-fire seed dispersal following a high-severity fire in a South

Dakota ponderosa pine forest was restricted to approximately 150

m from seed sources on the edge of residual forest. By this measure,

the centers of high severity patcheslarger than 10 ha with a minimum

lateral dimension of ≥300 m would require multiple generations for

propagulestoreachthe interiorbynaturaldispersalprocesses.Species

withlonger-rangedispersalorwithpersistingvegetativepotential(e.g.,

sprouting species) would have a significant advantage in capturing

post-fire occupancy (Haire and McGarigal 2010).

Local and Regional Species Pools—The pool ofspecies available

to recolonize a site following disturbance is a key determinant of postfire ecological trajectories(Menninger and Palmer 2006). Even when

suitable biophysical conditions exist for establishment, species must

be present and be able to reproduce to become part of the developing community. Clearly, species that can persist through disturbance

have a significant advantage over othersthat must disperse to the site.

Persistence is, in turn, a function of evolved life history attributes

that reflect a range of evolutionary responses to disturbance (Bond

and van Wilgen 1996). For example, the thick bark and lifted crowns

characteristic of ponderosa and Jeffrey pines arewidely understood as

adaptations to surviving low-intensity fires burning in surface fuels,

in which survivorship of mature individuals is maximized (Keeley

and Zedler 1998). Other species including Chihuahua pine (P. leiophylla) and many Madrean oaks (Quercus spp.) can resprout after

being top-killed, providing a persistence mechanism that may facilitate rapid post-fire occupancy (fig. 6). Aspen (Populus tremuloides)

clonessimilarly employ a vegetative life history strategy for post-fire

response, althoughbothoaks andaspenalsoreproduce sexually.These

reproductive strategies allow these native speciesto capture post-fire

environments quickly and to persist once established. For example,

Gambel oak (Q. gambellii) was observed resprouting within weeks

of both the 2000 Cerro Grande Fire and the 2011 Las Conchas Fire in

the Jemez Mountains, New Mexico. These evolutionary adaptations

to post-fire colonization suggests that type conversions (e.g., from

pine-dominated forest to oak-dominated scrub), however long they

may persist, are part of the historical post-fire behavior of Madrean

ecosystems. However, other factors (such as climate and landscape Figure 4—High soil and vegetation burn severity on steep slopes of 

Cochiti Canyon, 2011 Las Conchas fire, Jemez Mountains, New Mexico.

Figure 5—Large high-severity burn patch, 2011 Horseshoe 2 fire, 

Chiricahua Mountains, Arizona.

Figure 6—Rapid resprouting of oaks following 2011 Monument 

Fire, Huachuca Mountains, Arizona.
USDA Forest Service Proceedings RMRS-P-67. 2013 45

Are Madrean Ecosystems Approaching Tipping Points? Falk

change) may render these conversions permanent, or at least more

persistent than in pre-settlement environments.

	 By contrast, in upper Sonoran Desert communitiesin the Madrean

bioregion, the primary agents of abrupt and radical change in the native fire regime are invasive non-native grasses. These changes are

among the clearest expressions of tipping point behavior in Madrean

ecosystems.NaturalSonoranDesertvegetationprovidesverylowfine

fuel mass and connectivity due to the dominance of intermingling

woodyvegetationformuchoftheyear andwide spacingof established

plants. Under these conditions, ignitions rarely generate a spreading

fire due to the lack of sufficient spatially-continuous fine fuels to

sustain a chain reaction. As a result, fires in the Sonoran Desert are

rare and low intensity (heat per unit area-time). Consequently, most

species of the community lack adaptations to survive fire.

	 IntroducedAfrican grasses, particularly buffelgrass, have changed

the fuel environment in Sonoran communities by infilling interspaces

with increased, spatially continuous fine fuels (fig. 7). In addition,

the foliage of buffelgrass is persistent for many months after curing,

increasing the flammability ofthe fuel bed forthe driest months ofthe

year.Buffelgrassisfire tolerant, a formidablebelowgroundcompetitor

(Olsson and others 2011; Stevens and Falk 2009), and can rapidly

assume dominance in the herbaceous community. The net result is

that areas that have been invaded by buffelgrass become converted

to a novel community type with high herbaceous fuel loads of up to

600-1,200g/m2,twoordersofmagnitudemore thanun-invadeddesert

(McDonald 2009) and frequent high-intensity fires that are lethal to

manynativedesertspecies.Columnar cactisuchassaguaro(Carnegia 

gigantea) are particularly susceptible due to their thin epidermis that

is easily damaged by levels of heat typically generated in a wildfire.

Once the “grass-fire cycle” is established, the community is resilient

in its new configuration and resistant to return to the pre-invasion

state (Brooks 2008).

Climate Change—The “bioclimatic envelope” is a central determinant of species distributions and climate variables are key dimensions in a species’ niche space. Niches are non-spatial combinations

of multiple variablesthat define the conditions under which a species

can maintain a non-negative population growth rate (λ). The niche

is the product of species evolution, including phenotypic plasticity;

where λ ≥ 1, a population is self-sustaining (Colwell and Rangel

2009). When the hyperdimensional niche volume is mapped onto

geographic space, the result is the total potential spatial distribution

of a species in the absence of competition or other limiting factors.

Regional climate along with soils, topography, and hydrology also

provides the context for the distribution of biotic communities.

	 Although climate-species and climate-community relationships

are mechanistic, they are not deterministic; many more community

configurations are possible within a given climate envelope at a given

location and few species occupy their entire potential geographic

range. In niche theory, the realized niche is the portion of the total

fundamental niche hypervolume that a species occupies. Realized

niche space is narrower than the fundamental niche for any given

axis due to competition, dispersal limitations, and other factors. The

geographic expression of the realized niche is the potential habitat

of a species. Similar relationships hold for the distribution of biotic

communities as assemblages of multiple interacting species. These

non-deterministic relationships are the foundation for the potential

for alternative metastable states.

	 Bioclimatic envelope (BCE) models are used widely to generate

null hypotheses for potential species responses to climate change. In

a BCE framework, changing climate conditions require a species to

follow one oftwo possible pathways(Colwell andRangel 2009). One

alternative is that the species retains its current niche requirements.

In this case, the species must move geographically to remain within

its potential habitat zone of suitable climate. In an era of increasing

global and regional temperatures, this is usually interpreted as pressure to move upslope or poleward. Thus, in this potential adaptive

response, niche space is retained, and adaptation occurs by movement in geographic space. The second alternative isthe converse; the

species adapts its niche requirements in situ to new conditions but

retains its current geographic distribution; in other words, the species evolves. Many species contain alleles, present in the population

at low frequency, that may provide the basis for adaptation to novel

conditions (e.g., tolerance of higher temperatures or drought). As

climate changes, these alleles may increase in their frequency and the

species’ niche space is altered to fit current conditions. Most species

also retain some phenotypic plasticity, which retains fitness under

variable conditions within the current genotypes and may facilitate

local adaptive responses.

	 Climate variability and secular (anthropogenic) trend provide the

envelope in which threshold responses may occur (Alley and others

2003; McNeall and others 2011). Species distributionsrepresent current geographic range in most BCE modeling, and thusimplicitly, the

climate toleranceof establishedindividuals.However,theniche space

for recruitment—the “regeneration niche”—is not only smaller than

the niche space for established individuals but occupies the cooler,

wetter domain oftheniche (fig.8).Established individuals of a species

such as ponderosa pine or Douglas-fir (Pseudotsuga menziesii) can

persist through dry years and even sustained periods of drought and

high temperatures, but these same conditions are lethal for recently

germinated seedlings. Thus, a projected warmer and drier climate in

the Madrean ecoregion may not allow for post-fire regeneration of

previously dominant species, especially if there is high disturbancerelated mortality.

	 In addition to regulating species distributions and post-fire regeneration, altered climate also increases the probability of severe fires

(Flannigan and others 2000). Fire initiation and spread are facilitated

by warmer, drier, windier conditions that reduce fuel moistures and

promote the combustion chain reaction (Pyne and others 1996). Fire

severity may also increase under these climatic conditions, due to

decreased live fuelmoistures and the resulting increasedflammability

of vegetation. Finally, the length of fire season is projected to increase

Figure 7—Prescribed experimental burn in buffelgrass stand, Altar 

Valley, Arizona (photo: C. J. McDonald).
46 USDA Forest Service Proceedings RMRS-P-67. 2013

Falk Are Madrean Ecosystems Approaching Tipping Points? 

with changing climate and seasonality, as reflected in the increased

number of days with energy release component (ERC) greater than

critical values (Brown and others 2004). Thus, altered climate may

play an additional destabilizing role in Madrean ecosystems by direct

alteration of fire regimes.

	 AsFlanniganandothers(2000)observe, “The almostinstantaneous

response of the fire regime to changes in climate has the potential to

overshadowimportance of direct effects of globalwarming on species

distribution,migration,substitutionandextinction...[F]ire is a catalyst

for vegetation change.” The collective effect of disturbance-climate

interactions is to rapidly and persistently change landscape patterns

and processes including disturbance regimes, vegetation composition, carbon dynamics, and hydrologic balance (Breshears and others

2005; O’Connor and others 2011). The central challenge confronting

ecosystem scientists and managersisto understand how fire behavior

and legacies of prior disturbancemay interact with climate variability

to influence post-fire recruitment and ecosystemtrajectories, and how

these transitions can be managed to maintain sustainable ecosystems.

Broader Impacts

	 The potential for abrupt ecological and hydrologic change in

response to climate and disturbance is of central importance to

ecosystem management in a changing world. As climate change

progresses, species assemblages and ecosystem disturbance regimes

are already changing in ways that challenge the foundations of ecosystem management, especially in arid and semi-arid regions (Allen

and Breshears 1998; Aronson and others 1993; Breshears and others

2005; Quijada-Mascareñas and others, in press). Major fires, insect

outbreaks, forest dieback, and other phenomena are already creating

new and serious stressors to terrestrial ecosystems (Overpeck and

others 1990). Moreover, the character, function, and integrity of

ecosystems are fundamental to human society in innumerable ways.

Thus, if ecosystems begin to change abruptly and profoundly, the

impacts are likely to ripple through human society as well as the

biosphere itself.

Acknowledgments

	 Ann Lynch, Rocky Mountain Research Station, U.S. Forest

Service, Tucson Arizona, and Linda Wadleigh, Fire and Aviation

Management, Southwestern Region, U.S. Forest Service, Flagstaff,

Arizona, provided valuable reviews and suggestions on the original

manuscript. The author also acknowledges the contributions of

Craig Allen (U.S. Geological Survey, Los Alamos, New Mexico);

Christopher Baisan (Laboratory of Tree-Ring Research, University

of Arizona, Tucson); David Breshears (School of Natural Resources

and the Environment, University ofArizona);Corinne Dolan (School

of Natural Resources and the Environment, University of Arizona);

William Edwards (Coronado National Forest, Douglas, Arizona);

Calvin Farris (National Park Service, Klamath Falls, Oregon); Peter

Fulé (School of Forestry, Northern Arizona University, Flagstaff,

Arizona);BrookeGebow(TheNatureConservancyofArizona);Perry

Grissom (Saguaro National Park); Jose Iniguez (Rocky Mountain

Research Station, U.S. Forest Service Flagstaff,Arizona); Mark Kaib

(U.S. Fish and Wildlife Service,Albuquerque, New Mexico); Rachel

Loehman (Fire Sciences Laboratory, Rocky Mountain Research

Station, U.S. Forest Service, Missoula, Montana); Ellis Margolis

(Laboratory of Tree-Ring Research, University of Arizona, Tucson,

Arizona);Adrian Quijada-Mascareñas (School of Natural Resources

and the Environment, University of Arizona); Melissa Savage (Four

Corners Institute, Santa Fe, New Mexico); Christopher Stetson

(Coronado National Forest, Douglas, Arizona); Elaine Sutherland

(Rocky Mountain Research Station, U.S. Forest Service Missoula,

Montana); Thomas Swetnam (Laboratory of Tree-Ring Research,

University of Arizona, Tucson); Andrea Thode (School of Forestry,

Northern Arizona University, Flagstaff, Arizona); and Craig Wilcox

(Coronado National Forest, Douglas, Arizona).

References

AllenC.D.;D.D.Breshears. 1998.Drought-induced shift of a forest-woodland

ecotone: rapid response to climate variation. Proceedings of the National

Academy of Sciences US. 95(25):14839-14842.

Alley,R.B.;[and others]. 2003.Abrupt climate change. Science. 299: 2005-2010.

Andersen, T., J. Carstensen, E. Hernandez-Garcia, and C.M. Duarte. 2008.

Ecologicalthresholds andregime shifts: approachestoidentification.Trends

in Ecology and Evolution. 24:49-57.

Aronson.J.,C.Floret,E.LeFloc’h,C.Ovalle, andR.Pontanier. 1993.Restoration and rehabilitation of degraded ecosystems in arid and semi-arid lands.

I. A view from the South. Restoration Ecology. 1: 8-17.

Bond, W.J., and B. van Wilgen. 1996. Plants and fire. Chapman & Hall:

New York.

Bonnet,V.H.,A.W.Schoettle,andW.D.Shepperd,2005.Postfireenvironmental

conditionsinfluence the spatial pattern ofregenerationforPinus ponderosa.

Canadian Journal of Forest Research. 35:37-47.

Breshears, D.D., [and others]. 2005. Regional vegetation die-off in response

to global-change-type drought. Proceedings of the National Academy of

Sciences US 102(42):15144-15148.

Brooks, M.L. 2008. Plant invasions and fire regimes.In Zouhar, K.J.K. Smith,

S.Sutherland, and M.L. Brooks (Eds.), Wildland fire in ecosystems: fire

and nonnative invasive plants. Gen. Tech. Rep. RMRS-GTR-42, Vol. 6.

U.S. Forest Service, Ogden, UT.

Brown, T.J., B.L. Hall, and A.L. Westerling. 2004. The impact of twenty-first

century climate change onwildlandfire dangerin thewesternUnitedStates:

An applications perspective. Climatic Change. 62:365–388.

Figure 8—The climate state space in which established individuals 

can persist (E0), illustrated here on axes of minimum growing season 

soil water potential (SWP) and maximum growing season temperature. 

Most climate projections for the Madrean bioregion indicate generally 

warmer and drier conditions (E1). Established individuals can persist in 

the intersection of these two spaces. However, climate requirements for 

seedling recruitment (R) are in the cooler, wetter domain of the climate 

space, which no longer intersects climate regime E1.
USDA Forest Service Proceedings RMRS-P-67. 2013 47

Are Madrean Ecosystems Approaching Tipping Points? Falk

Chen,I.-C.,J.K.Hill,R.Ohlemüller,D.B.Roy, andC.D.Thomas. 2011.Rapid

range shifts of species associated with high levels of climate warming.

Science. 333:1024-1026.

Colwell, R.K., and T.F. Rangel. 2009. Hutchinson’s duality: The once and

future niche. Proceedings of the National Academy of Sciences US. 106

(Supplement 2):19651-19658.

Crimmins, T.M., M.A. Crimmins, and C.D. Bertelson. 2009. Flowering range

changes across an elevational gradient in response to warming summer

temperatures. Global Change Biology. 15:1141-1152.

DeBano,L.F.,D.G.Neary, andP.F.Ffolliott.1996.Fire’s effectsonecosystems.

John Wiley &Sons, New York.

Elmqvist,T., C. Folke, M. Nyström, G. Peterson,J. Bengtsson, B.Walker, and

J. Norberg. 2003. Response diversity, ecosystem change, and resilience.

Frontiers in Ecology and the Environment 1:488-494.

Falk, D.A.,E.K. Heyerdahl, P.M.Brown,C.A. Farris, P.Z. Fulé, D. McKenzie,

T.W.Swetnam,A.H.Taylor, andM.L.VanHorne. 2011.Multiscale controls

of historical fire regimes: New insights from fire-scar networks. Frontiers

in Ecology and the Environment. 9:446-454.

Flannigan, M., B. Stocks, and B. Wotton. 2000. Climate change and forest

fires. The Science of the Total Environment. 262:221-229.

Folke,C., S.Carpenter,B.Walker,MScheffer,T.Elmqvist,L.Gunderson, and

C.S.Holling. 2004.Regime shifts,resilience, and biodiversity in ecosystem

management. Annual Review of Ecology & Systematics. 35:557-581.

Gonzalez, P., R.P. Neilson, J.M. Lenihan, and R.J. Drapek. 2010. Global patterns in the vulnerability of ecosystems to vegetation shifts due to climate

change. Global Ecology and Biogeography. 19:755–768.

Haire, S.L., and K. McGarigal. 2010. Effects of landscape patterns of fire

severity on regenerating ponderosa pine forests (Pinus ponderosa) in New

Mexico and Arizona, USA. Landscape Ecology. 25:1055–1069.

Hart,S.C.,T.H.DeLuca,G.S.Newman,M.D.MacKenzie, andS.I.Boyle.2005.

Post-fire vegetative dynamics as drivers of microbial community structure

and function in forestsoils. Forest Ecology & Management. 220: 166-184.

Holmgren, C.A., J.L. Betancourt, and K.A. Rylander. 2006. A 36,000-yr

vegetation history from the Peloncillo Mountains, southeastern Arizona,

USA. Palaeogeography, Palaeoclimatology, Palaeoecology. 240: 405–422

Ice, G.G., D.G. Neary, and P.W.Adams. 2004. Effects of wildfire on soils and

watershed processes. Journal of Forestry 102 (6):16-20.

Iniguez,J.M.,T.W.Swetnam, andC.H.Baisan. 2009.Spatially and temporally

variablefire regime onRinconPeak,Arizona,USA.FireEcology5(1):3-21.

Jackson, S.T. 2006. Vegetation, environment, and time: The origination and

termination of ecosystems. Journal of Vegetation Science 17:549-557.

Keane, R.E., R.A. Loehman, and L.M. Holsinger. 2011. The FireBGCv2

landscape fire and succession model: A research simulation platform for

exploring fire and vegetation dynamics. Gen.Tech.Rep.RMRS-GTR-255.

U.S. Department ofAgriculture, Forest Service,Rocky MountainResearch

Station, Fort Collins, CO.

Keeley, J.E., and P.H. Zedler. 1998. Evolution of life histories in Pinus. In

D.M. Richardson (Ed.), Ecology & biogeography of Pinus. Cambridge

University Press: Cambridge, UK: 219-250

Kitzberger T., E. Araoz, J.H. Gowda, M. Mermoz, and J.M. Morales. 2011.

Decreases in fire spread probability with forest age promote alternative

community states, reduced resilience to climate variability and large fire

regime shifts. Ecosystems DOI: 10.1007/s10021-011-9494-y.

Kotliar, N.B., S.L. Haire, and C.H. Key. 2003. Lessonsfrom the fires of 2000:

Post-fire heterogeneity in ponderosa pine forests. In Omi, P. N. and L. A.

Joyce (Eds.),Fire ecology,fueltreatments, and ecologicalrestoration.Proc.

RMRS-P-29. Fort Collins, CO: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station: 277-280.

Lentile, L.B, P. Morgan, A.T. Hudak, M.J. Bobbitt, S.A. Lewis, A.M.S.

Smith, and P.R. Robichaud. 2007. Post-fire burn severity and vegetation

response following eight large wildfires across the western United States.

Fire Ecology. 3(1):91-108.

Margolis E.Q., T.W. Swetnam, and C.D. Allen. 2007. A stand-replacing fire

historyinuppermontane forestsofthe southernRockyMountains.Canadian

Journal of Forest Research. 37: 2227–2241.

Menninger, H.L., and M.A. Palmer 2006. Restoring ecological communities:

Fromtheory to practice.In Falk, D.A., M.A. Palmer, and J.B. Zedler(Eds.),

Foundationsofrestorationecology.IslandPress,Washington,DC.Pp.88-112

Mayer, G.G., and A.H. Khalanyi. 2011. Grass trumps trees with fire. Science.

334:188-189.

McDonald,C.J.2009.Managementofnon-nativeperennialgrassesinsouthern

Arizona: Effects of prescribed fire and grazing. Ph.D. Dissertation. School

of Natural Resources, University of Arizona, Tucson.

McKenzie, D., C. Miller, and D.A. Falk (Eds.). 2011. The Landscape ecology

offire.EcologicalStudiesSeriesNo.213,Springer,Dordrecht,Netherlands.

McNeall, D., P.R. Halloran, P. Good, and R.A. Betts. 2011. Analyzing abrupt

andnonlinearclimatechangesandtheirimpacts.ClimateChange.2:663-686.

Neary, D.G., K.C. Ryan, and L.F.DeBano (Eds.). 2008. Wildland fire in

ecosystems: effects of fire on soils and water. Gen. Tech. Rep. RMRSGTR-42-vol.4.Ogden,UT:U.S.DepartmentofAgriculture,ForestService,

Rocky Mountain Research Station. 250 p.

O’Connor, C., T.W. Swetnam, G. Garfin, and D.A. Falk. 2011. Human

pyrogeography: A new synergy of fire, climate and people is reshaping

ecosystems across the globe. Geography Compass. 5/6:329-350.

Olsson, A.D., J. Betancourt, M.P. McClaran, and S.E. Marsh. 2011. Sonoran

Desert ecosystem transformation by a C4 grass without the grass/fire cycle.

Diversity and Distributions. 18(1):10-21.

Overpeck J.T., D. Rind, and R. Goldberg. 1990. Climate-induced changes in

forest disturbance and vegetation. Nature. 343:51-53.

Palmer, M.A., R.F. Ambrose, and N.L. Poff. 1997. Ecological theory and

community restoration ecology. Restoration Ecology. 5:291–300.

Parmesan, C. 2006. Ecological and evolutionary responses to recent climate

change.AnnualReviewofEcology,Evolution, andSystematics.37:637–69

Parmesan, C., and G. Yohe. 2003. A globally coherent fingerprint of climate

change impacts across natural systems. Nature. 421:37-42.

Passovoy, M.D., and P.Z. Fulé. 2006. Snag and woody debris dynamics following severe wildfires in northern Arizona ponderosa pine forests. Forest

Ecology and Management. 223:237–246.

Pounds,J.A.andothers.2005.Widespreadamphibianextinctionsfromepidemic

disease driven by global warming. Nature. 439:161–165.

Pyne, S.J., P.L.Andrews, and R.D. Laven. 1996. Introduction to wildland fire.

2nd Edition. John Wiley & Sons, New York.

Quijada-Mascareñas,A., D.A. Falk,J.Weiss, M. McClaran,J. Koprowski, M.

Culver, S. Drake, S. Marsh, W. van Leeuwen,A.M. Lynch, and M. Skroch.

[In press]. Potential multi-scale ecological effects of climate change in Sky

Island ecosystems. The Open Ecology Journal.

Rehfeldt, G.E., N.L. Crookston, M.V. Warwell and J.S. Evans. 2006. Empirical analyses of plant‐climate relationships for the western United States. 

International Journal of Plant Sciences. 167:1123-1150.

Savage M., and J.N. Mast. 2005. How resilient are ponderosa pine ecosystems

after crown fires? Canadian Journal of Forest Research. 35: 967-977.

Scheffer, M., [and others]. 2001. Catastrophic shifts in ecosystems. Nature.

413:591-596.

Stevens, J., and D.A. Falk. 2009. Using invasion ecology theory to explain

buffelgrass success and restoration potential in the American Southwest.

Ecological Restoration 27:417-427.

Swetnam, T.W., C.H. Baisan, and J.M. Kaib. 2001. Forest fire histories in the

SkyIslandsofLaFrontera.InG.L.Webster andC.J.Bahre (Eds.),Changing

plant life of La Frontera: Observations on vegetation in the United States/

MexicoBorderlands.UniversityofNewMexicoPress,Albuquerque:95-119.

Temperton, V.M., R.J. Hobbs, T. Nuttle, and S. Halle (Eds). 2004. Assembly

rules and restoration ecology. Island Press, Washington, DC.

Van Devender, T.R.,J.L. Betancourt, and M. Wimberly. 1984. Biogeographic

implicationsof apackratmiddensequence fromtheSacramentoMountains,

south-central New Mexico. Quaternary Research. 22(3): 344-360.

WesterlingA.L., H.G. Hidalgo, D.R.Cayan, andT.W. Swetnam. 2006.Warming and earlier spring increase western U.S. wildfire activity. Science. 313

(5789): 940-943.

Zinck, R. D., M. Pascual, and V. Grimm. 2011. Understanding shifts in

wildfire regimes as emergent threshold phenomena. American Naturalist.

178(6): E149-E161.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
48 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Preliminary Assessment of Changes in a Lizard Assemblage 

at an Ecotone in Southeastern Arizona

Lawrence L. C. Jones

USDA Forest Service, Coronado National Forest, Tucson, Arizona

Abstract—The Madrean Archipelago and its associated valleys have the highest diversity of lizards in the 

United States. This is due to a convergence of ecoregions in an area that provides excellent environmental 

conditions for life history needs of terrestrial ectotherms. The study area, near Safford, Arizona, is known to 

have about 20 species of sympatric lizards, although only about one-half are common. The lizard community 

is represented by species of the Sonoran and Chihuahuan deserts, semi-desert grasslands, and lower Madrean 

and boreal woodlands. It has recently been suggested that lizard species are expected to decline globally due 

to climate change and other reasons. A study site representing an ecotone between desert and grassland/

montane/riparian vegetation types in a foothills situation was chosen, as ecotones are marginal habitats that 

are expected to be sensitive to environmental change. Study objectives were to assess the baseline lizard 

community and detect changes in the lizard assemblage over time due to climate change and other factors. 

During systematic road transects, a total of 3,889 lizards representing 13 species were recorded during 60 

visits in 2003 (n = 8 visits), 2010 (n = 12,) and 2011 (n = 40). Although this represents the early stages of a 

long-term monitoring program, preliminary observations show differences in the lizard assemblage between 

years consistent with climate change predictions.

Introduction

	 The Madrean Archipelago’s “sky island” mountain ranges and

valleys have the highest diversity of lizards in the United States of

America; nearly one-third of all U.S. species (about 37-39/120) are

found in southeastern Arizona and adjacent New Mexico (Jones and

Lovich 2009). There can be about 15-20 sympatric species in a small

area, where the mountains meet the valleys (Degenhardt and others,

1996 [p. 208]; Jones 2009; Lazaroff and others, 2006).

	 Literature suggests lizards are declining for a variety of reasons

including habitat loss, invasive species, and competition with other

species(Gibbons and others, 2000; Olson 2012). Sinervo and others

(2010) documented local extinctions of lizards from study plots in

Mexico, making a cause-and-effects case for the decline in diversity

due to climate change. They went on to describe how the life history

attributes of lizards are such that increased spring temperatures and

other factors made lizards, as a group, susceptible to extirpation and

extinction. They predicted that by 2080 about 40% of lizard species

wouldbecome locallyextinct and20%wouldbecomegloballyextinct.

	 When the Sinervo and others(2010) paper was published, I reinitiated surveys from 2003 (Jones 2009) to identify potential changes

in the local lizard assemblage since that time, as well as qualitative

changessince the Nickerson and Mays(1969)inventory ofthe Pinaleño

Mountains. In this paper I document some changes in the lizard assemblage, although surveys to date are only the beginning of what is

intended to be a long-term monitoring project. Thus, results reported

here are preliminary or observational, but do help establish a baseline

of distribution and relative abundance of commonly encountered

species, as well as highlighting some inter-year trends. 

Study Area

	 The study area is about 10 mi S of Safford, Graham County,

Arizona, along a 2.7-mi stretch of road (Forest Road 57) from its

junction with Swift Trail (Highway 366). This dirt road is usually

termedMarijildaRoad,because it accessesMarijildaCreek.Elevation

ranges from about 3,810 to 4,010 ft above mean sea level. Adjacent

vegetation is primarily semi-desert grassland, but the area is situated

at an ecotone between two deserts (Sonoran and Chihuahuan) in the

San Simon Valley and semi-desert grasslands and Madrean encinal

(oak) woodlands in the foothills of the Pinaleño Mountains (Brown

1994). Vegetation is primarily composed of grasses, shrubs, cacti,

and smalltrees.LehmannLovegrass(Eragrostis lehmanniana, a nonnative bunchgrass), Velvet Mesquite (Prosopis velutina), Engelmann

PricklyPear(Opuntia engelmannii), andSoaptreeYucca (Yucca elata)

are among the common plant species, but desertscrub, riparian, and

encinal woodland species are also present (Jones 2009, unpublished

data).Because the area isrocky,sections ofthe shoulders of Marijilda

Road are replete with boulders and boulder piles, partially an artifact

of road construction.The first 0.4 miles of road approach the foothills

from the desert valley, then the road turnsto parallel the lowerslopes

of the mountains. The transect ends at Marijilda Creek, a perennial

stream with deciduous riparian vegetation.
USDA Forest Service Proceedings RMRS-P-67. 2013 49

Preliminary Assessment of Changes in a Lizard Assemblage at an Ecotone in Southeastern Arizona Jones

Methods

	 In 2003, 2010, and 2011,I conducted a road-transect tally oflizards

alongMarijildaRoad, accordingtoaprotocol;themethodisdiscussed

in greater detail elsewhere (Jones 2009).Basically,the observer drove

very slowly along Marijilda Road, and stopped when a lizard was

seen. The lizard was then viewed through binoculars and identified

to species and age class(hatchling, juvenile,subadult, adult), and the

0.1 mi segment of road was recorded. Only the driver tallied lizards

seen; passengers (if present) were not allowed to point out lizards.

Lizards in the Desert Spiny Lizard complex (Sceloporus magister

and S. bimaculosus) were recorded simply as “Sceloporus magister

complex” because both species are sometimes difficult to distinguish

and likely hybridize in the area (Jones and Schwalbe 2009; Phelan

and Brattstrom 1955). Lizards of the Aspidoscelis sonorae complex

are recorded as A. flagellicauda. Without genetic confirmation, I

consider A. flagellicauda to be the only species in the A. sonorae

complex present, although HerpNet (an online resource of museum

records) reports A. sonorae, A. flagellicauda, and A. exsanguis from

the immediate vicinity. All species in the complex are similar and

there are taxonomic uncertainties (Jones and Lovich 2009).

	 Starting in June 2010, surveys were conducted year-round. There

was a minimum of one survey per month during November through

February, and one survey per week March through October. Surveys

were conducted when temperature, sunlight, wind, and cloud cover

were conducive to surface activity of lizards (temperatures between

75 and 95 oF, with mid-80’s being optimal, plus low wind and low

cloud cover). Starting time was as early as 0700 in the summer and

as late as 1400 in the winter.

	 Age class data were recorded differently between 2010 and 2011

(not at all in 2003), and there was apparent observer bias in 2011. In

order to compare results of reproductive output between 2010 and

2011, I limited observations to my own and transformed age class

codes to make them comparable. Age classes reported here are classified as Young (= hatchlings and young-of-year juveniles) or Adult

(= subadult or adult). The distinction is based on species-specific

phenotypic attributes (e.g., size, coloration, pattern, and secondary

sexual characters) and the appearance of young-of-year.

Results

	 Jones(2009) reported that 19 species have been documented along

the Marijilda Road and adjacent lower Swift Trail, but presently I

consider the total to be 20 (table 1), counting both species of the

Sceloporus magister complex. Elgaria kingii is a cryptic species

that may be present in the riparian area, but has not been detected

from Marijilda Creek, although it has been documented from nearby

Noon Creek (Nickerson and Mays 1969). Interestingly, Holbrookia

Table 1—Lizards of Marijilda study site and adjacent lower Swift Trail. Detections (counts) are for all systematic road-transect surveys in 

2003, 2010, and 2011. Species not detected during surveys were opportunistic or had been documented in the literature. Sceloporus 

magister complex includes S. magister, S. bimaculosus, and presumptive hybrids. See Jones and Lovich (2009) for a discussion of 

taxonomy and nomenclature, which is recognized here.

Standard English Name Scientific Name Code Count Percent

Ornate Tree Lizard Urosaurus ornatus UROR 1560 40

Common Side-blotched Lizard Uta stansburiana UTST 809 21

Desert Spiny Lizard complex Sceloporus magister complex SCMC 611 16

Tiger Whiptail Aspidoscelis tigris ASTI 277 7

Eastern Collared Lizard Crotaphytus collaris CRCO 191 5

Greater Earless Lizard Cophosaurus texanus COTE 187 5

Clark’s Spiny Lizard S. clarkii SCCL 94 2

Desert Grassland Whiptail A. uniparens ASUN 37 1

Regal Horned Lizard Phrynosoma solare PHSO 6 <1

Round-tailed Horned Lizard P. modestum PHMO 3 <1

Long-nosed Leopard Lizard Gambelia wislizenii GAWI 2 <1

Gila Spotted Whiptail A. flagellicauda ASFL 2 <1

Zebra-tailed Lizard Callisaurus draconoides CADR 1 <1

Hybrid horned lizard P. solare × P. modestum PHHY 1 <1

Unidentified lizards UNKN 108 3

Western Banded Gecko Coleonyx variegatus COVA 0 0

Gila Monster Heloderma suspectum HESU 0 0

Texas Horned Lizard P. cornutum PHCO 0 0

Greater Short-horned Lizard P. hernandesi PHHE 0 0

Great Plains Skink Plestiodon obsoletus PLOB 0 0

Yarrow’s Spiny Lizard S. jarrovii SCJA 0 0

Total 20 species 3,889 100
50 USDA Forest Service Proceedings RMRS-P-67. 2013

Jones Preliminary Assessment of Changes in a Lizard Assemblage at an Ecotone in Southeastern Arizona

maculata and Sceloporus undulatus complex seem to be absent, but

they have been documented from the vicinity (Nickerson and Mays

1969).

	 A total of 60 surveys were completed: 8 in 2003 (1 June to 3

August), 12 in 2010 (26 June to 11 December), and 40 in 2011 (all

months). The only complete calendar-year survey was 2011 because

that was the first complete year when long-term monitoring was an

objective. There were a total of 3,889 lizard detections(range 0 - 179

detections per visit, mean = 64.8 sd = 38.2) during the 3 survey years.

Thirteen species were recorded during road tallies(table 1).The most

commonly detected species (at least 5% of detections) were Ornate

TreeLizard(Urosaurus ornatus),CommonSide-blotchedLizard(Uta 

stansburiana), Desert Spiny Lizard complex, Tiger Whiptail (Aspidoscelis tigris), Eastern Collared Lizard (Crotaphytus collaris), and

Greater Earless Lizard (Cophosaurus texanus). Species not detected

during transects were assumed to be rare in the study area, nocturnal,

or cryptic. Some species, such as U. ornatus, U. stansburiana, and

parthenogeneticwhiptails are known to frequent disturbed areas,such

as the road margin (Jones and Lovich 2009; Wright and Lowe 1968).

	 Lizardsweredetectedyear-round,inagenerallybimodaldistribution

reflecting spring and summer (monsoonal) peaks of activity (fig. 1).

In 2011 (the complete year),seasonal activity varied by species. Only

U. stansburiana (especially) and U. ornatus were active during the

winter, and only then onwarmdays(generally in the 60’s). Sceloporus 

clarkii (n = 58) also had a long period of activity (March through

November). Aspidoscelis uniparens (n = 26) had the narrowest peak

of activity, with 65% of the detectionsin July, immediately following

the onset of summer rains. Aspidoscelis tigris was only active during

hot periods,with 88%(n=203) being active June throughAugust.The

other species showed a generally normal curve from spring through

fall, although there was a profound increase in surface activity for

some species with the onset of summer rains. This spike was apparent for A. uniparens, U. ornatus, U. stansburiana, and S. magister

complex. Horned lizards (Phrynosoma solare, P. modestum, and a

hybrid [Jones and Winsor, in press]) also were mostly detected after

the onset of the monsoon (71% [5/7] detections). Although sample

size was low for horned lizards, monsoonal peak activity was noted

in other years and from opportunistic observations on the adjacent

paved Swift Trail. There was no apparent monsoonal spike for A. 

tigris, C. collaris, or C. texanus.

	 A comparison of differences between years had to be limited to

June, July, and August, the only months that were sampled in all 3

years. Figure 2 shows the differences in the relative detections of

common species. There were observed decreases over time (2003

vs. 2010/2011) in C. texanus and C. collaris and an increase in A. 

tigris and U. stansburiana. In fact, U. stansburiana was considered

extremely rare on Marijilda road in the 1960’s despite extensive

searches for it (Nickerson and Mays 1969), but it is now extremely

abundant.

	 Figure 3 shows differences in age-class distribution for common

species between 2010 and 2011. The data clearly show a higher

proportion of Young to Adult categories in 2010 than 2011, with the

exception of U. stansburiana. This observation is consistent with the

expectationsfrom the winter drought pattern between years(e.g.,Vitt

and others, 1978).

	 One of the waysI tracked changesin the lizard assemblage was by

comparing distribution of detections by species along the transect.

Overall, the first mile or so of the transect had the highest detection

rate and presumably the highest density of surface-active lizards

(fig. 4). This pattern was similar in all years. The detection rate

dropped with increasing distance from the start, although there was

an increase in detections at the end of the road, which corresponds

to the Marijilda Creek riparian area. Most of the species were not

evenly distributed. The large number of detections in the first part

of the transect was primarily due to the three most common species,

U. ornatus, U. stansburiana, and S. magister complex. Urosaurus 

ornatus, C. texanus, and S. clarkii were responsible for most observations at the creek. Aspidoscelis tigris (2003-2011) and C. texanus

(2003 only, fig. 5) were fairly evenly distributed along the transect.

Crotaphytus collaris and A. uniparens, were most abundant in the

upper one-half of the transect. Sceloporus clarkii was usually found

in distinct segments. According to data, C. texanus was much more

abundant in 2003, but it is interesting to note that the distribution

along the transect also changed. In 2003, the species was common

and widely distributed, while in 2010 and 2011, there were far fewer

detections and the species was mostly detected in two orthree distinct

road segments (fig. 5).

Figure 1—Mean detections of lizards (Y-axis) by month in 2011, showing 

the bimodal pattern of spring and monsoon peaks of surface activity. In 

2010, there was another peak in late summer and early fall due to the 

emergence of young-of-year.

Figure 2—Mean detections per visit of common species by year for June 

through August, the only months surveyed in all 3 years. Species codes 

are shown in Table 1.
USDA Forest Service Proceedings RMRS-P-67. 2013 51

Preliminary Assessment of Changes in a Lizard Assemblage at an Ecotone in Southeastern Arizona Jones

Figure 3—A coarse comparison of relative reproductive output between 

2010 and 2011 (see text for explanation), June through October. There 

was good winter precipitation in 2009-2010, which translated to a good 

year for annuals and flowering perennials (hence, abundant insect prey), 

whereas 2010-2011 was a dry winter with few annuals (hence, reduced 

insect prey). Decreases in reproductive output due to frequent winter 

drought years could result in population declines and possibly extirpation 

for some species. However, the role of monsoonal moisture and reproductive success is poorly understood. Species codes are shown in Table 1.

Figure 4—Total detections of all species of lizards by road segments for a 

1-year subset in 2010-2011 (n = 1,380 detections). Lizards are recorded 

by 0.1-mi segments, but are combined here in 0.3-mi segments for display. Segment 6 (mile 1.3-1.6) is the area treated for invasive weeds, but 

notice that lizards are not particularly abundant in either side of the area.

Figure 5—Mean detections of Cophosaurus texanus by 0.3-mi segment 

by year. In 2003 this species was the second most abundant species 

(18.4% of detections) and was widespread across the study area. In 2010 

and 2011 there were considerably fewer observations (1.2% and 3.2%, 

respectively) and the lizards were in discrete areas (usually near washes 

and Marijilda Creek riparian area).

Discussion

	 Thisstudy is attempting to answerthreemajor questions:(1) which

lizard species compose the assemblage, (2) how they are distributed

spatially and temporally, and (3) how the assemblage will change

over time. The third part requires a cause-and-effect explanation,

especially as it relates to climate change.

	 For the first question, I reviewed the literature and HerpNet, conducted surveys, and spent many opportunistic hoursin the study area

between2003and2011.Thelizardassemblage shouldbeaccuratenow,

although other rare or cryptic species could be detected in the future.

Jones (2009) reported that the study area has the highest diversity of

lizard species (for such a small area) documented anywhere in the

United States. Other areas of southeastern Arizona or adjacent New

Mexico likelyhave a similarlevelofdiversity, but clearly theMadrean

Archipelago and surrounding valleys have the highest diversity in the

country. The explanation for the high diversity wasreported by Jones

(2009): the area is a convergence zone of physiographic provinces

that each has a high diversity of lizard species.

	 The second question—how lizards are distributed along the

transect—is starting to unfold. Distribution of most species is not

random and patterns are emerging. I am currently engaged in an

intensive habitat characterization study component along the transect

to determine why lizards are distributed as they are. Observations

suggest certain variables will likely help explain abundance, such as

distribution and type ofrock, vegetation community, topography, and

topographic features(e.g., desert approach,foothill,stream).Another

observation is that the segment of road from about milepost 1.3-1.6

has the least number of detections, despite having the best visibility

of the entire transect. This area was treated for invasive weeds and

regrowth is sparse. However, treatment of invasive weeds in a small

area to abate spread may be a very important conservation measure

that is best evaluated at a larger scale.

	 The third question is the crux of the study. The Sinervo and others

(2010) prediction of large-scale lizard declines wasthe impetusto reinitiate surveys. Marijilda was selected because of the high diversity

and apparent high abundance of lizards, and its setting in an ecotone.

Because ecotones are at the edge of vegetation communities, they

are expected to have sensitivity to show change from altered climate

patterns or other factors (Allen and Breshears 1998, Loehle 2000,

Noble 1993).Unfortunately,when the studywasinitiatedin 2003,that

was not a goal, so the 2003 data are limited, and the Nickerson and

Mays (1969) paper was qualitative. However, these provide the only

information we have on the area before 2010. The observations from

then to now are generally consistent with climate change predictions.

Cophosaurus texanus, a foothill species in the Marijilda area, has

gone from being extremely abundant and widespread to uncommon

with a disjunct distribution. Uta stansburiana, a desert species, has

gone from being “extremely rare” (Nickerson and Mays 1969) to

extremely abundant.

	 It is well established in the literature that lizard populations are

known to be naturally dynamic between years (e.g., Dunham 1982;

Fergusen and Fox 1984;Vitt and others, 1978); there aremany factors
52 USDA Forest Service Proceedings RMRS-P-67. 2013

Jones Preliminary Assessment of Changes in a Lizard Assemblage at an Ecotone in Southeastern Arizona

that influence population dynamics, including temperature,moisture,

and resource availability. Nevertheless, the difference seen between

years is a red flag that suggests there may be a changing lizard assemblage concomitantwith the frequent droughts and increasedmean

annual temperature in the past decade. The species that have shown

increases in this study are desert species, while some grassland/

foothill species have been stable or showing decreases. Of course,

there are factors besides climate change that can influence lizard distribution.Another hypothesis for a changing lizard assemblage is the

weed-species hypothesis for all-female whiptails (Wright and Lowe

1968). There are three species (at least) of whiptails (Aspidoscelis)

in the study area, including two parthenogenetic species, but only

the gonochoristic species (A. tigris, a desert associate) is common

and well-distributed along the transect, unlike A. uniparens and A. 

flagellicauda, which are typical of grasslands and woodlands.

	 This study is not limited to understanding lizard biology, as there

are inferred management implications. Lizards may function as a

“canary in a coal mine,” warning of impending vegetation change

or identifying other environmental parameters. Even with increased

temperature and decreased precipitation,some plants may be temporarily resilient (e.g., many seeds or shrubs can endure droughts, then

capitalize on occasional wet years).Tracking vegetation change itself

in an ecotone may be problematic (Noble 1993). Because of natural

history attributes of lizards (e.g., a rapid drop in fecundity due to

drought), it seems logical there will be changes in lizard populations

before dramatic shifts in vegetation are seen. Such an early warning

system can help land managers assess habitat vulnerabilities, so they

can adjust management strategies for programs such as range or fire

management (USFS 2011). Also, there are inherent uncertainties in

modelsthat predictthe outcome of a changing environment on species

assemblages. This study should help support or refute these predictions, refine variables and outputs, identify other factors contributing

to population shifts, and contribute to adaptivemanagement of habitat

for lizards and other organisms.

Acknowledgments

	 First and foremost, I thank Hunter Winsor for his many days of

volunteering to conduct lizard surveys. Mike McCarthy, Eastern

Arizona College, helped me to locate this valuable student resource.

Others who helped in the field include Anne Casey, Roger Joos, and

Betty Phillips. Rick Gerhart and Jennifer Ruyle helped support this

project as “targeted climate change monitoring” for the national

Forest Service climate change program (USFS 2011). Insight and

helpful suggestions for the project came from Southwest Partners in

Amphibian and Reptile Conservation, as well as several herpetologists,includingRoyAverill-Murray,TomJones,DavidGrandmaison,

Phil Rosen, Wade Sherbrooke, and Cecil Schwalbe. Esther Nelson

and Anne Casey reviewed an earlier version of this manuscript.

References

Allen, C.D.; and D.D. Breshears. 1998. Drought-induced shift of a forestwoodland ecotone: Rapid landscape response to climate variation. Proceedings of the National Academy of Sciences, USA 95:14839-14842.

Brown, D.E., ed. 1994. Biotic communities. Salt Lake City: University of

Utah Press. 342 p.

Degenhardt,W.G.;Painter,C.W.; andA.H.Price.1996.Amphibians andreptiles

of New Mexico. University of New Mexico Press. 431 p.

Dunham,A.E.1982.Demographicandlife-historyvariationamongpopulations

of the iguanid lizard Urosaurus ornatus: implications for the study of life

history phenomena in lizards. Herpetologica 38(1):208-221.

Fergusen, G.W.; and S.F. Fox. 1984. Annual variation of survival advantage

of large juvenile side-blotched Lizards, Uta stansburiana: its causes and

evolutionary significance. Evolution 38(2):342-349.

Gibbons, J.W.; Scott, D.E.; Ryan, T.J.; [and others]. 2000. The global decline

of reptiles, déjà vu amphibians. Bioscience 50(8):653-666.

Jones,L.L.C.2009.Lizarddiversity atMarijildaCanyon andlowerSwiftTrail,

Graham County, Arizona. Sonoran Herpetologist 22(9):94-98.

Jones,L.L.C.; andR.E.Lovich, eds. 2009.Lizards oftheAmericanSouthwest.

Tucson: Rio Nuevo Publishers. 567 p.

Jones, L.L.C.; and C.R. Schwalbe. 2008. Desert spiny lizard Sceloporus magister Hollowell, 1854. Pp.226-229. In: Jones, L.L.C.; and R.E. Lovich,

eds. Lizards of the American Southwest. Tucson: Rio Nuevo Publishers.

Jones, L.L.C.; and H. Winsor. In press. Phrynosoma modestum × P. solare

(Round-tailed Horned Lizard × Regal Horned Lizard). Putative hybrid.

Herpetological Review.

Lazaroff,D.W.;Rosen,P.C.; andC.H.Lowe,Jr.2006.Amphibians,reptiles, and

their habitats atSabinoCanyon.Tucson:University ofArizonaPress. 158 p.

Loehle, C. 2000. Forest ecotone response to climate change: sensitivity to

temperature response functional forms. Canadian Journal of Forest Research 30:1632-1645.

Nickerson, M.A.; and C.E. Mays. 1969.Apreliminary herpetofaunal analysis

of the Graham (Pinaleño) Mountain region, Graham Co., Arizona with

ecological comments. Transactions of the Kansas Academy of Science

72:492-505.

Noble, I. R. 1993. A model of the responses of ecotones to climate change.

Ecological Applications 3(3):396-403.

Olson, D.H. 2012. State of the lizard. Year of the Lizard News 1:1, 6-12.

Phelan, R.L.; and B.H. Brattstrom. 1955. Geographic variation in Sceloporus 

magister. Herpetologica 11:1-14.

Sinervo, B.; Méndez-de-la-Cruz, F.; Miles, D.B.; [and others]. 2010. Erosion

of lizard diversity by climate change and altered thermal niches. Science

328(5980):894-899.

USDA Forest Service (USFS). 2011. National roadmap for responding to

climate change.ForestServiceFS-957b.Washington,DC:U.S.Department

of Agriculture, Forest Service.

Vitt, L. J.; Van Loben Sels, R. C.; and R. D. Ohmart. 1978. Lizard reproduction: annual variation and environmental correlates in the iguanid lizard

Urosaurus graciosus. Herpetologica 34(3):241-253.

Wright, J.W.; and C.H. Lowe 1968. Weeds, polyploids, parthenogenesis,

and the geographical and ecological distribution of all-female species of

Cnemidophorus. Copeia 1968(1):128-138.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 53

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

It’s Lonely at the Top: Biodiversity at Risk to Loss from 

Climate Change

John L. Koprowski, Sandra L. Doumas, Melissa J. Merrick, Brittany Oleson, Erin E. Posthumus, 

Timothy G. Jessen, and R. Nathan Gwinn

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—Climate change is a serious immediate and long-term threat to wildlife species. State and federal 

agencies are working with universities and non-government organizations to predict, plan for, and mitigate 

such uncertainties in the future. Endemic species may be particularly at-risk as climate-induced changes 

impact their limited geographic ranges. The Madrean Archipelago is characterized by high levels of endemism, natural fragmentation, and increasingly poor connectivity among the mesic montane islands within 

an arid matrix of low elevation deserts and grasslands. The region already has experienced an increase in 

temperature and this trend is predicted to accelerate over the remainder of the century. We assessed patterns of elevational distribution of reptiles and mammals within the Madrean Archipelago. We examined 

incremental temperature increases and determined how much reptilian and mammalian biodiversity is at-risk 

to be lost from montane islands due to elevational and geographic restriction. We estimate that 15 to 20% 

of all reptile and mammal diversity is at risk of loss by 2100 based solely on predicted patterns of upslope 

migration of biotic communities driven primarily by climate change. With this significant proportion of native 

species, we suggest that this emphasizes the need to continue traditional management actions focused on 

habitat improvement, restoration and connectivity for the majority of species in combination with innovative 

and active management strategies directed at the subset that is most at risk of extirpation.

Introduction 

	 TheMadreanArchipelagois characterized byextremelyhighlevels

of diversity often attributed to the geographic location at the nexus

of the Chihuahuan and Sonoran Deserts and the Rocky Mountains

and Sierra Madre (Gehlbach 1993). Topographic diversity is also

exceptional with a basin and range landscape extending from sea

level to the west and above 3,000 m on numerous peaks that create

a habitat archipelago of >40 montane islands with patterns of diversity explained by island biogeographic theory (Lomolino and others

1989; Patterson 1995) and a lack of Pleistocene glaciation (Brown

and Davis 1995). The Madrean Archipelago is considered to be a

hotspotof evolution (Spector 2002) andbiodiversity (Mittermeier and

others 2005) and been labeled as a region ofmega-diversity (Warshall

1995). For a number of taxa, the region harbors exceptional species

diversity, often notseen atsuch northerly latitudes, especially among

mammals (Turner and others 1995) and reptiles (Stein 2002). 

	 General predictions for climate change suggest that most of the

westernUnitedStates and northwesternMexicowill experiencemean

temperature increases of 2-5 oCchangesinmean temperature by 2100

with increases in the variability of extreme events (Easterling and

others 2008; IPCC 2001). Upslope migration and changes in plant

community composition (Munson and others 2012), divergent fire

regimes(Flanniganandothers2000),novelinvasivespecies(Hellmann

and others 2008), and increased disease and insect infestation (Dale

and others 2001) are likely to occur and, in many cases, have already

been observed. As a result,Arizona and the MadreanArchipelago are

considered to be at medium to extreme risk of biome shifts(Gonzalez

and others 2010)that can have profound risksforfaunal communities

thatincrease the challenges ofmanaging for persistence under climate

change scenarios.

	 Herein, we focus on the patterns of diversity among reptiles and

mammals due to their elevated species richness in Arizona (Stein

2002; Turner and others 1995) and their reduced vagility and greater

difficulty in moving between “sky islands” or other isolated habitats.

Furthermore, we capitalize on the fact that these taxa are relatively

conspicuous as modest to large terrestrial vertebrates and distributional ranges are reasonably well known (Brennan 2008; Brennan

and Holycross 2006; Hoffmeister 1986). We conclude that a subset

of the mammalian and herpetofauna found in high elevation forests

are at elevated risk of extinction at least at the local level with predicted levels of climate change. However, the majority of reptiles

and mammals are in habitats that will likely remain contiguous over

the remainder of the century. We suggest that successful conservation strategies will require innovative approaches to deal with high

elevation speciesin combination with traditional landmanagement to

maintain connectivity and restore degraded habitats for the majority

of species.

Methods

	 We simulated general patterns of climate change using the digitized base map of major biotic provinces by Brown and Lowe (1981)
54 USDA Forest Service Proceedings RMRS-P-67. 2013

Koprowski, Doumas, Merrick, and others It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change

under the assumption that the lapse rate (10 oC for 1000 m in dry air,

6 oC for 1000 m in saturated air) will be the primary determinant of

habitat change across elevation. While simplistic, this appears to be

a reasonable approximation of the major influence on distribution of

biotic communities(McDonald andBrown 1992). We down-sampled

elevation data (1 arc-second resolution DEM; Sugarbaker and Carswell 2011) to ~1.2 km resolution by assigning the mean elevation

of 16 raster cells to the aggregate cell. We manually reclassified

elevation values into 250-m elevational intervals. To simulate the

impacts of predicted warming levels, effective elevation was dropped

(i.e., subtracted from current elevation) in three stages to represent

moderate to severe climate impacts. Breakpoints were defined at 250

m (simulating ~2-2.5 oC change in temperature), 500 m (~4-5 oC

change in temperature), and 750m(~6-7.5 oCchange in temperature).

Using a polygon GIS layer of vegetation communities in Arizona

(The Nature Conservancy 2004), we simulated concomitant upslope

migration of vegetation communities for the southeastern corner of

Arizona by assigning each polygon the most dominant vegetation

type in neighboring, lower-elevation areas.

	 Data on species range and elevation were collected from taxonspecific treatisesfor mammalian (Cockrum 1960; Findley and others

1975; Hoffmeister 1986) and herpetofauna (Stitt and others 2005;

Brennan and Holycross 2006; Brennan 2008). If elevations were

not provided in these sources, we used topographic maps of known

locationsto obtain an approximate elevation. We did not include data

on species associated with aquatic environments (two Thamnophis

snakes,threeKinosternonturtles,threemammals(Castor canadensis, 

Ondatra zibethicus, Lontra canadensis). Furthermore, we conducted

our assessments at the specieslevel and did not addresssubspecies or

otherwise unique population segments. We assessed the elevational

range of each species and habitat affinites from the taxon-specific

treatises and tallied the species at high or low risk of extirpation from

Arizona by 2100 based on projected elevational shifts. For species

at elevations with habitats that were likely to increase in land cover

or remain as dominant biotic communities within Arizona by 2100,

we considered them to be at low risk of extirpation within the State.

For species at elevations with habitats that were likely to decrease

in cover, we assessed the proportion of their range that would be at

risk of loss using an 80% decline as threshold for high risk of loss

by 2100.

Results

	 The area oflandscape acrossArizona at upper elevationsrepresents

a small proportion of the state (table 1) and upper elevations above

2250 m (~7500 ft) account for about 119,000,000 ha. Simulating

temperature increases of 2-2.5 oC (250 m depression in elevation),

4-5 oC (500 m depression in elevation), and 6-7.5 oC (750 m depression in elevation) results in a substantial change in the high elevation

environmental conditionsstatewide (fig.1) andeffectivelyreducesthe

area above 2250 m to < 13,000,000 ha (500 m scenario) by 2100. The

changesin high elevation conditions within theMadreanArchipelago

would result in the expansion of desertscrub and substantial decline

in the coverage of all montane forest types over the next century

(fig. 2). Desertscrub and grasslands would be the dominant biotic

communitiesin the MadreanArchipelago ofArizona by 2100 (fig. 3).

Given the likely increase of low elevation microclimates as well as

desertscrubandgrasslandcommunitiesinArizona,species associated

with these elevations are projected to persist until 2100 in the face of

climate change alone. This represents a majority of reptiles (83.9%,

fig. 4) and mammals (76.3%, fig. 4) with no difference between the

two major taxa (χ2=2.38, df = 2, p > 0.20). The risk of extirpation

associated with loss of habitat due to climate change was high or

unclear for less than 25% of reptiles and mammals.

Discussion

	 Climate change is likely to have significant and rapid impacts on

the Madrean Archipelago (McPherson and Weltzin 2000; Weiss and

Overpeck 2005). High elevation habitatssuch asmany coniferforests

dominatedbyfir(Abies),spruce(Picea),andDouglas-fir(Pseudotsuga)

are likely to decline significantly during the remainder of this century

(Kupfer and others 2005; McPherson andWeltzin 2000). The rugged

and diverse topography of the Madrean Archipelago will lead to the

loss of many montane island-top forests and the initiation of habitat

type conversion in the surrounding basins. Our simple elevational

models support the projections of others that vegetation change, especially at high elevation, is likely to be substantial across Arizona

by 2100.

	 The pace of distributional shifts is difficult to predict due to the

multitude of influences. Current elevational shifts for a variety of

animals suggest an upward elevation movement of 11.0 m per year

at present occurs in areas experiencing the greatest temperature

change, although mammals seem to be slower to move than other

taxa (Chen and others 2011). However, overall species assemblage

change may lag behind temperature change by decades (Menendez

and others 2006). This suggests a lag time in the visibility of change

and so proactive preventative management may be warranted with

the knowledge that such changes are underway.

	 Oneofthegreatest challengesispredictinghowbiotic communities

will move; anticipating movements of species is even more difficult.

Forests worldwide are showing a myriad of predictable and novel

responses to global climate changes (Allen and others 2010). The

Sonoran Desert is likely to expand although predicting the extent

of the expansion is difficult as it will depend greatly on patterns of

precipitation variability and increased frequency of such previously

uncommon disturbances such as fire (Weiss and Overpeck 2005;

Munson and others 2012). Detailed downscaledmodeling efforts will

provide insight to such changes (Garfin and others 2009; Tabor and

Williams 2010). However, variationsin phenology, phenotypic plasticity and other system responses that decouple important ecological

Table 1—Area (ha) of Arizona terrestrial environments 

within 250-m elevation classes that were used as 

the starting point of the elevation models.

Low elevation (m) Area (ha)

0 161404560

250 368010000

500 344429712

750 258707808

1000 293224752

1250 424730448

1500 605162160

1750 551220624

2000 248307696

2250 75723120

2500 31092048

2750 10116288

3000 1589328

3250 339552

3500 46656

3750 2304
USDA Forest Service Proceedings RMRS-P-67. 2013 55

It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change Koprowski, Doumas, Merrick, and others

Figure 1—Elevations of Arizona with color contours at 250 m intervals for present conditions and with elevations decreased by 250 m to simulate 

a 2.0-2.5 oC temperature increase, 500 m to simulate a 4.0-5.0 oC temperature increase, and 750 to simulate a 6- 7.5 oC temperature increase. 

Effective elevation presents elevations with similar colors that would be expected to have similar temperature profiles after adjustment.
56 USDA Forest Service Proceedings RMRS-P-67. 2013

Koprowski, Doumas, Merrick, and others It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change

Figure 2—Distribution of biotic communities (Brown and Lowe 1981) in the Arizona portion of the Madrean Archipelago at present and potential 

distribution after adjusting to simulate a 4.0-5.0 oC temperature change. Communities were adjusted upward in elevation by 500 m.
USDA Forest Service Proceedings RMRS-P-67. 2013 57

It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change Koprowski, Doumas, Merrick, and others

Figure 3—Potential availability of biotic 

communities (Brown and Lowe 1981) in the 

Arizona portion of the Madrean Archipelago 

at present and after adjusting to simulate a 

4.0-5.0 oC temperature change. Communities 

were adjusted upward in elevation by 500 m.

Figure 4—Proportion of mammal and 

reptile species that could lose >80% 

of the habitat within their current 

range in Arizona with a 4.0-5.0 oC 

temperature change simulated that 

would adjust communities upward 

in elevation by ~500 m. 

associations are unpredictable and of concern (Visser andBoth 2005).

For instance, some organisms respond to increasing photoperiod in

their annual cycles whereastemperature change can be the important

cue for others (Bale and others 2002). Phenotypic plasticity and

ecological flexibility in patterns of habitat use and dietary habits will

also be important. We will continue to gain insight into drivers over

time and such processes can clearly lead tomultiple and unpredictable

future stable states (Schneider 2004). Land managers must continue

to act in the present but plan for the future based on this uncertainty.

	 Upper elevation forests are likely to decrease significantly in coverage by the end of the century under any set of projections; relic stands

are likely to experience further degradation due to the plethora of

factors exacerbated by climate change (Hale and others 2001). Our

assessments suggest that 15-25% of the reptile and mammal species

are at risk almost exclusively due to habitat loss driven by climate

change. Ourresults did notinclude subspecies orisolated populations

and these unique population segments should be evaluated further in

the future. The vast majority ofspeciesthat have isolated subspecies

or unique populations are found in high elevations where the factor

that maintains isolation is elevational changes in microclimate and

habitat. However, unique and disjunct population isolates outside the

continuous range of low elevation species may also be identified for

future conservation efforts. Some low elevation species with unique

microhabitat and soil requirements such as the sand dune inhabiting

fringe-toed (Uma) and isolated rocky outcrop dwelling night lizards

(Xantusia) may also be targeted for more active management strategies. We emphasize that while appropriate biotic communities will

remain in substantial amounts withinArizona, the speciesthat inhabit

thesehabitatswill continue torequiremanagementtoconservehabitat

and mitigate current drivers of threats to biodiversity. Unpredictable

responses will also undoubtedly occur and contemporary evolution in

the face of climate change can in theory be significant (Hanson and

others 2012). While we cannot predict the unpredictable, we must

be aware that such events will occur and be vigilant.

	 The enormity of current and potential climate change impacts has

resulted in an effective paralysis in action by many land managers,

scientists, and politicians. Inaction is perhaps the least appropriate

response at this point in time. Our objective was to assess the magnitude of the impacts of habitat loss due to elevational changes in

comparison with latitudinal changes among two of the most diverse
58 USDA Forest Service Proceedings RMRS-P-67. 2013

Koprowski, Doumas, Merrick, and others It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change

yet least vagile vertebrate taxa (reptiles and mammals). Among these

taxa, a minority of species (15-25%) are at high risk of loss from

the fauna of Arizona by the turn of the century due primarily to the

disappearance of high elevation habitats (fig. 4). A similar pattern

was reported for lizards in Mexico; global loss of lizard species is

estimated to be about 20% by 2080 (Sinervo and others 2010). These

are the species upon which we might focus captive breeding, assisted

migration, andassistedtranslocationeffortsinthe shortterm(Richardson and others 2009). Long-term efforts at maintaining connectivity

and quality of habitat as well as habitat restoration using traditional

land management techniques are what we need to continue to do for

the majority of species. However, we must redouble our efforts for

the temporal and spatial scales of change and levels of collaboration

will not enable us to face the challenges ahead at the pace that global

change occurs. 

Acknowledgments

	 We thank Don Swann, Claire Zugmeyer, and Peter Lurz for their

helpful reviews that undoubtedly increased the quality of this manuscript. 

References

Allen, Craig D.; Macalady, Alison K.; Chenchouni, H.; [and others]. 2010.

A global overview of drought and heat-induced tree mortality reveals

emerging climate change risks for forests. Forest Ecology and Management 259: 660–684

Bale, Jeffrey S.; Masters, Gregory J.; Hodkinson, Ian D.; [nd others]. 2002.

Herbivory in global climate change research: direct effects of rising temperature on insect herbivores. Global Change Biology 8: 1–	 16.

Brennan, Thomas C. 2008. On-line field guide to the reptiles and amphibians

of Arizona. http://www.reptilesofaz.org/ (accessed 1 May 2012).

Brennan, Thomas C.; Holycross, Andrew T. 2006. Field guide to the amphibians and reptiles in Arizona. Phoenix: Arizona Department of Fish and

Game. 150 p.

Brown, David E.; Davis, R. S. 1995. One hundred years of vicissitude: Terrestrial bird and mammal distribution changes in theAmerican Southwest,

1890-1990.In:Biodiversity andManagement oftheMadreanArchipelago:

the Sky Islands of Southwestern United States and Northwestern Mexico.

Gen. Tech. Rep. GTR-RM-264. Fort Collins, CO: U.S. Department of

Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 669 p.

Brown, David. E.; Gonzalez, C. A. L. 2000. Notes on the occurrences of

jaguarsinArizona and New Mexico. Southwestern Naturalist 45: 537-542.

Cockrum, E. Lendell. 1960. The recent mammals of Arizona: their taxonomy

and distribution. Tucson: University of Arizona Press.

Dale,VirginiaH.;Joyce,LindaA.;McNulty,Steve;[andothers].2001.Climate

Change and forest disturbances. BioScience 51: 723-734.

Easterling, D.R.; Meehl, G.A.; Parmesan, C.; Changon, S.A.; Karl, T.R.;

Mearns,L.O.2008.Climate extremes: observations, modeling, and impacts. 

Science 289: 2068–2074.

Findley, James S.; Harris, A. H.; Wilson, D. S.; Jones, C. 1975. Mammals of

New Mexico. Albuquerque: University of New Mexico Press.

Flannigan, M.D.; Stocks, B.J.; Wotton, B.M. 2000. Climate change and forest

fires. Science and the Total Environment 262: 221-229.

Garfin,GreggM.;Eischeid,JonK.,Lenart,Melanie[andothers]. 2009.Downscaling climate projectionsto model ecological change on topographically

diverse landscapes of the arid southwestern United States. In: Charles Van

Riper III, Brian F. Wakeling, and Thomas D. Sisk (eds.), Colorado Plateau

IV: Shaping conservation through science and management. Tucson: University of Arizona Press. 355 p.

Gehlbach, Frederick R. 1993. Mountain islands and desert seas. College Station: Texas A&M University. 326 p.

Gonzalez,Patrick;Neilson,RonaldP.;Lenihan,JamesM.;Drapek,RaymondJ.

2010. Global patternsin the vulnerability of ecosystemsto vegetation shifts

due to climate change. Global Ecology and Biogeography 19: 755–768.

Hansen, Michael M.; Olivieri, Isabelle; Waller, Donald M.; Nielson, Einar

E.2012. Monitoring adaptive genetic responses to environmental change. 

Molecular Ecology 21: 1311-1329.

Hellman, Jessica J.; Byers, James E.; Bierwagen, Britta G.; Dukes, Jeffrey S.

2008. Five potential consequences of climate change for invasive species.

Conservation Biology 22: 534–543.

Hoffmeister, Donald F. 1986. Mammals of Arizona. Tucson: University of

Arizona Press. 602 p.

IPCC. 2001. Climate Change 2001: The Scientific Basis: Contribution of 

Working Group I to the Third Assessment Report of the Intergovernmental 

Panel on Climate Change. Cambridge: Cambridge University Press.

Kupfer,JohnA.;Balmat,Jeff;Smith,JacquelineL. 2005.Shiftsin the potential

distribution of Sky Island plant communitiesin response to climate change.

In: Gottfried, Gerald J.; Gebow,Brooke S.;Eskew,Lane G.; andEdminster,

Carleton B., compilers. Connecting mountain islands and desert seas: biodiversity andmanagement oftheMadreanArchipelago II. 2004May 11-15;

Tucson,AZ. Proceedings RMRS-P-36. Fort Collins, CO: U.S. Department

of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.

Lomolino, Mark V.; Brown, James H.; Davis, Russell. 1989. Island biogeography of montane forest mammals in the American Southwest. Ecology

70: 180-194.

McDonald, Kelly A.; Brown, James H. 1992. Using montane mammals to

model extinctions due to climate change.ConservationBiology 6:409-415.

McPherson, Guy R.; Weltzin, Jake F. 2000. Disturbance and climate change

in United States/Mexico borderland plant communities: a state-of-theknowledge review. Gen. Tech. Rep. RMRS-GTR-50. Fort Collins, CO:

U.S. Department ofAgriculture, Forest Service,Rocky MountainResearch

Station. 24 p.

Menéndez, Rosa; Gonzalez-Megias, Adela; Hill, Jane K.; [and others]. 2006.

Species richness changes lag behind climate change. Proceedings of the

Royal Society B: Biological Sciences 273: 1465-1470.

Mittermeier,RussellA.;Robles-Gil,Patricio;Hoffman,Michael;[and others].

2005. Hotspotsrevisited: Earth’s biologically richest and most endangered

terrestrial ecoregions. Arlington VA: Conservation International. 392 p.

Munson, Seth M.; Webb, Robert H.; Belnap, Jayne; [and others]. 2012.

Forecasting climate change impactsto plant community composition in the

Sonoran Desert region. Global Change Biology 18: 1083-1095.

The Nature Conservancy. 2004. Biotic communities of the Southwest (Brown

and Lowe Vegetation 1981). The Nature Conservancy, Center for Science

and Public Policy. Online: http://azconservation.org/downloads/category/

southwest_regional/

Patterson, Bruce D. 1995. Local extinctions and the biogeographic dynamics

of boreal mammals in the Southwest. In: Storm over a mountain island:

conservation biology and the Mt. Graham affair. Istock, C. A.; Hoffmann,

R. S., eds. Tucson: University of Arizona Press: 151-176.

Richardson,DavidM.;Hellman,Jessica J.;McLachlan,Jason S.;[and others].

2009. Multidimensional evaluation of managed relocation. Proceedings of

the National Academy of Sciences 106: 9721-9724.

Schneider, Stephen H. 2004.Abrupt non-linear climate change, irreversibility

and surprise. Global Environmental Change 14: 245–258

Sinervo, Barry; Mendez de la Cruz, Fausto; Miles, Donald B. [and others].

2010. Erosion of lizard diversity by climate change and altered thermal

niches. Science 328:894-899.

Spector, S. 2002. Biogeographic crossroads as priority areas for biodiversity

conservation. Conservation Biology 16: 1480-1487.

Stein,B. 2002. States ofthe Union:rankingAmerica’s biodiversity.Arlington,

VA: Natureserve.

Sugarbaker, L.J.; Carswell, W.J., Jr. 2011. The national map: U.S. Geological

Survey.FactSheet2011–3042.DigitalElevationModeldata at1arc-second

resolution. Online: http://viewer.nationalmap.gov/viewer/ [Accessed 26

Apr 2011].

Tabor, Karyn; Williams, John W. 2010. Globally downscaled climate projections for assessing the conservation impacts of climate change. Ecological

Applications 20: 554–565.
USDA Forest Service Proceedings RMRS-P-67. 2013 59

It’s Lonely at the Top: Biodiversity at Risk to Loss from Climate Change Koprowski, Doumas, Merrick, and others

Turner, Dale S.; Brandes, S.; Fishbein, M.; Hirt, P.W. 1995. Preserve design

for maintaining biodiversity in the Sky Island Region. In: Biodiversity and

management ofthe MadreanArchipelago: the Sky Islands of Southwestern

United States and Northwestern Mexico. Gen. Tech. Rep. GTR-RM-264.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Forest and Range Experiment Station. 669 p.

Visser, Marcel E.; Both, Christiaan. 2005. Shifts in phenology due to global

climate change: The need for a yardstick. Proceedings of the Royal Society

B: Biological Sciences 272: 2561-2569.

Warshall,Peter.1995.TheMadreansky-islandarchipelago.In:Biodiversityand

management oftheMadreanArchipelago:TheSky Islands ofSouthwestern

United States and Northwestern Mexico. Gen. Tech. Rep. GTR-RM-264.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Forest and Range Experiment Station. 669 p.

Weiss, Jeremy L.; Overpeck, Jonathan T. 2005. Is the Sonoran Desert losing 

its cool? Global Change Biology 11: 2065–2077.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
60 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Responding to Climate Change Impacts in the Sky Island 

Region: From Planning to Action

Louise W. Misztal

Sky Island Alliance, Tucson, Arizona

Gregg Garfin

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Lara Hansen

EcoAdapt, Bainbridge Island, Washington

Abstract—Addressing the increasing effects of climate change on natural resources requires multiple organizations, agencies, and institutions working cooperatively to incorporate climate change into resource 

management. In the Sky Island region of the southwestern United States and northern Mexico, Sky Island 

Alliance, a non-governmental organization, has convened a series of climate change adaptation workshops 

in cooperation with a variety of partners. This paper describes a process and methodology for bringing 

together federal and state agencies, local governments, non-profit organizations, tribal representatives, 

private landowners, and academic researchers in order to develop, on-the-ground and policy-level actions 

through climate change adaptation planning. Key outcomes of the workshops include: identification of 

climate change threats to and vulnerabilities of Madrean forest, riparian, desert, and grassland ecosystems 

in the Sky Island region; analysis of the effects of those changes in the region (both direct and indirect) and 

interacting factors; a list of ecosystem specific adaptation options for the region; a plan for implementation 

of an adaptation strategy; and development of a regional network of professionals working cooperatively to 

improve natural resource management under changing conditions. This paper highlights one approach for 

addressing the management and conservation challenges posed by climate change through collaborative 

engagement at a regional scale.

Introduction

	 The global oceans and atmosphere have changed due to human

activities, resulting in a warmer and moister atmosphere (Trenberth

2012).As a result, the southwestern United Statesis among the fastest

warming regions in the nation (Karl and others 2009). In the past 10

years, parts of the Southwest have warmed more than 2 °F relative

to average 20th century temperatures (fig. 1). Nestled in the heart of

this rapid warming is the Sky Island region of southeastern Arizona,

southwestNewMexico, andnorthernMexico.SkyIslands are isolated,

forest-topped, mountain ranges, surrounded by lowland desert and

grasslands. Characterized by steep elevation gradients, commonly

from 600 m at their bases to 3,000 m at their summits, they span the

gap between the Sierra Madre in Mexico and the Rocky Mountains

and overlap the boundary between the Sonoran and Chihuahuan

desert (fig. 2). They harbor some of the most biologically diverse

ecosystems in North America.

	 Effects associated with observed regional warming include a decrease in the fraction of winter precipitation falling as snow, less soil

moisture, changes in timing of species’life cycle events, widespread

vegetationmortality,andincreasedfrequencyoflargewildfires(Robles

and Enquist 2010). Temperature increasesinteract with other factors,

suchasdecadal-scaledrought,landuse andlandcover changes,habitat

fragmentation, and complex ecosystem interactions.

	 In the U.S. portion of the Sky Island region, land tenure is a patchwork, with approximately 34 percent managed by federal agencies,

30 percent by state agencies, 27 percent in private ownership, and

smaller portions managed by Native nations, local jurisdictions,

and conservation interests. Although plans and mandates exist for

considering climate change, individual management agencies and

private entities in the region are at different stages of incorporating

it into management. Many questionsremain about how to implement

adaptation strategies at the local level across different land management boundaries. Moreover there is a constantly expanding body of

scientific information, yet it is still a challenge for natural resource

managersto accessscience useful for planning and decision-making.

In this context, work to establish cross-jurisdictional and regional

coordination and to foster knowledge exchange within and across an

international border is essential for building the institutional adaptive

capacity needed to lessen the potential impacts of climate change

(Hansen and Hoffman 2010).

	 Addressing these myriad challenges is the goal of Adapting to a 

Changing Climate in the Sky Island Region, a project initiated by
USDA Forest Service Proceedings RMRS-P-67. 2013 61

Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action Misztal, Garfin, and Hansen

Figure 1—Composite temperature anomalies (F) Jan to Dec 2000 to 2009 

Versus 1895-2000 longterm average.

Figure 2—Map of Sky Island Region.
62 USDA Forest Service Proceedings RMRS-P-67. 2013

Misztal, Garfin, and Hansen Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action

Sky Island Alliance (SIA) in 2009, the principle elements of which

include a series of regionally focused climate change adaptation

workshops. Objectives of the three-part workshop series include (1)

develop and implement on-the-ground and policy-level adaptation

strategiesthat addresskeyecosystemmanagementvulnerabilities, and

(2) integrate climate change information into participants’ planning

and work. To support these objectives, a regional knowledge-action

network of professionals (e.g., Jacobs and others 2010), working cooperatively to improve natural resource management under changing

conditions, was created. The first two workshops in the series were

convened in 2010 and 2011 and the third workshop will be convened

in 2013. Participants at the first two workshops included personnel

from federal, state, and local agencies; non-governmental organizations; universities; and Native nations and private landowners (table

1). Results include development and implementation of adaptation

strategiesthatspanU.S jurisdictions.This paper describesthe process

used to develop and convene workshops, key workshop results, and

status of post-workshop implementation.

Process and Methodology

	 Climate change adaptation for natural systems can be defined as a

dynamic management strategy that involves identifying, preparing

for, and responding to predicted climate change in order to promote

systemresilience,maintainsystemfunction, andprovide thenecessary

elementsto support biodiversity, human communities and sustainable

ecosystem services (Theoharides 2009). To support development of

climate change adaptation strategies for the Sky Island region, we

worked with partners to develop and convene two workshops of a

three-partseries.The serieswasdesignedtoinvolve the same agencies

and individuals to provide continuity and allow for increasing depth

of involvement with each successive workshop and to be of mutual

benefit across jurisdictions and management mandates.

	 Before the first workshop, we surveyed selected natural resource

managers(table 2) to assess how potential participants gather and use

climate information, and to learn about their current work on climate

change adaptation, and what they see as the most pressing regional

climate change threats and vulnerabilities including barriers to and

needsforreducingvulnerabilities.Inthis context,reducingvulnerability means reducing exposure and sensitivity, and increasing adaptive

capacity. The survey was developed by SIAwith the ClimateAssessmentfortheSouthwest(CLIMAS),Ecoadapt andtheU.S.Institute for

Environmental Conflict Resolution. The survey allowed usto initiate

contact with potential workshop participants and gauge interest in our

workshops and an Arizona Climate Change Network—our concept

for a forum for communication with colleagues about adaptation and

cooperative work. Of 44 question respondents, 85% indicated strong

interest in both attending a climate change adaptation workshop and

joining the Network. Respondents identified the following greatest

threatsin the region: waterscarcity and drought, human pressures on

ecosystems, invasive and non-native species, and fire. The greatest

management needsincluded stable funding, a framework for dealing

with uncertainty, translation ofscience, and effective communication

among colleagues, partners and stakeholders.

	 In addition to our survey-development partners, we worked with

the University ofArizona School of Natural Resources and the Environment and Institute of the Environment, Sonoran Joint Venture, the

U.S. Bureau of Reclamation, and the U.S. Fish and Wildlife Service

through the Desert Landscape Conservation Cooperative (LCC) to

develop workshops that foster movement from climate change planning to action. A 2-day structure was created to deliver regionally

relevant climate science and adaptation case studies (information

push),followedbyaninteractivebreakoutgroupprocess(information

exchange). The science delivery component addressed threats and

greatest current needs identified in survey responses, while breakout

groups addressed the need for better communication and coordination

between jurisdictions within agencies and among different agencies

and organizations in the region.

	 Weco-convenedWorkshop1,Climate Change in the Arid Southwest

(September 2010), with the newly formed Desert Landscape Conservation Cooperative (LCC), a public-private partnership providing

scientific andtechnicalsupport andcoordinationtoresourcemanagers

to address climate change and other landscape-scale stressors. Although the area encompassed by the Desert LCC (Mohave, Sonoran

and Chihuahuan Deserts) is much larger than the Sky Island region,

it had similar landscape-scale objectives and an overlapping group

of participants. The first half-day of the workshop was dedicated to

presenting region-specific information on projected climate changes,

fire, water, wildlife range shifts, adaptation efforts, pre-workshop

survey results, and background about the Desert LCC. Participants

were pre-assigned into three facilitated breakout groups to address

the following vulnerabilities and needs: water scarcity, species and

habitat conservation, and research and monitoring. Each group included a diverse mix of disciplines, organizations, and management

jurisdictions. The groups discussed the following series of questions:

1. What is your management goal?

2. How might climate change affect your goal?

3. How might it affect your existing strategies and methods?

4. Brainstorm actions that can be taken to reduce vulnerability.

Each group then chose two vulnerabilities for in-depth discussion of

adaptation strategies.

	 InWorkshop 2,Between a Rock and a Hot Place (April 2011)—coconvened with EcoAdapt—participants were assigned to ecosystemspecific facilitated breakout groups as follows: Madrean forest,

semi-desert grassland, desert, and riparian. The first half-day of the

workshopwasdedicatedtopresentinginformationon(a)likelyclimate

changes in the region, (b) how those changes may affect hydrology,

Table 1—Climate change adaptation workshop 

attendees by affiliation.

Workshop 1 Workshop 2

Federal 31 (35%) 24 (35%)

NGO 17 (20%) 22 (32%)

State 7 (8%) 2 (3%)

University 15 (17%) 10 (15%)

Other 17 (20%) 10 (15%)

Total 87 68

Repeat attendees 34 (50%)

Table 2—Affiliation of climate change 

adaptation survey respondents

Government agency 18 (33%)

NGO 17 (32%)

University 15(28%)

Other 4 (7%)

Total 54
USDA Forest Service Proceedings RMRS-P-67. 2013 63

Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action Misztal, Garfin, and Hansen

fire,invasivespecies,andconnectivityandcorridors,(c)vulnerabilities

ofspeciesin the region, (d) a framework for dealing with uncertainty,

and (e) case studies ofmanagersincorporating climate change considerationsinto currentwork.We presented a framework for dealingwith

climate uncertainty in the same manner that managers and planners

deal with other uncertainties. Informalscenario planning was used to

consider the range of possible futures by using the models that best

capture climate processes in the region of interest, noting areas of

agreement while also considering extreme but plausible projections

to give a sense of the potential range and direction of change. We

shared an adaptive management example that specifies key uncertainties and research needed to address them, triggers for action, and

necessary science and institutional structures. For the remainder of

the workshop, participants developed preliminary adaptation plans

in ecosystem-specific breakout groups. Groups worked through the

following activities: (1) identify a specific management effort for

adaptation planning and prioritize a common goal, (2) determine

vulnerabilities of your goal to climate change, (3) identify a suite

of potential adaptation responses, and (4) create a set of adaptation

actions and next steps. They then developed hypotheses of change

by answering the following questions:

1. How might climate change affect your common goal or ecosystem directly?

2. How might it affect them indirectly (e.g., ecological effects,

interactions with existing stressors)?

3. How might changes outside your ecosystem influence your

common goal or ecosystem?

4. Which interacting factors influence vulnerability to climate

change (e.g. other physical stressors)?

	 At the end of the first day, there was an opportunity for each

breakout group to exchange information with other groups by sharing

their progress in a marketplace of ideas about key vulnerabilities

and means of addressing them. We structured the second day to

facilitate participants’ discussion of interactions across ecosystems,

landscapes, and stressors to ensure that each breakout group thought

aboutwaysinwhich different ecosystems and strategiesinfluence one

another.We did thisto prevent groupsfrom developing “maladaptive

strategies”—i.e., strategies that addressed issues pertaining to their

ecosystem, but which might adversely affect adjacent ecosystems.

For the concluding session of the workshop, each breakout group

presented summaries oftheir ecosystem goal, top five vulnerabilities,

and a fully formed plan for implementing one adaptation strategy.

Each plan included actions to make the ecosystem less vulnerable,

identification of resources to bring the plan to fruition (e.g. data,

skills, funding, materials, infrastructure, permits), identification of

partners with important resources or involvement necessary for plan

implementation, a timeline, and actions for monitoring success. The

workshop process is summarized in figure 3.

Key Outcomes

	 The chief outcomes of the first two workshops included enhanced

awareness of Sky Island climate change issues, an improved social network for communication and coordination, cross-agency/

cross-jurisdictionaldiscussionandcommonagreementonimpacts and

adaptation options. Workshop participants identified (a) elements of

climate change likely to occur in the region; (b) the effects of those

changes in the region (both direct and indirect) and vulnerabilities

these effects may cause; (c) non-climate stressors, interactions

between climate and non-climate factors, and interactions between

distinct ecosystems; and (d) prioritized initial adaptation strategies

(table 3) (reports available at www.skyislandalliance.org/adaptationworkshops.htm).

	 Themain climate threatsidentified by each group included increasing temperatures; precipitation factors, including amount, seasonal

timing, and intensity; and interactive effects on exposure to climate

through drought.The impact ofthe timing ofseasonal precipitation is

ofconcernbecauseoftheimplicationsofanextendedpre-monsoonarid

fore-summer season: increased fire risk in all ecosystems, decreased

connectivity in stream reaches, decreased dissolved oxygen in bodies of water and riparian pools, increased grass mortality, impacts on

migrating birds that rely on riparian vegetation, and shifts in species

composition. Participants highlighted the threat of “megadrought”

to which the Southwest—which is at the fringe of both winter and

summer moisture-bearing atmospheric circulation patterns and at the

center of strong interior West temperature increases—is particularly

sensitive. It was noted that megadrought could be a game changer

for ecosystems, due to the potential for rapidly crossing ecosystem

thresholds to new quasi-equilibrium states, such as from pine-oak

forests to oak scrub woodlands, or from productive Chihuahuan

Desert grasslands to semi-woody shrublands.

	 We note that uncertainties in climate change projections did not

impede discussions about climate change effects and identification

of adaptation strategies. Participants did identify critical uncertainties about what to monitor, and the necessary frequency and timing

of monitoring. Through thoughtful discussion, participants identified

actionable adaptation strategies (table 3) that build on existing

management, restoration, and public education priorities, and—with

sufficient resources—can be implemented in the short-term using

familiar management tools. These are “win-win, no regrets” options

for Sky Island ecosystem management.

	 In this brief paper, we cannot give an exhaustive accounting of

the nuanced articulations of vulnerabilities, impacts, and adaptation options for each of the four Sky Island ecosystems; however,

the following overview touches on important common factors and

gives examples of some unique challenges. In a nutshell, Sky Island

ecosystem vulnerabilities center on maintenance of ecosystem health

Figure 3—Flow chart of workshop process.
64 USDA Forest Service Proceedings RMRS-P-67. 2013

Misztal, Garfin, and Hansen Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action

and function, keystone ecological processes(e.g., fire),speciesshifts,

erosion, habitat fragmentation, and maintaining traditional economic

land-use practices (e.g., grazing). It is interesting to note that discussions of low elevation ecosystems (grassland and desert) raised concerns about the human community’s indifference to climate change.

Development of alternative land uses(e.g., energy development) was

identified as increasing the vulnerability of these ecosystems and

as a potential impediment to development of sufficient biological

adaptive capacity. Certainly desert and riparian environments have

greater exposure tohumanactivities,suchasdevelopment,recreation,

immigration, and infrastructure.

	 Participants’ articulations of direct and indirect impacts and

adaptation options accounted for much of the workshop activity.

The following example from the Madrean Forest group illustrates

a workshop output. The group identified more than a dozen unique

direct and indirect impacts and interactive factors, related to each of

four climate threats (increased temperature, increased frequency of

warm/dry winters, increased summer precipitation variability, and

megadrought).Asample cascade ofimpacts and interactionsfollows:

Direct impacts: increased frequency of warm/dry winters leads

to altered snow hydrology (more winter rain), diminished watershed

moisture retention, increased insect invasions, increased fire risk, and

vegetation shifts that favor woody species

Indirect impacts: altered phenology of aquatic at springs and

increased grazing pressures

Interacting factors: policy shifts to address lack of precipitation

through cloud seeding

Table 3—Key workshop outcomes.

Madrean Forest Riparian Areas Semi-Desert Grassland Desert

Threats

átemperatures; áfrequency 

of warmer 

and drier winters;

ásummer precipitation 

variability and mega 

droughts

átemperature; 

áfrequency of warmer 

and drier winters;

hotter and longer 

foresummers; changes 

in monsoon season 

precipitation

ádry winters; 

átemperatures; 

ávariability in 

precipitation events, 

changes in seasonality

átemperatures; changes 

in hydrology

Vulnerabilities

Forest health and 

function;

áfire risk;

shifts in wildlife and 

vegetation;

loss of soil and potential 

for forest regeneration; 

áinsect infestations

Habitat fragmentation; 

âbiodiversity; alterations in 

physical processes, stream 

morphology, and 

water table; 

ârecharge; 

âecosystem services

Altered or diminished land 

use practices;

flossing;

âviable ranching;

soil loss & erosion;

lack of community 

concern for climate 

change effects on 

grasslands

Public disconnect with 

climate change impacts;

changes in land pressure;

áwater use, 

átemperature and energy use:

áinvasive species

Adaptation Options

Manage for resilience on a 

landscape scale; 

Manage human uses of 

public lands; 

Focus resources on 

maintaining, and 

protecting resilient areas; 

Protect corridors for 

species connectivity; 

Close sensitive areas to 

prevent further disturbance; 

Plan for beetle detection 

and treatment

Capitalize on drought to 

reduce invasive species; 

Promote restoration, work 

with planners to build and 

design infrastructure that 

helps maintain ecosystem 

processes; 

Pursue different water 

policies

Show communities 

alternative futures (climatic/

landscape changes); 

Incorporate past water &land 

allocation information, with 

potential climate changes 

into future management; 

Harness mass flooding 

events for water reserves, 

âchannelization;

Change grazing time and 

location; and 

Install stabilizing features

Conduct a climate change 

education and awareness 

campaign; 

Engage public through citizen 

science projects, Harvest 

rainwater, 

Increase public transportation 

and bike lanes, 

Increase energy efficiency;

Develop awareness campaign to 

explain relationship between fire 

and invasive species

Selected 

Adaptation Goal

Maintain ecosystem 

services and function 

of montane forests and 

woodlands to preserve 

biodiversity and adaptation 

potential where possible, or 

facilitate transition

Conserve the function and 

integrity of riparian systems 

in a changing climate for the 

Upper San Pedro basin

To maintain and restore 

grasslands and the species 

in them through community 

empowerment and 

engagement

Reduce human impact on desert 

ecosystems by awareness and 

outreach

Adaptation 

Strategy

Initiate a process to manage 

the Sky Island region at a 

landscape scale through 

the National Environmental 

Policy Act (NEPA) 

“Stop the Stupid, Start the 

Smart” outreach campaign 

that places the value of water 

and riparian systems in terms 

that all different groups of 

people can understand

Work together to cultivate 

resilient, native seed sources 

to prepare for likely flooding 

and soil loss associated with 

climate change impacts in the 

region

Incorporate climate change 

into the Saguaro National Park 

BioBlitz event of 2011, specifically 

into the Biodiversity University
USDA Forest Service Proceedings RMRS-P-67. 2013 65

Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action Misztal, Garfin, and Hansen

	 To address this set of issues, which participants classified as

“decreased forest health and ecosystem services,” six options were

identified: (1) transition from project-specific planning and implementation of National Environmental Policy Act (NEPA) requirements to landscape level planning and implementation, (2) evaluate

ecosystem function to prioritize management strategies, (3) restore

fire appropriate regimes, (4) shift fire damage liabilities, (5) manage

invasive grasses, and (6) protect resilient areas. Adjusting the scope

of NEPAassessments wasidentified asthe highest priority adaptation

strategy, and a feasible plan was laid out (table 4).

	 A “marketplace of ideas” session, in which participants were able

to interact between breakout groups, yielded insights on cross-cutting

issues and interactions between ecosystems. Many ideas echoed

those raised during the first workshop, but in the more action-focused

context of the second workshop, they were honed and articulated

more specifically. For instance, riparian areas, which exist in each

of the ecosystems, are affected by watershed uplands. Disturbance to

uplands and upstream reaches,such as wildland fire and water extraction, affect downstream reaches through erosion, sedimentation, and

decreases in the number and extent of perennial segments. Another

theme common to all ecosystems was intervention of non-climate

stressors, such as water law and policy, land use and development,

energy and mining policy, recreation and tourism. These stressors

can increase exposure to climate changes by fragmenting habitat

and reducing connectivity (as in the case of alternative energy development), or increasing fire area and intensity (as in the case of

expanding wildland-urban interface). Non-climate factors are also

important determinants of biological adaptive capacity. Participants

identified hindrancesto developing adequate institutional capacity to

prepare for and respond to climate changes, including (1) attitudes of

the public and key decision makers toward climate change; (2) lack

of coordination and information exchange between isolated effortsto

restore ecosystemfunction and/or prepare for climate change;(3)lack

of consistency in data collection, coupled with a lack of coordination

in sharing data; (4) lack of staffing, resources, and expertise to plan

for and implement experimental treatments and initiatives; and (5)

ineffective water and land use laws that impede efforts to enhance

ecosystem resilience.

Implementation

	 Other outcomes from the first two workshops include the incorporation of climate change considerations into planning and project

development andamore coordinatedapproachtopreparingfor climate

change and restoring ecosystem function.There are a variety of adaptation planning and implementation processes currently underway in

the region (table 5). Our workshops and theArizona Climate Change

Networkhave createda regionalnexusforinformationsharing,project

planning, and cooperative implementation that did not previously exist. For example, funding wassecured to complete a Spring and Seep 

Inventory, Assessment, and Management Planning Project to gather

dataonbiological, hydrological, geomorphological, andmanagement

status of springs and seeps in the Sky Island region. Information will

then be applied to the management of sensitive and invasive aquatic

species, the prioritization of restoration and conservation monies,

and management of wildlife that rely on surface water. The project

is being implemented by SIA in coordination with regional resource

managers, including Pima County, the U.S. Forest Service, and the

Spring Stewardship Institute and is receiving funding through the

Desert LCC. The project seeks to reduce the vulnerability of water

resources and species of concern by developing in-depth knowledge

of regional spring resources, fostering cross-jurisdictional management of those resources, and prioritizing where to focus restoration

and protection (table 6).

Conclusions and Next Steps

	 Our workshops have filled a void of regional coordination and

communication about climate change effects and management responses.Assessing natural resource managers’needs and knowledge

before developing the workshops ensured effective engagement and

a focus on the issues of highest importance to participants. We took

a unique approach, focusing adaptation planning efforts on the entire

Sky Island region rather than individual management units. This approach was beneficial in that it generated discussion and subsequent

coordination acrossjurisdictions andmanagementtypes, and resulted

in identifying key common vulnerabilities and resource issues across

Table 4—Madrean forest adaptation plan.

Goal Maintain ecosystem services and function of montane forests and woodlands to preserve biodiversity and adaptation 

potential where possible, or facilitate transition.

Strategy: Manage the Sky Island region at a landscape scale by implementing a programmatic National Environmental Policy 

Act (NEPA) analysis.

Conduct a landscape vulnerability assessment, define project areas, and use a facilitated process with a conflict 

resolution expert. 

Rationale: Consider landscape scale instead of single places in specific jurisdictions. 

Develop a programmatic NEPA analysis to manage for resilience. 

The FireScape model could inform management at a landscape scale. 

Resources 

Needed:

Working group for coordination: State agencies, Government agencies, Counties, Non-profit organizations, such as 

Sky Island Alliance, Land Conservation Trusts, Tribes, Sonoran Joint Venture, Universities, such as the University of 

Arizona, State forestry groups, Private land owners

Lead: U.S. Forest Service

6-9 months:

6-24 months:

24-36 months:

Define and assess landscape using FireScape tool, identify resources, build partnerships.

Organize and engage working group to develop landscape NEPA analysis. 

Draft programmatic NEPA analysis, stakeholder outreach, implement public comments, and prepare final. 

Monitor Success: Track the timeline and process in getting the NEPA decision adopted in the region. Track implementation 

and revise if necessary.
66 USDA Forest Service Proceedings RMRS-P-67. 2013

Misztal, Garfin, and Hansen Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action

the region. It also afforded managers a sense of how their activities

mayaffectneighboring resources.Conveningworkshops andcreating

a knowledge-action network has fostered ongoing sharing of project

work, information and expertise. One year after the 2011 workshop,

it is clear that adaptation project implementation is most successful

when there is a dedicated lead organization with time and resources

to advance the project. Continually integrating emerging science on

regional climate change and its impacts into management planning

and project implementation will be an ongoing challenge.

	 The next steps for this initiative include: a final workshop in 2013

to share updates on project implementation from the previous two

workshops, further development and expansion of the Arizona Climate Change Network to the entire Sky Island region, and further

implementation of adaptation projects, and incorporation of climate

change information into regional management.

Table 5—Related adaptation efforts.

Adaptation Efforts Lead Organization

Sonoran Desert Conservation Plan Pima County

Climate Change and Natural Resources in Pima County Pima County

Las Cienegas National Conservation Area Climate Change

 Adaptation and Scenario Planning Bureau of Land Management and The Nature Conservancy 

Greater Southlands Habitat Conservation Plan City of Tucson Office of Conservation and Sustainable

 Development

Cuenca Los Ojos –Restoration Cuenca Los Ojos

Firescape University of Arizona and Coronado National Forest

Coronado National Forest Land and Resource Management Plan Revision U.S. Forest Service Rocky Mountain Research Station 

 and Coronado National Forest

Sky Island Region Climate Change Adaptation Workshop Series Sky Island Alliance

Sky Island Spring and Seep Inventory, Assessment and Management 

 Planning Project Sky Island Alliance

Table 6—Implementation case study.

Spring and Seep Inventory, Assessment, and Management Planning

Threats átemperatures, á aridity, á scarcity of water that supports wildlife and biological diversity

Vulnerabilities Lack of data on condition of springs/seeps, alteration of springs/seeps for human uses, likely inability of 

 managers to maintain water where it currently exists

Adaptation 

Strategy Conduct field-based assessment of spring/seep condition, species present, water quality and quantity, solar 

 exposure and human alteration; identify appropriate restoration and protection activities

Project Partners Lead- Sky Island Alliance, Spring Stewardship Institute, Pima County, Pima Association of Governments, Arizona 

 Game and Fish Department, The National Park Service, The Nature Conservancy, U.S. Bureau of Land 

 Management, U.S. Fish and Wildlife Service, Ft. Huachuca, Coronado National Forest, U.S. Geological Survey,

 Arizona Water Resources Research Center, Desert LCC

Implementation Activities

Nov 2011 – May 2012 Determine areas of high management priority for conducting assessments with project partners

Apr 2012 Train volunteers and agency personnel in spring/seep assessment protocols

May 2012 - Aug 2013 Utilize volunteers to assess 50 springs in high-priority areas 

Apr 2012 – Ongoing Work with agency personnel and complementary projects to assess springs/seeps being visited for other projects

Nov 2011 – Aug 2013 Develop a regional spring/seep online database accessible to all jurisdictions

Aug 2012 - Ongoing Direct restoration and protection money and efforts to newly prioritized springs, and incorporate new spring data

 in project planning (e.g. prescribed fire)

Jan 2013 Implement restoration of natural flow and vegetative structure on 12 priority sites

References

Hansen, L.J. and J.R. Hoffman. 2010. Climate savvy: Adapting conservation

and resource management to a changing world. Washington, DC: Island

Press. 256 pp.

Jacobs, K., L. Lebel, [and others]. 2010. Linking knowledge with action in

the pursuit ofsustainable water-resources management. Proceedings of the

National Academy of Sciences. Online: www.pnas.org/cgi/doi/10.1073/

pnas.0813125107.

Karl, T.R., J.M. Melillo, and T.C. Peterson, eds. 2009. Global climate change

impacts in the United States. U.S. Global Change Research Program (USGCRP).Washington,DC:CambridgeUniversityPress.Online:http://www.

globalchange.gov/publications/reports/scientific-assessments/us-impacts/.

Overpeck, J. 2011. Global climate change and the Southwest. Presentation

given at Between a Rock and a Hot Place: Climate Change Adaptation

and Resource Management for the Sky Island Region. April 13, 2011,

Tucson, Arizona. Online: http://www.skyislandalliance.org/adaptationworkshop2011.htm. [Accessed March 15, 2012].
USDA Forest Service Proceedings RMRS-P-67. 2013 67

Responding to Climate Change Impacts in the Sky Island Region: From Planning to Action Misztal, Garfin, and Hansen

organizations and institutionsin the United States. TheAssociation of Fish

and Wildlife Agencies, Defenders of Wildlife; The Nature Conservancy;

and The National Wildlife Federation.

Trenberth, K.E. 2012. Framing the way to relate climate extremes to climate

change. Climatic Change. DOI 10.1007/s10584-012-0441-5.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Overpeck, J., and B. Udall. 2010. Dry times ahead. Science 328:1642-1643.

Robles, M. D., and C. E. F. Enquist. 2010. Managing changing landscapes in

the southwestern United States.Areport of the Southwest Climate Change

Initiative. The Nature Conservancy, Tucson, AZ. 26 pp.

Theoharides,Katie,G.Barnahrt andP.Glick. 2009.Climate change adaptation

across the landscape: A survey of federal and state agencies, conservation
68 USDA Forest Service Proceedings RMRS-P-00. 2012
USDA Forest Service Proceedings RMRS-P-67. 2013 69

Southwestern Cienegas
70 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 71

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Flora and Vegetation of the Saint David and Lewis Springs 

Cienegas, Cochise County, Arizona

Elizabeth Makings

School of Life Sciences, Arizona State University, Tempe, Arizona

Abstract—In the Sky Island region, cienegas are rare marshlands amidst arid surroundings where groundwater perennially intersects the surface. Their unique physical properties give rise to a characteristic plant 

community dominated by wetland graminoids. Evaporation usually causes the water to be alkaline, and 

vegetation around a cienega commonly includes halophytes and other unusual species. Depending on their 

age and size, they may also harbor high levels of endemism. Cienegas are far from pristine, and like many 

wetlands and riparian areas, they have probably been exploited for millennia. In the American Southwest, 

these habitats are hypothesized to have been in severe decline over the past 150 years due to a variety of 

man-made factors. In light of their biological and cultural importance, it is important to learn as much as we 

can about their current condition and conservation potential. 

Introduction

	 Cienegas, described by Hendrickson and Minckley (1985), are

mid-elevation wetlands (3280-6560 ft.) characterized by permanent,

scarcelyfluctuatingwatersources amidaridsurroundings.The aquifer

dynamics,floral, andfaunal charactersof cienegas areunique andvary

according to regional geomorphological and ecological constraints.

Cienega-type marshes and wetlands form where layers of rock or

impervious clay hold water at the surface or through the continuous

upwelling of numeroussmallsprings and seeps(USDI 2012). Cienegas can be very large and harbor high levels of endemism, such as

Cuatro Cienegasin Coahuila, Mexico, with an area of approximately

208,000 acres (325 mi2); or small and cryptic <2 acres (0.003 mi2),

such as the Lewis Springs Cienega in Cochise County, Arizona.

	 The soils of cienegas are typically finely textured, and highly organic. Moisture, salinity, and other abiotic factors are also important

influences on species composition. The result is a characteristic suite

of plantsspecially adapted to these conditions, a rare plant community

dominated by wetland graminoids(monocots)in three main families:

grasses (Poaceae), rushes (Juncaceae), and sedges (Cyperaceae).

Saint David Cienega

	 The San Pedro NationalConservationArea (SPRNCA)is a narrow

stripwith approximately 40miles ofstreamrunning fromtheMexican

border to the town of St. David, Arizona. Among the plant communities of the SPRNCA are two separate cienega habitats: the St. David

Cienega at the northern tip and a little known, much smaller wetland

near the middle associated with the Lewis Springs and Government

Draw drainages (fig. 1).

	 The St. David Cienega is a marshy flat of approximately 350 acres

with a perimeter of 2.3 miles. Seasonality is an important factor, and

cienegas can vary dramatically depending on time of year. The main

portion of the wetland with permanent water is about 75 acres (figs.

2, 3).

	 Surrounding vegetation is a mixture of mesquite grasslands,

bosques, andChihuahuan desertscrub.SaintDavidCienegawould be

completely vegetated with no visible surface water; however, there is

currently active management (vegetation removal and earth moving)

in a northwest portion to provide open water for native fish and frog

habitat (Simms, personal communication). Figure 4 is a landscape

schematic of the vegetation zones of the St. David Cienega. Species

associations are influenced by a moisture gradient from perennially

saturated soils in the middle, where the water intersects the surface,

to a seasonally dry and increasingly saline periphery. Schoenoplectus 

americanus (chairmaker’s bulrush)forms nearly impenetrable stands

throughout the middle portion, associated with Lythrum californicum 

(California loosestrife), and Berula erecta (cutleafwaterparsnip).This

zonehas ankle-deepwaterthroughoutwithorganic,blacksoils(fig.5).

Moving outward, dense clumps ofCarex praegracilis(clustered field

sedge) dominate along withEleocharis rostellata (beaked spike rush)

(fig. 6).Amix of Muhlenbergia asperifolia (alkali muhly), Distichlis 

spicata (salt grass), and Juncus arcticus var. balticus (Baltic rush) can

be found in the “Juncus-Distichlis” zone; transitioning into an area of

tall grasses such as Sporobolus wrightii (sacaton), Panicum obtusum

(vine mesquite), and Bothriochloa laguroides ssp. torreyana (silver

beardgrass). Mesquite (Prosopis velutina) and sacaton outline much

of the perimeter of St. David Cienega.

	 There are naturally various species that can be patchy and locally

abundant, such as Anemopsis californica (yerba mansa), Asclepias 

subverticillata (whorledmilkweed) and Helianthus annuus(common

sunflower).InthenorthwesternsectionofthecienegaTypha domingensis 
72 USDA Forest Service Proceedings RMRS-P-67. 2013

Makings Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona

Figure 1—St. David and Lewis Springs Cienegas 

within the San Pedro National Conservation Area.

Figure 2—St. David Cienega looking northwest; Whetstone Mountains 

in the background (photo taken July 2005). 

Figure 3—Similar perspective of St. David Cienega (fig. 2) taken April 2012.
USDA Forest Service Proceedings RMRS-P-67. 2013 73

Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona Makings

Figure 4—Schematic of St. David Cienega vegetation zones.

Figure 5—Nearly impenetrable stands of chairmaker’s bulrush (Schoenoplectus americanus) in Middle portion of St. David Cienega. Figure 6—Mats of clustered field sedge (Carex praegracilis) in the ‘CarexEleocharis’ zone of the St. David Cienega. 

(southern cat-tail) replaces S. americanus as the dominant wetland

graminoid.

	 Cycles of saturation and evaporation tend to make soils alkaline,

and vegetation in cienegas almost always includes halophytes and

other specialized plants. The St. David Cienega has numerous examples of species with high fidelity to cienegas—their vernacular

often indicating salt tolerance e.g., Almutaster pauciflorus (alkali

marshaster),Distichlis spicata (saltgrass),Muhlenbergia asperifolia,

Pluchea odorata (saltmarsh fleabane), Symphyotrichum subulatum 

var. ligulatum (salt marsh aster), Suaeda moquinii (seablight), etc.

	 A floristic inventory was conducted during the early 2000’s of the

SPRNCAand625taxawerevoucheredfromthatstudy(Makings2006).

Table 1 is a list of 23 of those from the St. David and Lewis Springs

Cienegas that were found nowhere else but in these two habitats.

Some, no doubt, can be dismissed as simply rare and/or overlooked

in the greater SPRNCA, but others might be considered true cienega

indicators. For example, in Cochise County, Almutaster pauciflorus,

Ammannia coccinea, and Muhlenbergia asperifolia have only been

collected from cienegas of the region. Using biodiversity occurrence

data accessed through the Southwest Environmental Information
74 USDA Forest Service Proceedings RMRS-P-67. 2013

Makings Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona

Figure 7—Lewis Springs Cienega looking northeast. Most abundant species are Juncus arcticus var. balticus and Lythrum californicum (California 

loosestrife) (post-monsoon photo taken in October 2011). Grass along 

perimeter is mostly alkali sacaton (Sporobolus airoides).

Figure 8—Eryngium sparganophyllum (Arizona eryngo).

Network (SEINet 2012), this overlapping distribution pattern is

consistent with many other taxa in table 1. In addition to the cienegas

of the San Pedro, they regionally tend to be found in such places as

Empire Cienega in Las Cienegas National Conservation Area (Pima

County), O’Donnell Cienega in the Canelo Hills (Santa Cruz Co.),

San Rafael State NaturalArea (Santa Cruz County), Hooker Cienega

(Cochise County), and Pipe Springs Cienega in the San Bernardino

National Wildlife Refuge (Cochise County).

Lewis Springs Cienega

The Lewis Springs Cienega is approximately 1300 ft. east of the

San Pedro River and just south of the Lewis Springs/Government

Draw drainage.It is an unlikely wetland situated on the side of a small

slope, probably maintained by gravity-driven subterranean flow from

surrounding uplands. The margins of the wetland are abrupt and the

contrast of desert vs. wetland vegetation is especially evident in the

dryer months (fig. 7).

	 The Lewis Springs Cienega is a little less than 2 acres, but has

yielded several species of interest, including ones that appear to be

limited to this tiny wetland. For example, Lobelia cardinalis (cardinalflower), an obligate wetland species, is only known from this

isolated population at Lewis Springs, and nowhere else along the

SPRNCAfloodplain.Also notable, isAndropogon glomeratus(bushy

bluestem), a tall perennial grass that is rare regionally, but abundant

here.‘Cienega indicators’such as Almutaster pauciflorus, Anemopsis 

californica, Lythrum californicum, Mentha spicata (spearmint), and

Muhlenbergia asperifolia are also only present here orin theSt.David

Cienega, and not in the greater SPRNCA.

Table 1—Species found only in cienega habitats within SPRNCA.

Species Cienega location

Almutaster pauciflorusa St. David / Lewis Springs

Ammannia coccineaa St. David

Anemopsis californicaa St. David / Lewis Springs

Berula erectaa St. David

Calystegia sepiuma,b St. David

Carex praegracilisa St. David

Chamaesyce prostrata St. David

Cirsium vulgare St. David

Cyperus spectabilis St. David

Distichlis spicataa St. David

Eleocharis rostellataa St. David

Elymus trachycaulusa St. David

Epilobium ciliatuma St. David

Eryngium sparganophylluma Lewis Springs

Leptochloa fusca ssp. fascicularisa St. David

Lobelia cardinalis Lewis Springs

Lythrum californicuma St. David / Lewis Springs

Mentha spicataa St. David / Lewis Springs

Muhlenbergia asperifoliaa St. David / Lewis Springs

Pluchea odorataa St. David

Pluchea sericea St. David

Samolus valerandi ssp. parviflorus St. David

Sisyrinchium demissum St. David

a

High fidelity to cienegas and similar wetlands.

b

State record.

	 The most noteworthy species in the Lewis Springs Cienega is

Eryngium sparganophyllum (Arizona eryngo), an unusual dicot in

the Apiaceae. Eryngium sparganophyllum is a perennial herb with a

basal rosette of long, parallel-veined leaves. The flowers are cream

and clustered in dense heads at the end of the branching, scapose

inflorescences(fig. 8). Eryngium sparganophyllum is a rare, regional

endemic that occurs in marshes and other riparian habitats. SEINet

data shows only 15 collections of vouchered material from the

region—In Arizona, from Pima and Cochise Counties; In Mexico,

one locality each from Sonora, Coahuila, and south to Durango

(fig. 9). Other biodiversity data such as NatureServe (2012) report

the global range to include Zacatecas and Jalisco, which would be

slightly farther south, but regardless of its distributional limits, the

scant number of collections suggests a rare life history strategy for

this species. Eryngium sparganophyllum can reach a height of >5

ft. and is conspicuous when flowering, and therefore, it is hard to

imagine that this plant is overlooked in the field. In fact, there are
USDA Forest Service Proceedings RMRS-P-67. 2013 75

Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona Makings

anecdotal comments on several labels noting itslocalized abundance

(e.g., Yatskievych in 1982: “dominant in marshy meadow”; Reina in

2003: “very common herbaceous perennial”;Wiensin 2008: “locally

common herbaceous perennial...” The limited distribution, coupled

with such narrow ecological parameters, also make E. sparganophyllum especially susceptible to local extinction, earning its status as

“critically imperiled” in Arizona, and “possibly extirpated” in New

Mexico (NatureServe 2012). Indeed, Wooton and Standley, in their

treatment of the New Mexico flora in 1915, note the only collection

of E. sparganophyllum, a Charles Wright type specimen from “Las

PlayasSprings, neartheSierrade lasAnimas,NewMexico,”probably

the present day Playas Valley east of the Animas Mountains in the

New Mexico “Bootheel” (Wooton and Standley 1915). In addition

to this population, probably many others have been extirpated due

to dewatering of wetlands in the region.

Discussion

	 Cienegas and their environs have been inhabited for millennia and

have great archaeological and biological value. However, the future

of cienegas is uncertain, having been greatly reduced over the last

150 years mainly due to the influences of humans. Cienegas may be

one of the most endangered habitats in the American Southwest and

northwesternMexico,manyonlyremnantsoftheirhistoricalcondition.

Hendrickson and Minckley (1985) estimate habitat loss of cienegas

to be upwards of 95%, making them aquatic islandsin an arid matrix.

FrankCrosswhite, editor of Desert Plants at the time, noted, “Cienega

siteswere thefirstto be usurped by land-hungryHispanics andAnglos

alike who developed large herds of cattle to devourthe vegetation and

drink the water. Overgrazing made the cienega locations among the

mostmistreated sites on earth.Avariety ofmisfortunes, brought about

either knowingly or unconsciously by man, have resulted in drainage, arroyo cutting, and general destruction of these unique habitats”

(Crosswhite1985).Cattlegrazing,development, andassociatedwater

extraction (groundwater pumping, draining, diverting, impounding,

etc.) continue to be problems and have increasingly placed pressures

on the ecological integrity of many cienegas, including those along

the San Pedro River valley.

	 Cienegas are relatively small ecosystems with important roles on

the landscape,yetironically,poorlyunderstood.Inordertoassesstheir

importance and conservation potential, better knowledge regarding

proper functioning of these sitesis critical, including investigation of

their extent, ecology, and abiotic parameters, as well as monitoring

their trends. The survival of the St. David Cienega, Lewis Springs

Cienega, and other rare wetlands of the region will ultimately depend

on (1) a demand for thoughtful choices about where and how much

groundwater is pumped, (2) greater emphasis on research, and (3)

our support of efforts to protect and restore them.

Figure 9—Distribution of Eryngium sparganophyllum with locality (collector and date).
76 USDA Forest Service Proceedings RMRS-P-67. 2013

Makings Flora and Vegetation of the Saint David and Lewis Springs Cienegas, Cochise County, Arizona

References

Crosswhite, F.S. 1985. Editorial. Desert Plants 6(3):130.

Hendrickson, D. A. and W. L. Minckley. 1985. Cienegas—vanishing climax

communities of the American Southwest. Desert Plants 6:130-175.

Makings, E. 2006. Flora of the San Pedro Riparian National Conservation

Area. Desert Plants 22(2): 104 p.

NatureServe. 2010. NatureServe Explorer: An online encyclopedia of life.

Version 7.1. Arlington, Virginia. http://www.natureserve.org/explorer

[April - June 2012].

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

SEINet. 2012. Southwest Environmental Information Network,

http//:swbiodiversity.org/seinet/index.php [April – June 2012].

Simms, J. 2012. Personal communication. Fishery Biologist, United State

Bureau of Land Management, Tucson, Arizona.

Wooton,E.O.andP.C.Standley.1915.FloraofNewMexico.Contributionsfrom

the United States National Herbarium Volume 19. Washington, DC. 794 p.

U.S Department of the Interior. 2012. Land Corral/St. David Cienega Trail.

Washington, DC: U.S. Department of the Interior, Bureau of Land Management. Online: http://www.blm.gov/az/st/en/prog/recreation/hiking/

stdavid.html [June 2012].
USDA Forest Service Proceedings RMRS-P-67. 2013 77

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Paleoenvironmental Framework for Understanding the 

Development, Stability, and State-Changes of Ciénegas in 

the American Deserts 

Thomas A. Minckley

Department of Geography and Program in Ecology, University of Wyoming, Laramie, Wyoming

Andrea Brunelle

Department of Geography, University of Utah, Salt Lake City, Utah

Dale Turner

The Nature Conservancy, Tucson, Arizona

Abstract—As persistent wetlands in arid regions, ciénegas represent important resources for the maintenance 

and preservation of regional biodiversity. The history of ciénegas in the American Southwest over the last 

8,000 years provides information on the dynamics of growth, longevity, and stability of these habitats under 

previous climate conditions. Proxy data such as sedimentology, pollen, charcoal, and isotopes preserved in 

ciénega sediments provide information on the formation, disturbance, resilience and state changes within 

these systems. This long-term perspective is compared to the recent history of degradation observed in the 

region. Once formed, ciénega surfaces alternate between wetland and dryland phases, identified by changes 

in pollen preservation and isotopic signatures. These phase changes are hypothesized to be controlled by 

groundwater-table depths. The degraded state of many extant ciénegas may be similar to the dryland phase, 

but may also require active management to initiate the natural hysteresis of wetland and dryland phases. 

We present a conceptual model on the controls for different ciénega states and how the paleoenvironmental record of change can be used in conservation, restoration, and management of these critical habitats.

Introduction

	 Arid and semi-arid environments occupy 40%ofthe terrestrialland

area and represent regions with the highest human population growth

rates (18.5% between 1990 and 2000; Hassan and others 2005). This

population growth places great pressure on the natural environment

and the ecosystem services it provides, particularly in terms of water

provisioning and water and erosion regulating services. Additional

future pressure will come with warming temperatures projected to

reduce globalstreamflow by 15% with every 1°C of temperature rise

due to the combined effects of increased evaporation and decreased

precipitation (Henderson-Sellers and McGuffie 2012). Beyond fundamental ecosystem services, wetlands (ciénegas) of the American

Deserts have long been identified as regions of high conservation

concern (Abell and others 2000; Hendrickson and Minckley1985;

Leopold 1949; W. Minckley 1969, 1992).

	 High endemism associated with some ciénegas, mostfamously the

wetland complexes of the Cuatrociénegas Valley, Coahuila Mexico

(Abell and others 2000; Badino and others 2004; W. Minckley 1969,

1992), have made these environments a conservation priority. Less

considered are smaller ciénegas across the Chihuahuan, Sonoran,

and a lesser extent Mojave deserts. For example, in Arizona where

wetland environments occupy ~2% of the land area they are critical

habitat for at least 19% of the threatened, candidate, or endangered

species within the state (Baker Jr. and others 2004; T. Minckley and

others in review). However, beyond species of concern, simply by

providing different habitat in otherwise arid regions, desert ciénegas

and riparian corridors may increase regional biodiversity by up to

50% (Sabo and others 2005).

	 At present, there is no systematic understanding of the dynamics

that promote the natural formation and resilience of ciénegas as a

biogeomorphic unit within fluvial systems. However, the literature

on ciénega establishment, development, and resilience has increased

in recent years(Brunelle and others 2010; Heffernan 2008; Heffernan

and others 2008; Heffernan and Fisher 2012; Minckley and Brunelle

2007;T.Minckley and others 2009, 2011).Herewe presentthe current

understanding of ciénega growth and development in the American

Deserts. We focus on the paleoenvironmental record of ciénega

origination and use those data to construct a model of natural ciénega

dynamics compared to their current degraded state, which form the

basis for the management and restoration challenges we presently

face.

	 PriortoEuroAmerican settlement, theAmericanDeserts contained

many ciénegas, near piedmonts, asin Cuatrociénegas, or as emergent

features within regional river systems as is seen in many ciénegas in

Arizona and New Mexico (fig. 1) (Hendrickson and Minckley 1985;
78 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley, Brunelle, and Turner Paleoenvironmental Framework for Understanding the Development, . . . 

T. Minckley and others, in review). The loss and degradation of these

habitats increased rapidly with human settlement, influenced by removal of beaver and their associated dams, draining and conversion

to pasture and agricultural fields, down-cutting along nick points

in wagon and rail roads, and active channelization and draining to

reduce regional malarial threats in the late 19th and early 20th centuries (Bryan 1925, 1928; Fonseca 1998; Hendrickson and Minckley

1985; Leopold 1949). Later impacts on ciénegas include headcuts

resulting from artificial dam installations, which concentrated flood

pulses and increased associated erosive force of those events (fig. 2).

Current threats are associated with both these legacy impacts and

groundwater overdrafts that result in a loss of surface connectivity

between the watertable and root zone.The pattern ofregional ciénega

degradation is shown by comparison of historic vs. extant ciénegas

in southeastern Arizona (fig. 1). Many of these former ciénegas are

located in the major agricultural region ofCochiseCounty,AZ (fig. 1).

Cumulatively the activities associatedwithsettlementhave resultedin

a general degradation of ciénegas by convertingmany into grasslands

andshrub-landsdominatedbymesquite (Prosopis sp.) andotherdesert

trees and shrubs.

Geomorphic Settings Of Ciénegas

	 Ciénega refers to a set of freshwater environments in the North

American deserts and semi-arid grasslands that are typically permanently wetted, either by springs or by water forced to the surface by

channel constrictions or sub-surface features such as bedrock or sills

(Heffernan 2008; Hendrickson and Minckley 1985; T. Minckley and

Figure 1—The distribution of the Apache Highland Grasslands and historic 

and extant ciénegas in southern Arizona, USA, southeastern New Mexico, 

USA, northern Sonora, Mexico, and northeastern Chihuahua, Mexico.

others,inreview).Flow

of subsurface water is

distributed laterally

and longitudinally, allowing ciénegas to

occupy areas 100’s of

meters wide and long.

For example, the San

Bernardino Ciénega,

spanning the border

between Arizona and

Sonora, is estimated

to have been 1.0-3.4

km wide and 6 km

long (Rosen and others

2005), while Canelo

Hills Ciénega is 0.1-

0.14 km wide and ~2

Figure 2—Downstream view from head cut of incised channel near 

Cloverdale Ciénega, NM. Brown, thin, paleo-ciénega soils are evident 

on left bank downstream margin. Poorly sorted and angular rocks and 

cobbles make up the bed material nearest to the head cut.

km long . Because the subsurface controls of ciénega formation are

not well known, the proposed spring classification systemof Springer

and others (2008) presently best describes these environments as

‘low gradient wetlands with indistinct or multiple [water] sources.’

Ciénegas, though aggradational (Minckley and Brunelle 2007; T.

Minckley and others 2011), do not appear to be formed through

anastomosing streams with associated terracing (Leopold and others

1994). Rather, ciénegas appear largely planar, occupying nearly the

entire widths of valley bottoms with little large-scale morphology to

distinguish sub-units within these surfaces. However, preferential

flow channels can exist across the surface between hummocks of

grasses, sedges, and other herbaceous taxa. Aerial photographs of

extant ciénegas show that these features within low-ordered streams

and rivers have distinguishable paleo-stream channels disappearing

above and dendritic channels emerging below permanently wetted

ciénega, creating a more deltaic-like form than a distinct channel

form.

	 Ciénega surfaces appear to have a strong biogeomorphic control

on their formation. Heffernan (2008) proposed that ciénegas represent an alternative stable state to gravel bottom stream channels that
USDA Forest Service Proceedings RMRS-P-67. 2013 79

Paleoenvironmental Framework for Understanding the Development, . . . Minckley, Brunelle, and Turner

characterize many of the river corridors of the American Deserts.

Heffernan(2008)observedthat after4yearswithoutgrazingpressure,

wetlands had developed in over 20% of Sycamore Creek, Tonto NationalForest,Arizona.Largefloodevents(>20m3 s-1)withinSycamore

Creek were able to remove up to 76% of the nascent vegetation cover

in single events, but most lower magnitude flood events (<20m3 s-1)

only affected 20% of the vegetation cover. How longer established

ciénegas respond to floods is not known, but the few analyses in

progress argue for long-term surface stability.

	 Extant ciénegas,such asCanelo HillsCiénega,show post-flooding

evidence of laminar flow avulsing from a “main-channel” that lays

aboveground vegetation down across the entire surface with no evidence of scouring to bedrock (fig. 3). This suggests that a combination of lateral spreading of flood-pulses and aboveground biomass

attenuatesthe erosive strength of floods and protectssoftersediments

of the ciénega surface. Flood events bring in sands,silt and clays, and

may be an important source of fine organic detritus from upstream

sources (Heffernan and others 2008). During non-flood season, the

permanently wetted soils and slow movement of water through the

vegetation matrix of a ciénega provides an environment for capturing

and retaining pollen and other micro and macro botanical remains

that allow for the reconstruction of paleoenvironmental histories

(T. Minckley and others 2011). Based on consistencies in organic

content from a transect ofsediment coresfrom Canelo Hills Ciénega,

it appears that the entire vegetated surfaces of ciénegas may grow

synchronously when climatic and hydrologic conditions are right for

wetland initiation (Berg-Mattson and others, in prep).

Paleoenvironmental Study

	 T. Minckley and others (2011) looked to expand upon Heffernan’s

(2008) multiple stable state hypothesis by applying the interpretation

of multiple stable states to the 8000 year long San Bernardino Ciénega paleoenvironmental record. Instead of a gravel bottom stream

transitioning to a wetland, stabilization of the sand bottom matrix

appears to have occurred ~8.0 ka (ka = calibrated kiloannum years

before1950CE)duringadryphasewherepollenwaspoorlypreserved

and herbaceous taxa and desert trees dominated (fig. 4). Evidence of

perennial water on the surface begins ~7.0 ka and persisted until ~4.0

ka based on pollen preservation and observed aquatic snail shells in

the sediments.After 4.0 ka, pollen preservation decreasessuggesting

aerial exposure of the ciénega until ~2.4-1.6 ka BP. Isotopic analysis

of thissection confirmsthat the surface waslikely occupied by grassland vegetation (fig. 4) (T. Minckley and others 2009). A return to a

wetland state occurs after 2.4-1.6 ka BP based on pollen preservation

and isotopic analyses, which likely persisted until the late 19th to early

20th century (Mearns 1907; T. Minckley and others 2011).

Discussion

	 Extending observations of modern riparian and ciénega dynamics

into the past requires placing interpretations of available proxy (pollen, isotopes and sediment characteristics) into a testable framework

(fig. 5a, b) where there are three possible ciénega states: historically

natural dryland and wetland phases, and a recent degraded phase.

Dryland phases and degraded phases appear similar in the pollen

record having poor pollen preservation and, therefore, only provide

minimal presence-absence vegetation data based on the most hardy

pollen grains (T. Minckley and others 2011). These data are not considered quantifiable by the authors. Pollen preservation is enhanced

during thewetland phase, providingafirst order estimationofwhether

a surface was saturated in the past. Wetland phases, then, are those

that provide the best data for composition of the plant communities

for both ciénegas and surrounding uplands.

	 Isotopic analysis of ciénega sediments provides evidence of state

changes based on identification elemental and isotopic sources of organic material (Meyers 1994; T. Minckley and others 2009). Though

not taxonomically specific, isotopes and specifically the C:N ratio

provide information on past ciénega surface conditions(T. Minckley

and others 2009) with lower C:N valuesindicating wetted conditions

(fig. 4).Isotope data are particularly usefulfor understanding changes

in pollen preservation and were instrumental to the identification of

state changes in the paleoenvironmental record of San Bernardino

Ciénega (T. Minckley and others 2009, 2011).

Reference State Models for Setting 

Restoration Goals in Desert Wetlands

	 Based on our observations of recent and past state changes in

ciénegas, we propose a process-based ecohydrologic model of state

transitions in ciénegas. This model integrates water balance, vegetation structure (above and below ground) and channel form as key

variables that influence the trajectory of biogeomorphic change, and

provides a framework for ciénega restoration and management. This

modelis comparable tostate-and-transitionmodelsusedtounderstand

and manage arid-upland sites that have experienced heavy grazing

pressure resulting in surface erosion, and encroachment of woody

vegetation (Bestelmeyer and others 2003; Laycock 1991).

	 Depending on historic and present water availability, the potential

naturalstates of ciénegasmay be dryland orwetland phases (fig. 5a,b).

Although these phases differ in the relative depth to groundwater and

the persistence of surface waters, both states are characterized by

surfaceswithrelativelyhomogeneoustopography,highorganicmatter

content and high resilience to erosion by floods. The current state of

many ciénegas may be considered in a degraded phase, characterized

by an active geomorphic surface, discrete channels, minimal surface

water, and encroachment of woody vegetation, especially proximal

to incised channel margins.

	 Throughputs for ciénega development are mainly hydrologic:

groundwater (GW) through, surface flow (SF) in and out, and actual

evapotranspiration (AE), which should be a net loss from the system

Figure 3—“Flow-Lines” across the Canelo Hills Ciénega, March 2009. 
80 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley, Brunelle, and Turner Paleoenvironmental Framework for Understanding the Development, . . . 

Figure 4—Scatterplot of C:N and δ13C isotopic values for the past 4000 years next to the pollen data for San Bernardino Ciénega 

(T. Minckley and others 2011) showing the isotopic shift in sediments with poor pollen preservation, indicated by triangles in the first 

three columns, as compared to sediments with good pollen preservation, indicated by shaded pollen abundance curves.

as evaporative potential (PE) is high throughout the year (fig. 5a,b).

However, upstream and upland sediment(SED)sources contribute to

organic detritus, inorganic carbon, and sand, silts and clays (Heffernan

and Fisher 2010; Heffernan and others 2008).

Dryland Phase—The dryland phase is the natural reference state

that occurs when groundwater levels have likely decreased >25 cm

below the surface for obligate wetland species(Stromberg and others

1996), allowing for grass and woody taxa encroachment (fig. 5b) (T.

Minckley and others 2009). Presumably lowered groundwater tables

result in no perennial surface flow, which decreases growth rates

of the ciénega surface (Minckley and Brunelle 2007; T. Minckley

and others 2011). Pollen preservation is reduced during the dryland

phase because of the periodic exposure of deposited pollen grains to

oxidizing conditions. Groundwater flow out of the system is lowered

becauseofplantusage andevapotranspiration.Floodpulse attenuation

occurs in the dryland phase, because surface flow is not channelized.

Rather, broad ciénega surfaces disperse seasonal flood pulses into

sheet floods, preventing channelization. Sedimentary organic content

is high during the dryland phase because of belowground competition

for water resources. This competition further stabilizes the natural

surface from degradation. Surface soil moisture is higher because of

surface shading by woody and grassland taxa. AE is lower than PE

in the dryland phase, but likely greater than in the degraded phase

because of transpiration from vegetated surfaces and hydrologic lift

associated with both the woody and grassland vegetation.

Wetland Phase—The wetland phase is the natural reference state

that occurs when groundwater levels are at or above the vegetated

surface resulting in standing water and saturated soils within and

adjacent to a ciénega (fig. 5a). Most woody taxa are excluded from

the ciénega surface because of the saturated soils. The combination

of surface flow and groundwater flow through the system results in

rapid and consistent aggradation of plant biomass and capture and

burial of fine sedimentsfrom upstream sources(Heffernan and Fisher

2010). Pollen preservation is good during thewetland phase due to the
USDA Forest Service Proceedings RMRS-P-67. 2013 81

Paleoenvironmental Framework for Understanding the Development, . . . Minckley, Brunelle, and Turner

Figure 5—Conceptual models for natural ciénega phases based on hydrologic flux and vegetative responses. Model states form a basis for 

interpreting paleoenvironmental records from ciénega environments as well as provide management and restoration goals for present and 

future stable states. The neutral state forms a null hypothesis for comparing the three observed conditions for ciénegas in the American Desert.

A

B
82 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley, Brunelle, and Turner Paleoenvironmental Framework for Understanding the Development, . . . 

anaerobicdepositional environment.Floodpulse attenuationoccursin

the wetland phase, because of surface characteristics and protection

of soft sediments by the structure of dense above- and below-ground

biomass(Heffernan 2008;Heffernan andothers 2008).Seasonalflood

pulses are slowed and sheet flooding acrossthe entire surfacesresults

in increased inputs of sands, silts and clays (T. Minckley and others

2011). The moisture surplus results in a local balance of AE and PE.

Degraded Phase—Based on this conceptual model, the generic

degraded state of ciénegas can be characterized asincised, having no

surface water, poor groundwater connection for shallow rooted taxa,

and experiencing woody taxa encroachment. Flood pulses are not attenuatedinthedegradedphasebecauseof channelizationconcentrates

flow and reduces sheet-flooding. AE is much lower than PE because

of surface drying and low vegetation densities reduce near surface

soil moistures (fig. 5b). The reconfiguration of the degraded state,

relative to the naturalstates, may be extremely resilient to movement

from this state, since flood power will be increased by concentrated

flows within the channels and the incised channels maintain water

levels well below the rooting zone of the adjacent terraces. Returning to one of the natural states, whether dryland or wetland, may

be difficult based on present water availability, sediment flux, and

vegetated surface-stability (DeBano and Schmidt 1990; Heffernan,

2008; Heffernan and others 2008), but is clearly possible asthe statechange in Sycamore Canyon, Arizona, has demonstrated (Heffernan

2008). Natural re-establishment of ciénegas from the degraded state

may occur primarilywithin recently formed (last 100 years) andmuch

narrower active channels, while re-establishment of ciénegasto their

former extents across entire valley channel widths may require active

interventionthatbothredistributessurfaceflowandraisesgroundwater

levels (i.e., check-dams placed within incised channels) (Heffernan

2008).

Management for the Paleoecological 

Perspective

	 State-changesindesertwetlands occuronmultiple temporalscales;

however, the current conditions of ciénegas in the American deserts

are an artifact of the last 100-125 years of EuroAmerican settlement.

Early research in changes in vegetation associated with lowered

groundwater table (Bryan 1928) was the first indication of change

in the region that favored deep rooted plant species. That changes in

vegetation and hydrology are recent and upon the release of grazing

pressure ciénegas can naturally reform (Heffernan 2008), suggests

the natural resilience of these systemsremains. Stromberg and others

(2008) put forth that the responsiveness of vegetation in riparian corridors to changes in water availability reflected the constant renewal

of the seed bank from both upland and riparian taxa. The constant

double-sourcing of the seed bank would provide a mechanism for

observedrapidvegetationchangesinciénega systemsorstate-changes

over the long-term in both modern and paleoenvironmental studies.

The rapid response may be based simply on changing precipitation

timing and amount, and the consequent groundwater recharge, asthe

depth to water table is likely the ultimate control on natural ciénega

states (Stromberg and others 1996).

	 Setting restoration targetsforthe management of ciénegasrequires

using the best information available. Unfortunately, reference conditions are poorly known asthe degradation of ciénegasin theAmerican

Desertswas a post-hoc observation of a region changed by settlement,

grazing, andland-use (Bryan1925,1928;Leopold1949;Turner2007;

Turner and others 2003). The paleoecological record does provide

insight into how ciénegas behaved under different climate regimes

providing achievable restoration and management targets, based on

current climate conditions andwater availability,bothnatural andlegal

based on water rights. Based on the hydrologic flow and evidence

of two potential natural states, ciénega management and restoration

can target the dryland or wetland state as equally viable and within

the natural range of variability for these environments (Millar and

Woolfenden 1999).

	 Management could be focused on restoration of surface stability

as either a grassland or wetland phase as a first order goal. In either

state, the potential ecosystem services of ciénegas and the potential

for natural state changes are restored. Passive management could

follow allowing the natural seeding of the system to adjust the vegetation composition based on short and long-term climate variability

(Stromberg and others 2008).Ciénegas have the potential to represent

a great success story in conservation given that the degradation of

these systems is relatively recent and we observe a great resilience

in ciénega vegetation released from disturbance pressure (Garnett

and Lindenmayer 2011; Heffernan and Fisher 2012; Heffernan 2008;

Heffernanandothers2008).Optimistically,our expandingunderstanding of the mechanisms controlling ciénega form and function in the

Madrean Archipelago can serve as a model for management of these

resources in arid regions around the world (T. Minckley and others,

in review).

Acknowledgments

	 This manuscript has benefited from many formal and informal

conversations about ciénegas over the past decade. We appreciate

the reviews and work on ciénegas by Jim Heffernan and Bill Radke

whose comments improved the manuscript itself and our thinking

about these systems every time we interact. Thomas Minckley and

Andrea Brunelle thank Cuenca los Ojos, The Nature Conservancy,

Sky Island Alliance, and U.S. Fish and Wildlife Service for permission and facilitation of this research. Funding for these studies has

been provided by the Southwest Center for Environment Research

and Policy (NR-04-02, NR-05-03, NR08-03, NR-09-02).

References

Abell, R., Olson, D.M., Dinerstein, E., [and others]. 2000. Freshwater

Ecoregions of North America: A Conservation Assessment. Island Press,

Washington, DC.

Badino, G., Bernabei, T., De Vivo,A., Giulivo, I., Savino, G. 2004. Under the

desert: The mysterious waters of Cuatro Ciénegas, 1st ed. Associazione

Geografica La Venta, Treviso, Italy.

BakerJr., M.B., Ffolliott, P.F., DeBano,L.F., Neary, D.G. 2004.Riparian areas

of the Southwestern United States: Hydrology, Ecology, and Management.

Lewis Publishers, New York.

Berg-Mattson,N.,Minckley,T.A.,Brewer,S.,in prep.Biogeomorphic controls

on cienega development and growth.

Bestelmeyer, B.T., Brown, J.R., Havstad, K.M., Alexander, R., Chavez, G.,

Herrick, J.E. 2003. Development and use of state-and-transition models

for rangelands. Journal of Range Management 56:114-126.

Brunelle, A., Minckley, T.A., Blissett, S., Cobabe, S.K., Guzman, B.L. 2010.

A ~8000 year fire history from an Arizona/Sonora borderland ciénega.

Journal of Arid Environments 74: 475-481.

Bryan, K. 1925. Date of channel trenching (arroyo cutting) in the arid Southwest. Science 62: 338-344.

Bryan, K. 1928. Change in plant associations by change in water table. Ecology 9: 474-478.

DeBano, L.F., Schmidt, L.J. 1990. Potential for enhancing riparian habitatsin

the southwestern United States with watershed practices. Forest Ecology

and Management 33–34: 385-403.
USDA Forest Service Proceedings RMRS-P-67. 2013 83

Paleoenvironmental Framework for Understanding the Development, . . . Minckley, Brunelle, and Turner

Fonseca,J.1998.Vegetationchanges atBinghamCienega,theSanPedroRiver

valley, Pima County, Arizona, since 1879. Journal of the Arizona-Nevada

Academy of Science 31: 103-116.

Garnett, S.T., Lindenmayer, D.B. 2011. Conservation science must engender

hope to succeed. Trends in Ecology & Evolution 26: 59-60.

Hassan, R., Scholes, R., Ash, N. 2005. Ecosystems and human well-being:

Current state and trends; findings of the Condition and Trends Working

Group, The millennium ecosystem assessment series. Island Press, Washington, DC. p. 917.

Heffernan, J., Fisher, S. 2012. Plant–microbe interactions and nitrogen dynamics during wetland establishment in a desert stream. Biogeochemistry

107: 379-391.

Heffernan,J.B. 2008. Wetlands as an alternative stable state in desertstreams.

Ecology 89: 1261-1271.

Heffernan, J.B., Sponseller, R.A., Fisher, S.G. 2008. Consequences of a

biogeomorphic regime shift for the hyporheic zone of a Sonoran Desert

stream. Freshwater Biology 53: 1954-1968.

Hendrickson,D.,Minckley,W.L.1985.Ciénegas-vanishingclimaxcommunities of the American Southwest. Desert Plants 6: 130-176.

Laycock, W.A. 1991. Stable States and Thresholds of Range Condition on

NorthAmerican Rangelands:AViewpoint.Journal of Range Management

44: 427-433.

Leopold,A. 1949.ASandCounty almanac and sketches here and there.Oxford

University Press, Oxford.

Leopold, L.B., Wolman, M.G., Miller, J.P. 1994. Fluvial processes in geomorphology. Dover Publications, New York.

Mearns, E.A. 1907. Mammals of the Mexican boundary of the United States:

Adescriptive catalogue ofthe species of mammals occurring in thatregion;

with a general summary of the natural history, and a list of trees. Government Printing Office, Washington, DC.

Meyers,P.A.1994.Preservationof elemental andisotopic source identification

of sedimentary organic-matter. Chemical Geology 114: 289-302.

Millar,C.I.,Woolfenden,W.B. 1999.The role of climate change in interpreting

historical variability. Ecological Monographs 9: 1207-1216.

Minckley, T.A., Brunelle, A. 2007. Paleohydrology and growth of a desert

cienega. Journal of Arid Environments 69: 420-431.

Minckley,T.A.,Brunelle,A.,Blissett,S.2011.Holocene sedimentaryandenvironmentalhistoryof anin-channelwetlandalongthe ecotoneoftheSonoran

and Chihuahuan Desert grasslands. Quaternary International 235: 40-47.

Minckley, T.A., Clementz, M., Brunelle, A., Klopfenstein, G.A., 2009. Isotopic analysis of wetland development in the American Southwest. The

Holocene 19, 737-744.

Minckley, T.A., Turner, D., Weinstein, S.R., in review. The relevance of wetland conservation in arid regions: A re-examination of vanishing climax

communities in the American Southwest. Journal of Arid Environments.

Minckley, W.L. 1969. Environments of the Bolsón of Cuatro Ciénegas,

Coahuila, México. Texas Western Press, University of Texas at El Paso.

Minckley, W.L. 1992. Three Decades near Cuatro Ciénegas, México: Photographic Documentation and a Plea for Area Conservation. Journal of the

Arizona-Nevada Academy of Science 26: 89-118.

Rosen, P.C., Radke, W.R., Caldwell, D.J. 2005. Herpetofauna of lowland

bottomlands of southeastern Arizona: a comparison of sites. In: Gottfried,

Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; Edminster, Carleton B.,

comps. Connecting mountain islands and desert seas: biodiversity and

management of the Madrean Archipelago II. Proc. RMRS-P-36. Fort Collins, CO: U.S. Department ofAgriculture, Forest Service, Rocky Mountain

Research Station: pp. 112-117.

Sabo, J.L., Sponseller, R., Dixon, M., [and others]. 2005. Riparian zones

increase regional species richness by harboring different, not more, species. Ecology 86: 56-62.

Springer, A.E., Stevens, L.E., Anderson, D.E., [and others]. 2008. A comprehensive springs classificationsystem:integratinggeomorphic,hydrochemical, and ecological criteria. In: Stevens, L.E., Meretsky, V.J., eds. Aridland

springs in North America: Ecology and conservation. The University of

Arizona Press, Tucson, AZ. pp. 49-75.

Stromberg,J.C., Boudell,J.A., Hazelton,A.F. 2008. Differencesin seed mass

between hydric and xeric plantsinfluence seed bank dynamicsin a dryland

riparian ecosystem. Functional Ecology 22: 205-212.

Stromberg,J.C.,Tiller,R.,Richter,B. 1996.Effects of groundwater decline on

riparian vegetation ofsemiarid regions:TheSanPedro,Arizona.Ecological

Applications 6: 113-131.

Turner, R.M. 2007. Confessions of a repeat photographer. In: Felger, R.S.,

Broyles,B., eds. Dry borders: Great naturalreserves ofthe Sonoran Desert.

University of Utah Press, Salt Lake City. pp. 50-57

Turner, R.M., Webb, R.H., Bowers, J.E., Hastings, J.R. 2003. The changing

mile revisited: An ecological study of vegetation change with time in the

lower mile of an arid and semiarid region. The University of Arizona

Press, Tucson.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
84 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 85

Northern Jaguar Reserve
86 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 87

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G., comps. 2013. Merging science and management in a rapidly changing 

world: Biodiversity and management of the Madrean Archipelago III; 2012 

May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research 

Station. 

When Will Female Jaguars Cross the Border? 

Socio-Demographics of the Northern Jaguar

Peter Warshall

Northern Jaguar Project, Tucson, Arizona

Abstract—Conservation biologists, NGOs, and the USFWS have established a goal of returning a viable 

population of jaguars into the United States. The source population for this recovery will come from Sonora, 

Mexico, the closest sub-population of the species. To maintain a viable population there must be females 

and an active corridor that allows passage of jaguars between Sonora, Arizona, and New Mexico. While 

considerable attention has been paid to the corridor, little attention has been paid to the potential rate of 

expansion of the existing population and the importance of female jaguars to dispersal. This paper’s purpose is 

to highlight what is known of the socio-demographics of northern jaguars, the differing role that females play 

in dispersal, the possible return-times for females to the U.S. Sky Islands, as well as conservation priorities.

Introduction

	 Tothebestofourknowledge,jaguars(Panthera onca),inthe1800s,

lived year-round in some areas of Arizona and New Mexico. About

60 sightings have been more or less confirmed in Arizona and New

Mexico in the twentieth century (Brown and Lopez-Gonzalez 2001).

Most recently (between 1996 and 2012), five male jaguars have been

photographed in theAnimas, Peloncillos,Whetstones,Atascosas and

Baboquivari Sky Islands of the United States. Dispersal male jaguars have been known to range widely; by analogy with other large

cats, hundreds of miles per year (Brown and Lopez-Gonzalez 2001;

Hoogesteijn and Mondolfi 1992).

	 No female jaguars have been recorded in the United States since

1963.The lastfemale with a cub wasshot in 1910 inChevlonCanyon,

Arizona. The nearest female jaguar to the U.S. border was shot south

of Moctezuma, Sonora in 1998.Afemale with cub has been recorded

by camera-traps on the Northern Jaguar Reserve (about 125 miles

from the border) in 2011. As opposed to adult males, adult female

jaguars are not known to range widely (Brown and Lopez-Gonzalez

2001; Hoogesteijn and Momdolfi 1992).

	 Jaguar males have large home ranges that may contain smaller

female home ranges (Hoogesteijn 1992; fig. 1). In addition, there appear to be nomadic malesthat are dispersing and wandering and may

not have any settled home range. The dispersal of female jaguars can

be visualized as the creation of home ranges within the boundaries

of one or two male home ranges (fig. 1).

	 A 1,100 square mile area in Sonora (Lopez-Gonzalez and others

2000, Lopez-Gonzalez and Brown 2002) has been delineated as the

area with the highest jaguar numbers and the most likely source area

for U.S. jaguars. The population of jaguars in this and adjacent areas

has been estimated at 80 to 120. Two general “corridors” that may be

used by Sonoran jaguars to return to the U.S. Sky Island mountains

have been suggested (fig. 2; Valera-Aguilar and others, abstract, this

volume). However, the rates of population expansion and dispersal

through these corridors are unknown. This paper presents the social

and demographic factorsthatmay contribute to population expansion

and dispersal, including very simple math-based scenariosto provide

various timeframes for return of adult female jaguars to the United

States. The results of these scenarios pose two conservation biology

questions:Would it be betterto re-introduce female(s)to the potential

jaguar habitat in the United States (pro-active restoration) or wait for

dispersing females to reach the United States (passive restoration)?

What does knowing probable timeframes contribute to the decisions

necessary for return of a U.S. reproductive population?

Figure 1— Home ranges of jaguars (based on Hoogesteijn and Mondolfi 1992). The yellow circles are male home ranges; the red circles 

are female home ranges; and the dotted blue lines are male wanderers 

without home ranges. 
88 USDA Forest Service Proceedings RMRS-P-67. 2013

Warshall When Will Female Jaguars Cross the Border? Socio-Demographics of the Northern Jaguar 

Figure 2—Locations of Northern Jaguar Reserve, core area of northern jaguar population 

(green polygon) and probable dispersal corridors to the Sky Islands in the United States. 

Blue dots are recent jaguar sightings.

Methods

	 To estimate the probable minimum times for return of female

jaguars, I first assembled what is known and not known of their life

history phenomena in their northern range or, if unavailable, for

southern jaguars in the most open or arid habitats (table 1). Southern

jaguars, especially those living in tropical humid forests (as opposed

to thornscrub, pinyon-juniper woodland, riparian and montane biotic

communities), have differing life-history phenomena (Hoogesteijn

and Mondolfi 1992). Life-history phenomena include (1) sex ratio

at birth, (2) the number of cubs (litter size), (3) age of first reproduction, (4) average lifespan, (5) number of years between litters, and

(6) pre-reproductive mortality of cubs. 

	 In addition, I assembled what is known about home range size and

dispersal(ageatdispersal,overlapwithmother’shomerange,residency

time in home range and direction of dispersal; table 2). For each data

point(tables 1 and 2), the degree ofreliability is noted.These two sets

of admittedly incomplete data were combined into a simple iterative

model to estimate the miles per generation that a female might move

north (table 3). “Optimistic” and “conservative” scenarios were used

to frame the probable number of miles per generation. The numbers

of years for return was calculated from the closest female sightings

to the border to the nearest U.S. Sky Island in which males have been

seen.

	 The scenarios include many assumptions that require further

refinement: (1) more accurate life-history phenomena and dispersal

data; (2) the verification of the postulated corridors and pathways;

(3) further understandings of directional choice and female home

range use; and (4) the use of a circle to define home range shape.
USDA Forest Service Proceedings RMRS-P-67. 2013 89

When Will Female Jaguars Cross the Border? Socio-Demographics of the Northern Jaguar Warshall
90 USDA Forest Service Proceedings RMRS-P-67. 2013

Warshall When Will Female Jaguars Cross the Border? Socio-Demographics of the Northern Jaguar 

Results

	 To the best of my knowledge, table 1 summarizes life-history

knowledge and table 2 presents information on dispersal of female

jaguars.Since nothing is known ofsex ratios at birth, pre-reproductive

mortality, or how females chose which direction to extend their home

ranges, these numbers are subject to significant revision. A young

female could theoretically create her home range in any direction.

Asfew as 25% of all daughters might extend the female population’s

range in a northward direction.

	 Other numbers may also be significantly altered as more is known

of the impact of the northern jaguar’s arid landscape on home range

size and shape and litter size. Jaguar home ranges change widely

with season, the location and number of roads, prey size, and the

first months of raising cubs (Hoogesteijn and Mondolfi 1992). The

quantitative impacts of prey density, human settlements and riparian habitat on northern jaguar home range shape and size have just

begun to be investigated. Two “wildcards” exist: a female could be

discovered much closer to the border or a female might (although

never reported) wander with a male a much greater distance.

Scenarios

	 For the purposes of these scenarios, it is important to emphasize

that young females do not appear to move great distances from their

natal home range. There is some evidence that a female cub may be

given perhaps 17 to 40% oftheirmother’s home range and,should the

mother die, may take over 75-85% of her home range (Hoogesteijn

and Mondolfi 1992). It is not known how mother jaguars divide up

their home ranges when they have two female cubs at the same time

or in sequential litters.

	 A northern jaguar female’s home range size will be considered 25

square miles (a diameter of approximately 5.6 miles) to 37 square

miles (a diameter of approximately 6.8 miles). This is about half the

known home range size of a male on the NJR (Gomez-Gutierrez and

others, abstract, this volume) and about the size reported for several

more open landscapes in South America (table 2). The scenarios assume that each mother has a new daughter every 3 years, which is

the spacing between a mother’s pregnancy and the first litter by the

daughter.

	 Three scenarios (table 3) for the smallest and largest home ranges

were used to frame the number of years for female jaguars to return

to the U.S. Sky Islands. The first assumed that the daughter’s home

range did not overlap with her mother’s but was adjacent to it. The

second assumed a 40% overlap. The third assumed an 85% overlap

(the mother dies). All scenarios make the optimistic assumptions

that home ranges moved north and every litter included at least one

female cub. They all assume the home range’s shape is a circle.

	 When the mother and daughter home ranges are adjacent to each

other (do not overlap), a daughter moves north 5.6 to 6.8 miles every

3 years. In the more conservative scenario a dispersing subadult female shares 40% of her home range with her mother and the daughter

moves north 4.4 to 5.2 miles every 3 years. The most conservative

assumes an 85% overlap with a northward crawl of 1.6 to 2.8 miles

every 3 years.

	 Since 1990, there have been six verified sightings of females

with a cub in northern Sonora (Brown and Lopez-Gonzalez 2001).

The direct, straight-line distances to the U.S. border Sky Islands for

females have ranged from about 100 to about 125 miles and could

easily be twice as large given the rugged terrain. Given all the assumptions, it may be reasonable to speculate that the fastest female

inter-generational lineage might return to the U.S. Sky Islands in 45

to 70 years; conservatively 60 to 85; and if many of our assumptions

are too optimistic, from 100 to 250 years (table 3).

Discussion

	 From these scenarios, it is clear that, if passive restoration is

preferred, jaguar conservation in the U.S. Sky Islands will require a

long-range view, and U.S. lands and prey densities of deer, javelin,

cattle, and smaller prey may need to be protected for a half-century

or more before a female appears. Although not studied in any detail,

the pathways north to the U.S. Sky Islands may become more difficult over the next 50 to 150 years as borderland habitats become

increasingly fragmented, settled by humans, and the border fence

“permanently” blocks some potential routes. In addition, the border

highway in Sonora will be widened within the next few years.

	 A bi-national discussion of how conservation funding should be

spentmost effectively is necessary.Should funding be spentinSonora

to improve dispersal from the core zone in all directions or spent

primarily on helping northern jaguar females move north? Which is

the more effective way to raise the population from the present 40 or

50 female jaguars to a more viable population size? Should funding

preferentially address land protection within the “corridor” to the

south from the Northern Jaguar Reserve to Sinaloa, which restocks

the small northern population and may provide genetic diversity?

Should active restoration by transferring females north into the U.S.

Sky Islands be attempted?

	 The scenarioswillhopefullyimprove inaccuracyinthenextdecade

or two. Perhaps a female will be photographed much closer to the

border. The scenarios provide a framework for decision-making that

elevatesthe importance offemales and rates of dispersalin addressing

conservation issues.

Acknowledgments

Thanks to Carlos Lopez Gonzalez, Carmina Gutierrez-Gonzalez,

Miguel Gómez-Ramírez, Megan Southern, Northern Jaguar Project

and Naturalia. All ideas and concepts are, of course, my own.

References

Brown, David E.; López González, Carlos A. 2001. Borderland jaguars. Salt

Lake City: University of Utah Press. 170 p.

Hoogesteijn,Rafael;Mondolfi,E.1992.TheJaguar.ArmitanoPublishers.183p.

Lopez-Gonzalez, C. A.; Brown, D.E.; Lorenzana, G. 2000. El jaguar en Sonora, ¿desapareciendo o solamente desconocido? Especies. Revista sobre

Conservación y Biodiversidad. 9(3): 19–23.

Lopez-Gonzalez, C. A.; Brown, D.E. 2002. Distribución y estado de conservacio ́n actuales del jaguar en el noroeste de Mexico. In: R. A. Medellın,

C. Equihua, C. L. B. Chetkiewicz, [and others], eds. El jaguar en el nuevo

milenio. Fondo de Cultura Economica Universidad NacionalAutónoma de

Mexico. Wildlife Conservation Society. Mexico, D. F. 379–391.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 91

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Bird Ecology and Conservation on the Northern Jaguar 

Reserve: Recent Lessons 

Peter Warshall

Northern Jaguar Project, Tucson, Arizona

Aaron D. Flesch

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—The Northern Jaguar Reserve is in the western foothills of the Sierra Madre Occidental 

and in a broad transition zone between Nearctic and Neotropical faunal realms. We have assessed the 

distribution and abundance of birds across all four seasons in foothills thornscrub, oak woodland, and 

adjacent riparian areas, and discuss issues relevant to conservation and management. Since 2007, we 

have observed 205 species of birds including 131 that are Neotropical migrants, 104 that likely breed 

on the reserve, and 35 species of binational conservation concern. Observations suggest this 20,140 -ha reserve 

has significant conservation value to migratory birds that traverse an inland migratory flyway that 

includes the Sky Islands; breeding species that use foothills thornscrub, oak woodland, and riparian 

woodlands along the Rio Aros and its tributaries; and numerous species and populations of significant 

conservation value. Existing conservation efforts include land purchase, cattle exclusion, invasive plant 

removal, and habitat restoration on both neighboring private ranches and the reserve.

Background

	 The20,140hectareNorthernJaguarReserve (NJR)sits atthenorthwestern foothills of the Sierra Madre Occidental, the “mainland” of

mountains south of the Madrean Archipelago. It lies in the transition

zone between the Nearctic/Neotropical faunal areas of the planet and

between the coastal and high sierra montane regions of Sonora. The

nearest Sky Islands that support pine-oak woodland are the Sierra la

Madera and the Sierra Bacadehuachi.

	 The NJR is surrounded on three sides by the deep canyons of the

RíosAros andYaqui.This difficult-to-accessregion is among the largest and least fragmented regions in northwest Mexico and supports a

population of jaguars (Panthera onca) (Brown and Lopez-Gonzalez

2001),whichbecame the centralfocusfor creatingthisprivate reserve.

Naturalia (a Mexican conservation organization) owns the reserve

and the Northern Jaguar Project and Naturalia manage it.

	 Before 2007, few ornithologists had visited this region of foothills

thornscrub, oak woodland and shrubland, and riparian woodlands in

east-central Sonora. Thus, there were few data on the occurrence,

abundance, breeding and residency status of birds in the interior

foothills of Sonora on the western edge of the Sierra Madre and little

information to guide conservation and habitat management efforts.

Between 2007 and 2011, we surveyed birds on and around the NJR

and estimated the status and seasonal abundance of all bird species.

This article discusses lessons we have learned about birds and their

conservationimplications.Estimatesofseasonal abundance andstatus

of birds will be presented in a companion paper(Flesch andWarshall,

in preparation). 

Methods

	 Detailed information on sampling methods and locations are described in Flesch (2007) and are not repeated here. To estimate bird

abundance, we used distance sampling along eight line transects

surveyed during each of four seasons over 1-2 years; seasons were

defined by the breeding and migratory phenology of local species.

To assess breeding, migratory and residency status, we observed bird

behavior, conductednestsearches andusedmist-nettingandincidental

observations from this and other efforts across the region. 

Lesson 1: An Inland Migratory Corridor for 

Birds 

	 Data gathered during our efforts on theNJRsuggest the importance

of an inland (vs. coastal) flyway formany types of Neotropicalmigratory birds in Sonora. We observed 78 species of passage migrants,

which represented 38% of all 205 species observed in the region. In

addition to passage migrants that often move long distances between

their winter and summer ranges, the reserve supports some species

of short-range migrants including many that likely originate in the

southwestern United States.Although determining the origin ofthese

short-range migrants is difficult without extensive banding efforts or

moleculartechniques, probable short-rangemigrantsthat occurin the

NJR include Turkey Vulture, Black Vulture, Common Blackhawk,

Bald Eagle, Elegant Trogon, Costa’s Hummingbird, Lucy’s Warbler,

Buff-collared Nightjar, Inca Dove, and Common Ground-Dove.

	 During winter, some short-distance migrants that occur on the

NJR are elevational migrants that likely breed to the north and east

in montane habitats of the Sky Islands and adjacent Sierra Madre.
92 USDA Forest Service Proceedings RMRS-P-67. 2013

Warshall and Flesch Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessons

These species include: White-eared Hummingbird, Buff-breasted

Flycatcher,TuftedFlycatcher,GreaterPewee,Slate-throatedRedstart,

and Brown-backed Solitaire. At this relatively northerly latitude,

Tufted Flycatcher and Slate-throated Redstart that breed locally to

the east and southeast may migrate downslope, north and west into

the NJR to winter.

	 Finally, there are migratory species of more tropical affinity many

of which do not occur as far north as the Sky Islands of the United

States, but may breed in the Mexican Sky Islands. They breed or are

suspected to breed on the NJR reserve at or near the northern margins

of their range and most migrate south in winter. These species may be

short-distancemigrants(MilitaryMacaw,White-tippedDove,Whitestriped Woodcreeper, Rose-throated Becard, Thick-billed Kingbird,

SinaloaWren,BlueMockingbird,Rufous-cappedWarbler,Fan-tailed

Warbler, and Five-striped Sparrow) orlong-distance (Sulphur-bellied

Flycatcher, Yellow-green Vireo).

	 The seasonal bird assemblages are additionally complex because

differentsub-populations of the same species may be permanent residents or dwell on reserve only as breedersin summer, over-winterers,

or passage migrants. At times, it is not certain which sub-population

(the migratory, resident) is present, although for some species such

as Lucy’sWarbler mid-summer passage migrants often arrive several

weeks after most spring breeders have departed. Without additional

study andwiderspatial coverage, the exact origin ofmigrantsremains

unknown.

	 In summary, the reserve isthe first area in the foothills of the Sierra

Madre Occidental in Sonora to be surveyed intensively for birds.

Among seasons, bird populationsinclude speciesthat areNeotropical

passagemigrants,Neotropicalsummerbreeders, elevationalmigrants

that breed in Madrean environmentsin nearby mountains, permanent

residents, and short-range over wintering migrants (table 1). Differences in density among seasons indicate large influxes of migratory

species dominated by flycatchers, vireos, and warblers during spring

and again during late summer and fall that together represented large

proportions of total bird abundance compared to winter and summer

residents (Flesch and Warshall, in preparation). About 60% (n =

131) of bird species have been categorized as Neotropical migrants

by the U.S. Fish and Wildlife Service. When elevational migrants

and subtropical summer breeders are considered, about 75% of bird

species observed had migratory habits.

	 These fall and spring bird assemblagesreveal the great complexity

(table 1)that occursin the Neotropical/Nearctic transition zone.They

pose asyet-to-be-researchedconservationchallengestounderstanding

Sky Island bird assemblages. For instance, does the reserve region

represent a weak or strong “link” in the annual cycle of one or more

populations of birds? Which habitat features and resources promote

high rates of winter and summer resident survival and/or population

growth? What are the high-quality foraging plant associations for

passage migrants? Do the same species use the flyway in both spring

and fall? Do migratory flyways shift away from the coastal plain to

the foothills and mountains in fall? What species and habitats are

most critical for conservation? To address some of these questions,

we are now studying over-winter survival of migratory populations

and continuing to monitoring abundance across space and time.

	 The potential importance of an inland corridor for migrants has

changed conservation thinking about land purchase and habitat protection in inland Sonora. How many “stepping stones” are protected?

How wide is the flyway? The closest reserves in the inland corridor

to the NJR are the state reserve (Cerro Las Conchas/Arivechi) and

the Ajos-Bavispe federal reserve, which includes the nearby Sierra

la Madera and upper Rio Bavispe, and the more distance Sierra los

Ajos, Buenas Aires and Purica. The NJR is the only protected area

where logging, land clearing, and cattle grazing cannot occur. A

proposed Presidential decree would also protect it from mining. In

addition to the reserve, the Sanctuario Aquila Calva (that covers part

of the NJR) and Sierra Huerfana have been proposed (Burquez and

Martínez-Yrízar 2006). Elsewhere, habitat protection in east-central

Sonora is de facto in origin and driven largely by the inaccessibility

of these lands to people and livestock.

Lesson 2: The Importance of Foothills 

Thornscrub and other Plant Associations

	 Vegetation in the uplands oftheNJRis composed predominantly of

foothills thornscrub (FT), which is drought deciduous, in leaf during

and after the summer monsoon season from July through September,

and defoliated during much of the winter and spring (Brown and

Lowe 1980). The northern edge of foothills thornscrub occurs less

than 50 km to the north and 100 km to the northwest, yet important

dominants such as Lysiloma divaricatum and L. watsonii, Bursera 

fagaroides, and Ceiba acuminata occur at scattered localities as far

north as the Baboquiviri and Rincon Mountains (Brown and Lowe

1980; Turner and others 1995). Our observations suggest that FT is

an important and undervalued biotic community for migratory and

breeding birds. The NJR is the only reserve in Mexico that protects

extensive areas of FT.
USDA Forest Service Proceedings RMRS-P-67. 2013 93

Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessonsr Warshall and Flesch

	 Foothills thornscrub is a diverse vegetation community with various discrete plant associations that occur at varying elevations and

soils (Burgess 2009). Given this diversity, some plant associations

and plant species within FT and their value to specific species and

foraging guilds need to be defined so that this information can be

applied to developing more efficient conservation plans and priorities. Based on our knowledge of the natural history and ecology of

birds and their habitats in this region, we began, for the first time,

to define these various plant associations and assess their potential

value to birds (table 2). For example, the tree ocotillo/tree morning
94 USDA Forest Service Proceedings RMRS-P-67. 2013

Warshall and Flesch Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessons

glory/Justicia association (Fouquieria macdougalii /Ipomoea 

arborescens//Justicia sp.) flowers in late winter and provides 

valuable nectar resources to migratory hummingbirds and 

orioles. Monsoon rains provide hillslope FT, open woodland 

FT, xeroriparian FT, mesic phase and rocky-bottom FT with 

abundant foliage and insects for foliage gleaners and aerial 

foragers. Guacima, Lysiloma, Bursera, Cebia and the fig species

Ficus pertusa and F. insipida provide fruit for Military Macaws 

and other frugivores.

	 This new attention to FT has brought up new conservation and

management questions. They include how fire (both natural and

human-caused), freezing temperatures, and drought affect needs of

various species of birds; how over-grazing and over-abundant native

and non-native invasive plants affect bird species composition and

density, and how harvesting of wild agaves affect populations of

nectivores. As seasonal utilization of the plant associations and plant

species that occur within FT becomes better known, restoration of

previously grazed or cleared lands can bemore intelligently designed.

Thusfar NJPand Naturalia have focused on more mesic riparian sites

that support woodlands.

Lesson 3: Importance of the Rio Aros and 

Riparian Vegetation

	 The reserve includes 37 km of waterfront along the Aros River,

one of the few remaining large free-flowing rivers in Mexico. NJR

and a federal reserve in its headwaters (Tutuaca) are the only areas

with protected status along the river (Burguez and Martínez-Yrízar

2006). The Rio Aros on and around the NJR supports among the

highest densities of breeding Common Black-Hawks in northern

Mexico (Rodríguez-Estrella 1990; O’Brien and others 2006), the

southernmost breeding population of Bald Eagle in the interior of

Mexico, the northernmost breeding population of Military Macaw,

and populations of many other species at the northern or southern

end of their breeding and wintering ranges (Flesch and Warshall in

preparation). One obvious lesson is that the vast lowland corridor

formed by theRioAros,Yaqui, andBavispe provides valuable habitat

for birds that require riparian woodlands, tall trees, beachfront and

riparian shrub communities, and that consume aquatic prey (table 2).

	 Effortsto protect habitat on and around the NJRhave included land

purchase, especially valuable riparian habitats; fencing of alluvial

terraces and shorelines from cattle in partnership with neighboring

ranches; establishing a plant nursery to support restoration work;

partnershipswithMexican naturalresource agencies and neighboring

landowners to restore riparian vegetation including sycamore woodlands, andconversionof cattle tanksintomorewildlife-friendlyponds.

The goal of these efforts is in part to augment habitat for birds such

as Bell's Vireo, Yellow and Wilson's Warblers, and Yellow-breasted

Chat that are associated with or obligated to riparian habitats(Skagen

2005). On a river basin scale, conservation objectivesinclude educating the public, collecting data to help prevent the construction of a

dam on the Rio Aros, and limiting potential toxic chemical pollution

into the river from mining.

Lesson 4: Birds of Conservation Concern

	 Birdsofconcernareoftenclassifiedatlocal,regionalandcontinental

scalestoaddressvariablethreatstodifferentpopulationsamongspecies

and subspecies (Shuford and Gardali 2008). We identified 34 birds

on the NJR (table 3) that are listed as birds of concern by USFWS,

NationalAudubonSociety,PartnersinFlight andtheMexicanFederal

Government (Berlanga and others 2010; Partners in Flight 2006).

These species include 28 migrants as well as subtropical breeders.

The NJR also supports molt migrants. Habitat protection for moltmigrants is a new and potentially important conservation objective.

Many birdsthat breed in western NorthAmerica leave their breeding

grounds in July and August and migrate to the “Mexican monsoon

region” where they use seasonally abundant resources to molt and

refuel before continuing to their wintering grounds. Because molting

is among the most energy-demanding events in the annual cycle of a

bird, maintenance of high quality stopover habitat for these species

is crucial to their persistence (Pyle 2009). Little was known of this

phenomenonintheinlandfoothillsofSonorauntilNJP’srecentstudies.

	 Of 34 species of likely molt-migrants in the broader region (Pyle

2009), NJP has caught, measured, and banded 14 (40%) in 2011

(Hanunselka, personal communication;table 3).Nine ofthese species

(table 3) are classified as Birds of Conservation Concern or Priority I

species by Partners in Flight (2006). In addition to confirmed moltmigrants (Hooded Orioles, Black-headed Grosbeaks, and Summer

Tanagers which all showed clear molting), we have also recorded

eight ”candidate” species of molt migrants (including two common

passage migrants, Warbling Vireo and Wilson’s Warbler).

	 Thesebirds-of-concernlists,StateofSonora,Mexicanandbinational

conservation initiatives aswell asstrategic surveys andmonitoring on

the NJR are crucial conservation toolsto aid in recovery and forestall

further listings.
USDA Forest Service Proceedings RMRS-P-67. 2013 95

Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessonsr Warshall and Flesch

Conclusions 

	 The fully protected NJR provides essential wintering and stopover 

habitat for migratory and important breeding habitat for over 200 species of birds. Foothills thornscrub, the Rio Aros, arid oak woodlands

and an inland migration flyway are very likely essential components

in the annual cycle of many Neotropical migratory species. Foothills thornscrub is apparently an undervalued biotic community for

migratory birds. Data collected by NJP and their cooperators have

established abaseline formonitoring long-termtrendsin the presence,

abundance, and dispersal of birdsin thisregion that may be due to the

impacts of climate change or, we hope, to habitat protection efforts.

Acknowledgments

	 We would like to thankAlberto Burguez, Diego Ezrre,Alan Craig,

Megan Southern, Richard Hutto, and Noel Snyder for help on this

manuscript, and many birders (Flesch and Warshall, in preparation),

especially Sky Jacobs. Support for this project was provided, in large

part, by grants by U.S. Fish and Wildlife Service, Aros/Yaqui Rivers

HabitatConservationNMBCAGrantAwards3543,4072,4520,4770,

5054 and the University ofArizona, School of Natural Resources and

the Environment. 

References

Brown, D.E. and C.H. Lowe, Jr. 1980. Biotic communities of the Southwest.

USDA Forest Service and University of Utah Press, Salt Lake City.

Brown, David E., López González, Carlos A. 2001. Borderland Jaguars.

University of Utah Press. 170 p.

Burgess, Tony. 2009. Framework to Interpret Vegetation of the Los Pavos,

Northern Jaguar Reserve. Unpublished paper on file at: Northern Jaguar

Project, Tucson, AZ. 12 p.

Búrquez,Alberto andA. Martínez-Yrízar. 2006. Conservation and Landscape

Transformation in Northwestern Mexico. In: Felger, R.S. and Broyles, B.

Table 4—Cont.
96 USDA Forest Service Proceedings RMRS-P-67. 2013

Warshall and Flesch Bird Ecology and Conservation on the Northern Jaguar Reserve: Recent Lessons

(Eds Dry Borders: Great Natural Reserves of the Sonoran Desert. The

University of Utah Press. Salt Lake City. 750 p.

DeSante, D., Romo de VivarAlvarez, C., Morales, S. 2009. MoSI (Monitoreo

de Sobrevivencia Invernal = Monitoring Overwintering Survival) Manual.

Contribution 214, The Institute for Bird Populations. Pt. Reyes, CA.

Flesch, Aaron D. and P. Warshall (in preparation). Status and seasonal variation in bird abundance in thornscrub and riparian vegetation in east-central

Sonora, Mexico.

Flesch, A. D. 2008. Migratory and resident birds of the Northern Jaguar Reserve of east-central Sonora, Mexico. Report to Northern Jaguar Project

and Naturalia A.C. On file at: Northern Jaguar Project, Tucson, AZ.

Hanunskela, Adam. 2011. [Letters to Northern Jaguar Project]. Five data

sheets, September and November 2011. On file at: Northern Jaguar Project,

Tucson, AZ.

Hutto, R. L. 2000. On the importance of en-route periods to the conservation

of migratory landbirds. Studies in Avian Biology 20: 109-114.

O’Brien, C,A. D. Flesch, E. Wallace, M. Bogan, [and others]. 2006.Biological Inventory of the Rio Aros, Sonora, Mexico: A river unknown.

Final

Report to T&E, Inc by School of Natural Resources and U.S. Geological

Survey, Sonoran Desert Research Station.

Partners in Flight. 2006. Species Assessment Data Base. Global Scores.

Online: www.rmbo.org.

Pyle, Peter, W.A. Leitner, L. Lozano-Angulo, [and others].. 2009. Temporal,

spatial, and annual variation in the occurrence of molt-migrant passerines

in the Mexican monsoon region. Condor 111: 583-590.

Rodríguez-Estrella, R. and B. T. Brown.1990. Density and habitat use of

raptors along the Rio Bavispe and Rio Yaqui Sonora Mexico. Journal of

Raptor Research 24: 47-51.

Shuford, W. David, and Gardali, T., editors. 2008. California bird species of

special concern: A ranked assessment of species, subspecies, and distinct

populations of birds of immediate conservation concern in California.

Studies of Western Birds 1. Western Field Ornithologists and California

Department of Fish and Game.

Skagen, Susan K., J.F. Kelly, C. Van Riper, [and others]. 2005. Geography of

spring landbird migration through riparian habitats in southwestern North

America. Condor 107: 212-227.

Turner, Ray M.,J. E. Bowers, and T. L. Burgess. 1995. Sonoran Desert plants:

an ecological atlas. University of Arizona Press, Tucson, AZ.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 97

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Buying Land for Conservation Purposes in Sonora, Mexico

S. Lucía Perez-Weil, Juan Carlos G. Bravo

Naturalia, Comité para la Conservación de Especies Silvestres, A.C., Northern Jaguar Project, Hermosillo, 

Sonora, México

Abstract— The Northern Jaguar Reserve is 50,000 acres and one of the largest privately owned wildlife 

preserves in Sonora. Buying land in remote parts of Sonora takes special knowledge as ownership rules 

may not be clear and boundaries may not be defined in the records. There are complex legal procedures 

to guarantee ownership in which letters of intent play a crucial role, and there are special complexities for 

ejido lands. In addition, only a presidential decree as a natural protected area can prevent mining in the long 

term. The lessons learned at the Northern Jaguar Reserve for conservation land purchases will be described.

Introduction

Land purchase by organizations or individuals with the purpose of 

preserving its natural resources is only one of many other mechanisms 

for conservation with its advantages and disadvantages like any other. 

In the case presented here, it has been the most suitable mechanism 

because of the land tenure conditions in the Sierra of Sonora and 

because of the conflict between the main economic activity in the 

region, cattle-raising, and the main objective of the project developed 

by Naturalia, A.C. and the Northern Jaguar Project, protection of 

carnivores and their habitat. The expansion acquired by Naturalia 

and Northern Jaguar Project in the Sahuaripa municipality increases 

the number of acres to 51,346.10, becoming the largest private reserve 

in the State.

This paper presents a glance of the road followed to acquire that 

property, the obstacles found, and the remaining work to strengthen 

this conservation strategy, through appropriate legal means, which can 

guarantee our conservation objective in perpetuity. Land purchase is 

shown as a means to a goal—conserve jaguar habitat— not as a goal 

by itself. Learning from its limitations we must find alternative ways 

to preserve jaguar habitat, through community work or implementing 

public and governmental mechanisms that can achieve this goal.

Biodiversity Conservation In Mexico

Nature Preserves known as Natural Protected Areas (NPA) in 

Mexico have been the most common mechanism to protect representative portions of natural environments in the different biogeographic 

and ecological regions of the country’s territory when these portions 

deserve to be preserved or restored because of their good condition 

and their fragility or relevance on the maintenance of the regional 

ecological balance. For that purpose, the Mexican Legal System assigns 

to them a special protecting regime by the issuing of a presidential 

decree that limits land and natural resources use. Most of the NPAs 

are being established on private or social property, such as marine 

areas and island territories, with only a few terrestrial areas (e.g., 

the San Pedro’s Sierra National Park in Baja California) demarcated 

in national property. More than 70% of the territory within NPAs is 

owned by private parties (Gutiérrez and others 2002). States and 

municipalities also have faculties to crate NPAs in their own territories 

and by their own competence. Those areas, like the federal ones, can 

be established on private and social property.

The government has tacitly recognized that it is incapable of being in charge of protecting or restoring all the areas of biological 

importance that require it. This is evident in the increasing new rules 

directed at recognizing and encouraging civil society participation 

on conservation matters. Legal reforms to environmental Mexican 

Law (LGEEPA, its acronym in Spanish) in 1996 opened the door for 

registering units for wildlife management and conservation (UMA), 

which are private properties where the environmental authority allows 

wildlife management for restoration, protection, research, exhibition, 

recreation, and environmental education or sustainable development 

activities. With this mechanism, habitat preservation is also promoted, 

not just wildlife populations.

Since 1996, the 59th Article of LGEEPA provided for the possibility for owners and possessors who have biologically relevant lands 

to promote, by themselves, the establishment of a federal NPA for 

them. However, a decree process can take a long time and requires 

certain qualitative and quantitative land specifications to be able to 

be considered for some of the existing NPA categories. To improve 

and make this mechanism more inclusive, LGEEPA’s regulations on 

NPA matters were issued in 2001. The Seventh Title of this regulation 

provided specific regulation for the referred article so that interested 

private owners could voluntarily request a certification of their properties as designated for protection, preservation, or restoration purposes 

of their land’s natural resources. In contrast with federal, state, or 

municipal NPA, while those areas do not imply a decree process, they 

can be established temporarily for no less than10 years. They do not 

require a minimum land extension but only well preserved conditions 
98 USDA Forest Service Proceedings RMRS-P-67. 2013

Perez-Weil and Bravo Buying Land for Conservation Purposes in Sonora, Mexico

and it is the owner who defines the measure of the restriction for 

the use of his land and its resources. The Environmental Ministry 

(SEMRNAT) issues a certificate to the owner, which can be revoked 

if the conservation objectives are not met.

In some states, including Sonora, legislation has replicated these 

certification schemes with a few differences. Moreover, some states 

have recognized through their environmental laws some other mechanisms such as conservation easements—contractual commitments 

between private owners or between them and a government entity 

for natural resources protection in which the owners impose limits to 

their properties’ use as recognition of the environmental services they 

provide; rural reserves— social properties destined for conservation 

purposes by the agrarian groups (from ejidos and communities) who 

are their owners; and private gardens with the objective of advancing 

native species conservation and regeneration.

Although governments have shown interest in integrating society 

to national conservation efforts, regulation is almost non-existent. 

Furthermore, paperwork requirements for getting a certification or 

a NPA decree are linked to the time and budget of the Public Administration. Another concern is that there is not any guarantee that 

a decreed NPA is effectively protecting the land and its resources 

because of many reasons including the lack of a designated budget, 

territory management issues, and diverse disagreements on public 

policies. Because of this, organized civil society has tried to apply 

or develop alternative mechanisms using existing civil regulations 

such as voluntary easements.

Any of those mechanisms require the owner’s willing participation 

because the owners are the ones who will ultimately commit to manage 

land in an environmentally responsible way. However, it is common 

for owners’ interests to change or to be incompatible with natural 

resources conservation. It is in these cases when non-governmental 

organizations with conservation purposes seek ownership of strategic 

properties to guarantee their long-term protection. In this manner, 

land purchasing constitutes another alternative mechanism to obtain 

the protection of ecosystems.

Northern Jaguar Reserve’s Case

As part of the “borderlands jaguars” research efforts between 

Mexico and the United States, David E. Brown and Carlos A. López 

G. found that most of the available jaguar records of dead, captured, 

or photographed animals in Sonora between 1996 and 2000 were 

within a 50-mi radius where the Aros and Yaqui Rivers converge—

they called this area Huasabas-Sahuaripa. The jaguar population 

in the area was still self-sustainable even though it had been over 

exploited (Brown and López 2001). Since then, the area has attracted 

the interest of researchers and has been visited or explored regularly. 

The area was also defined as the most northern region of jaguars in 

Mexico. Moreover, it has been cataloged as a Priority Terrestrial Region for Conservation by the National Commission for Biodivesity’s 

Knowledge and Use (CONABIO) (Arriaga and others 2000). One 

of the land owners near the Aros River, close to where the Bavispe 

joins it to form the Yaqui River, offered one of his properties for sale 

in 2003, opening a unique land-purchase opportunity.

To obtain the first property for creating a “private reserve” where 

jaguars along with other associated species could be monitored and 

preserved, Naturalia, A.C. bought the property known as Los Pavos de 

Arriba and Los Pavos de Abajo, an area of 10,001.48 ac, on July 15, 

2003. This is the how “Northern Jaguar Reserve” (RJN) was created. 

In the closing process, Naturalia was supported by the newly-formed 

Northern Jaguar Project. This purchase involved relatively little 

paperwork because the owner initiated the negotiation and there was 

a proper title issued by Agrarian Ministry (SRA), which is not common in the region. Most of the lands are only “possessions,” that is, 

irregular settlements where ranchers have lived for several generations. 

The Mexican Government recognizes these ranchers as “possessors” 

but not as owners because the properties that have not been titled in 

someone’s name but rather are called original properties belong to 

the Nation. The process for obtaining a property title issued by the 

SRA begins through the possessor’s request for buying the possession. The rancher must then demonstrate that he has been living on 

the land or that he has used that land for at least 3 consecutive years; 

that nobody argues against the boundaries of the requested land; and 

that the possession has to be paid to the Nation at a price depending 

on the land’s quality or aptness for agricultural or forestry activities.

The Nation usually does not take actions to claim its property rights 

and, therefore, the possessors seldom need more legitimacy than that 

which already results from the acknowledgement of neighbors and 

the local authorities, the Ayuntamientos, which in fact recognize possession rights by registering them at the local cadastral office and by 

giving them access to productive subsidies. This way possessors do 

not feel compelled to regularize their possessed land while in practice 

they act as owners without needing to spend money they do not have 

in buying what they consider to be their own land.

Once Naturalia acquired the referred property, the livestock were 

removed and wildlife monitoring began though camera-traps placed 

on it and the neighboring properties once a signed collaboration agreement was in place. The results from the monitoring effort confirmed 

the presence of jaguar and other carnivores such as puma, ocelot and 

bobcat. Moreover, some other endemic species like eared trogon or 

endangered animals like military macaw occurred on the property. All 

of this increased the interest in this area and motivated the expansion 

project.

Expansion

After buying Los Pavos, Naturalia with its partner, NJP began 

negotiation with the owner of Zetazora, a group of four ranches 

adjacent to the south-side of Los Pavos. This negotiation was complicated, however, since the owner’s expectations after the Los Pavos 

purchase had increased considerably. On the other hand, although 

all the properties had a proper title, not all of them came from SRA. 

They had diverse registration records and had belonged to different 

owners with some of them “regularized” by a declaratory judgment 

of ownership though a civil law procedure.

It is common among the possessors that have not obtained title 

by an agrarian procedure to go before a civil judge to ask for the 

“prescription” of their possessing rights through a voluntary jurisdiction— a non-contentious trial. Sonora ́s Civil Code and most of the 

civil law from other states allow a possessor to ask for a prescription 

that recognizes the possessor as the owner when they possess the 

land as if they were owners. That possession is recognized by society 

and, importantly, the owners of adjacent properties. The possession 

does not have to be registered under a person’s name at Property and 

Commerce Public Register (RPP). In this case, the possession had 

been exerted for at least 3 or 10 years without anyone discussing.

Possessors usually do not have a problem in demonstrating their 

lawful possession as it is already recognized by municipal authority. 

The Municipality, moved by raising more taxes, gives them a cadastral number, but they do not have a public statement even though the 

land belongs to the Nation and, because of that, are not prescriptible. 

While there is not a Civil Code, a binding procedure that calls SRA to 

defeat Nation’s interests when there is no public records at the RPP, a 
USDA Forest Service Proceedings RMRS-P-67. 2013 99

Buying Land for Conservation Purposes in Sonora, Mexico Perez-Weil and Bravo

judge can assess the requirements specified by that Code and grants 

land to the possessors applicant by prescription.

Having to deal with the owner through the different negotiation obstacles, Naturalia with NJP support acquired the group of four ranches, 

34,904.87 acres in total size, on July 2, 2007, with an agrarian title 

and civil regularization. This purchase increased the area of the NJR 

three times. In 2008 and 2010, a search was made of the Sahuaripa’s 

Municipal RPP and the Cadastre and Registration Institute of Sonora 

State (ICRESON) in Hermosillo, Sonora, to determine the status of 

each of the properties next to the NJR. This search was necessary 

because of the interest in acquiring some of these properties, and, 

therefore, knowledge of their legal status was necessary to determine 

that the lands were free from any lien or trial. Lands were removed 

from consideration when they lacked any kind of agrarian or civil 

regularization; were titled twice or had unfinished agrarian and civil 

procedures; were regularized by a civil procedure that had on-going 

agrarian procedures; had been decreed officially as public property; 

had prescription rights with non-viable limits or boundaries on reality; or lacked a will, were mortgaged without any kind of record, or 

had a lien.

This scenario led to negotiations with owners or possessors who 

had the “best titles,” that is, at least had a civil title. This purchasing 

strategy has led to obtaining those ranches that were best located, 

favored reserve integrity, allowed for better management, and whose 

owners had wanted to sell. The last acquisitions were done on 2011. 

El Carricito, an area of 393.86 ac, was acquired as a possession in 

February so it had to be regularized though a civil procedure that 

was obtained by a favorable judgment on October of the same year. 

The property’s use had been granted for life to her possessor to avoid 

destroying Sierra’s culture and taking into account that livestock activity would be reduced considerably. The acquisition of Los Tezotaz’ 

5,873.62 ac followed in August. This ranch was partially titled because 

the will that entitled to the seller as possessor of the land referred to 

the “approximate surface” that coincided with the acreage surface for 

which a property tax was paid at the Municipality. It was necessary, 

however, to make a surface rectification. Finally, a small paddock 

of 171.66 ac was purchased. This paddock was inside Zetazora, one 

of the ranches already titled in Naturalia’s favor and with an agrarian title. Even though the paddock was Naturalia’s property, it had 

to be paid for to avoid a conflict by a possession that existed before 

Zetazora was purchased. Moreover, the paddock was a possession 

of the same person who sold Los Tezotaz; an opportunity of solving 

two negotiations in just one operation.

On-Going and Future Actions

Steps are being carried out to obtain an agrarian title from SRA for 

a ranch of about 5,239.17 acres called Los Pavos de Abajo or Cajón 

Babizoso. We are also waiting for the owner of the only legally titled 

ranch, Tinaja Ahogadora, to complete a divorce process so that he 

can dispose of the property as sole owner. In parallel, since about 

a year ago, we have been correcting the previous justification study 

through which Naturalia had proposed as NPA to the NJR under the 

category of a “flora and fauna protection area” to Environmental 

Ministry. The main purpose of this request is to protect the area from 

one of its principal threats—mining—since planned gold extraction 

is becoming more imminent.

The Mexican Constitution, in its 27th Article, states that mining 

veins, ledges, masses, or deposits are national property. If it is accepted that both mining and environmental protection are public 

utility causes, it is desirable to avoid getting into a debate of which 

one of them should prevail. Moreover, an NPA establishment will 

allow a guaranteed federal zone or a riverbank strip of 10 m wide 

measured starting from the highest water river level destination of 

Aros’ riverbed on the adjacent portion to the NJR for conservation 

purposes. This is because once the area had a formal decree, the 

National Commission of Natural Protected Areas would be able to 

request it for its own destination. At this time, Naturalia could only 

request it as a concession that must be paid periodically, being unaffordable on the long term.

Establishing NJR as an NPA perpetuates the area purposes even 

if Naturalia is absent. The actions of NPA administered by public 

administration are compensated because there is only one owner 

who is committed to conservation purposes. The owner would be in 

charge of its management having the advantage of possibly obtaining more financial resources, capturing greater economic incentives, 

and having access to some technical support and participation of the 

regional conservation strategy.

Conclusions

Even though there have been advances in regulations for encouraging participation of organization and civil society in environmental 

conservation matters, irregularities confronting land tenants are still an 

impediment for its consolidation. A clear title is required for developing 

or carrying out any environmental conservation activities. However, 

clear titles do not always exist, so buying possessions always carries 

a risk that not everybody is willing to afford or finance. Meanwhile, 

while inefficiency in the regulations of land possessions often forces 

an immediate review, it also allows working with local community 

since a different perspective, promoting that way an important social 

change.

While land purchases may not be the only means nor be isolated from 

other mechanisms to integrate governmental participation at almost 

any level, the purchases allow participation by the local population in 

an area’s conservation. While the acquired properties may be a core 

area, their connection though biological corridors should be resolved 

with more than one organization and with people’s support. Land 

purchases should be conceived as an additional protective layer that 

becomes a whole piece with all other environmental efforts directed 

to ecosystems’ and priority regional species’ conservation efforts. 

References

Arriaga, L., Espinoza, J., Aguilar, C., Martínez, E., Gómez, L., and Loa, E. 

2000. Regiones Terrestres Prioritarias de México. México: Comisión 

Nacional para el Uso y Conocimiento de la Biodiversidad.

Brown, D., and López, C. 2001. Borderland Jaguars. Salt Lake City: The 

University of Utah Press.

Gutiérrez, M., Bacmeister, A., Ortiz, G.,and Montesionos, J. A. 2002. Herramientas legales para la conservación de tierras privadas y sociales en 

México. México: Pronatura, A.C.

Roldán, M., Carminati, A., Biganzoli, F., and Paruelo, J. M. 2010. Las reservas 

privadas son efectivas para conservar las propiedades de los ecosistemas? 

Ecología Austral. 20(2):185-199.

Consulted Regulations

Constitución Política de los Estados Unidos Mexicanos.

México, Ley General del Equilibrio Ecológico y la Protección al Ambiente, Diario 

Oficial de la Federación, 28 de enero de 1988.

México, Ley General del Equilibrio Ecológico y la Protección al Ambiente, Decreto 

que reforma, adiciona y deroga diversas disposiciones de la, Diario Oficial de 

la Federación, 13 de diciembre de 1996.
100 USDA Forest Service Proceedings RMRS-P-67. 2013

Perez-Weil and Bravo Buying Land for Conservation Purposes in Sonora, Mexico

México, Reglamento de la Ley General del Equilibrio Ecológico y la Protección al Ambiente en Materia de Áreas Naturales Protegidas, Diario Oficial 

de la Federación, 30 de noviembre de 2000.

México, Ley General de Vida Silvestre, Diario Oficial de la Federación, 3 

de julio de 2000.

México, Ley Agraria, Diario Oficial de la Federación, 26 de febrero de 1992.

México, Reglamento de la Ley Agraria en Materia de Ordenamiento de la 

Propiedad Rural, Diario Oficial de la Federación, 4 de enero de 1996.

México, Ley de Aguas Nacionales, Decreto por el que se reforman, adicionan 

y derogan diversas disposiciones de la, Diario Oficial de la Federación, 

29 de abril de 2004.

México, Ley Minera, Diario Oficial de la Federación, 26 de junio de 1992.

México, Código Civil para el Estado de Sonora, http://www.congresoson.gob.

mx/Leyes_Archivos/doc_1.pdf, fecha de consulta: 20 de abril de 2012.

México, Ley No. 171 Del Equilibrio Ecológico y Protección al Ambiente del 

Estado de Sonora, Boletín Oficial del Gobierno del Estado de Sonora, 25 

de septiembre de 2008.

México, Reglamento de la Ley del Equilibrio Ecológico y Protección al Ambiente del Estado, en materia de Áreas Naturales Protegidas, Boletín Oficial 

del Gobierno del Estado de Sonora, 25 de febrero del 2011.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 101

Amphibian Conservation
102 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 103

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

A Comparison of the Herpetofaunas of Ranchos Los 

Fresnos and El Aribabi in Northern Sonora, Mexico

James C. Rorabaugh

Saint David, Arizona

Jeffrey M. Servoss

U.S. Fish and Wildlife Service, Tucson, Arizona

Valerie L. Boyarski

Arizona Game and Fish Department, Phoenix, Arizona

Erin Fernandez, and Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Carlos Robles Elías

Rancho El Aribabi, Imuris, Sonora, Mexico

Kevin E. Bonine

Department of Ecology and Evolutionary Biology and School of Natural Resources, University of Arizona, 

Tucson, Arizona

Abstract—To compare and contrast herpetofaunas at Ranchos Los Fresnos and El Aribabi in north-central 

Sonora, México, we conducted herpetological surveys during 2006-2011, contacted others working in these 

two areas, and queried 27 museums and collections for specimens collected at or near these ranches. Based 

on this work, nine and seven amphibian, and 27 and 24 reptile species are known to occur at Ranchos 

El Aribabi and Los Fresnos, respectively. Significant findings at Los Fresnos include presence of presumed 

Sonora tiger salamanders (Ambystoma mavortium stebbinsi), of which there are only three localities in 

México; Arizona treefrog (Hyla wrightorum) at several sites; and Mexican gartersnake (Thamnophis eques); 

and at El Aribabi, Neotropical whipsnake (Coluber mentovarius), a range extension of 144 km, Tarahumara 

salamander (Ambystoma rosaceum) near the northern edge of its range, and Mexican gartersnake. Both 

ranches support non-native American bullfrogs (Lithobates catesbeianus) and non-native fishes. Non-native 

crayfish (Orconectes virilis) are also present at Rancho Los Fresnos. In total, 46 species of amphibians and 

reptiles were found, 22 of which occur on both ranches. Rancho Los Fresnos supports high grassland species 

not found at El Aribabi, while a number of typically Sonoran Desert or foothills thornscrub species found at 

El Aribabi were not encountered at Los Fresnos.

Introduction

	 Sonora is the second largest of the Mexican states and extremely

diversetopographicallyandinregardtobiologicalcommunities.Mesic

mixed conifer forests in the mountains of the northeast contrast with

the aridGranDesierto deAltar ofthe northwest and tropical deciduous

forests of the southeast. Sonora’s herpetofauna reflects this diversity

in that 37 and 141 species of amphibians and reptiles, respectively,

have been documented in the state exclusive ofitsislands(Rorabaugh

2008). However, many areas of Sonora are poorly explored herpetologically and focused work often reveals significant range extensions, unexpected faunal associations, or other relationships. From

2006-2011, we conducted focused herpetological inventories at two

ranches in northeastern Sonora: Rancho Los Fresnos and Rancho El

Aribabi.

Study Sites 

	 The 4,050-ha Rancho Los Fresnos, located about 35 km northnorthwest ofCananea on theU.S.-Mexico border(figs. 1, 2), is owned

by Naturalia, a conservation non-governmental organization. The

ranch is characterized by rolling plains grasslands at 1,460-1,600 m,

and is notable not only for its intact plains grassland community, but

also its well-developed ciénegas and riparian corridors along Portrero

del Alamos, Arroyo Los Fresnos, Arroyo Los Alisos, Agua Dulce,

and other drainages (Varela-Romero et al. 1992). Oak savannahs in

the northeastern portion of the ranch and occasional rocky outcrops

increase the heterogeneity of amphibian and reptile habitats.
104 USDA Forest Service Proceedings RMRS-P-67. 2013

Rorabaugh, Servoss, Boyarski, and others A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . .

Figure 1—Locations of Ranchos El Aribabi and Los Fresnos in the context 

of north-central Sonora, México.

Figure 2—Rancho Los Fresnos, approximately 35 km NNW of Cananea, 

Sonora, México.

Figure 3—Rancho El Aribabi, Approximately 16-31 km ENE of Imuris,

Sonora, México.

	 Rancho El Aribabi, owned by the Robles family, extends over

13,000 ha and islocated approximately 16-31 km northeast of Imuris

(figs. 1, 3). El Aribabi includes biotic communities ranging from Sonoran desertscrub at the very lowest elevations (960 m) to mesquite

grasslands, Madrean oak woodlands, and pine-oak woodlands at the

highest elevations(about2,000m).TheRíoCocospera,whichincludes

an impressive riparian corridor and associated ciénega, runs through

the western portion of the ranch. Much of our work was focused on

the approximate 4,046 ha managed by Carlos Robles.

	 NaturaliarecentlyacquiredRanchoLosFresnosandwasinterestedin

inventoryingthe ranch’snaturalresources,includingitsherpetofauna.

The U.S. Fish and Wildlife Service (USFWS) and Arizona Game

and Fish Department (AGFD) were interested in the ranch as well,

because of ongoing conservation and species recovery activities in

the San Rafael Valley, Arizona, and the potential for complementary

conservation at Los Fresnos, which is located in the southern portion of that valley. Carlos Robles was interested in pursuing federal

reserve status for Rancho El Aribabi through México’s federal La

Comisión Nacional de Áreas Naturales Protegidas (CONANP), and

describing the ranch’s biological diversity was one of the first steps

in that process. That federal reserve status was assigned to the ranch

in May of 2011.

Materials and Methods

	 Weusedthe followingthreemethodstoinvestigate theherpetofauna

of Ranchos Los Fresnos and El Aribabi.

Herpetological Inventories

	 Rancho Los Fresnos was visited 22-25 May, 23-25August, and 3-5

October 2006; and 17-19 September 2007. An initial reconnaissance

was also made 20-21 December 2005 and additional herpetological

recordswereobtainedduringamphibianworkshopsheld19-21August

2008, 7-9 July 2009, and 20-23 September 2010; and a bat workshop

held 6-10 June 2011.RanchoElAribabiwas visited 10-12May, 20-22

September, and 6-8 October 2006; 20-22 April, 19-21 June, and 7-9

August2007;14-18July2008; and18-20November2011.Inventories

at both ranchesincluded day and night visual encountersurveys, driving roads (especially just before and after dark), listening for calling

amphibians, and hoop nets and Gee minnow traps placed in streams

and pondsto capture tadpoles, turtles, and gartersnakes(Thamnophis 

sp.). We also employed a 10-m seine at cattle tanks at Los Fresnos to

survey for amphibians.Wemade incidentalobservations of native and

non-native fishes and introduced invertebrates, which can influence

distributionorpresenceofnative riparianor aquaticvertebrates(Rosen

and Schwalbe 2002). Cover board arrays were used on a limited basis

at Rancho El Aribabi, and at that ranch, concrete cattleguard boxes

were found to act as opportunistic pit fall traps. One of us (C. Robles

Elías)frequentlyvisitedElAribabiandperiodicallyphoto-documented

amphibians and reptiles in the course of other work. Due to good
USDA Forest Service Proceedings RMRS-P-67. 2013 105

A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . . Rorabaugh, Servoss, Boyarski, and others

accessibility, we were able to sample all biotic communities and most

areas of the ranch at Los Fresnos; however, at Rancho ElAribabi, the

majority of our work was done in the lowlands, and few trips were

made into the higher montane woodlands. Due to poor access, we

did not visit the pine-oak woodlands at Rancho El Aribabi, which

occur in the extreme southeastern corner of the ranch in the Sierra

Azul. Total person hours spent searching for amphibians and reptiles

at Ranchos Los Fresnos and El Aribabi were approximately 381 and

343, respectively. Numbers of minnow trap and hoop net trap days

(24-hour periods) were 17.0 and 49.0 at Rancho Los Fresnos and 5.3

and 28.5 at Rancho El Aribabi, respectively. Six 1-m2 cover boards

were also run for 2 days (12 trap days) along the Río Cocospera at

Rancho El Aribabi. We attempted to photo voucher at least one of

each species found at the ranches. Photo vouchers were deposited at

the University of Arizona herpetological collection (UAZ).

Museum and Literature Searches

	 The museums listed in table 1 were queried for specimen data at

and near Ranchos Los Fresnos and El Aribabi.

Interviews of Other Workers

	 We contactedothersworkinginthe areaofthe tworanchesregarding

their herpetological observations, including for Rancho Los Fresnos,

biologistswithBIDA(BiodiversidadyDesarrolloArmónico),students

from the Universidad de Sonora, Sally Stefferud (retired U.S. Fish

and Wildlife Service), and Peter Warren (The Nature Conservancy);

and for Rancho El Aribabi, Dale Turner (The Nature Conservancy),

Sergio Avila, Trevor Hare, and Tom Van Devender (Sky Island Alliance), and Tom Wood (Southeastern Arizona Bird Observatory).

Results

	 Our investigations, including herpetological inventories, review

of museum collections, and interviews with others, documented nine

and seven species of amphibians, and 27 and 24 species of reptiles at

Ranchos El Aribabi and Los Fresnos, respectively (table 2). In total,

46 amphibian and reptile species were found at Ranchos Los Fresnos

and ElAribabi, of which 22 occurred at both ranches(table 2), resulting

in a Jaccard Similarity Index (Baroni-Urbani 1980) of 22/46 = 0.48.

All species were documented via our own observations, except for

six found by others working in the area. No museum collections were

definitively from the two ranches, although KUNHM specimens

collected primarily by Joseph R. Alcorn in 1957 and 1958 at “9 mi

NNE of Imuris” (table 3) may have come from Rancho El Aribabi.

Alcorn’s locality plots in the Sierra Los Pinitos were about half way

between the Río Bambuto and the Río Cocospera; the latter river is

on Rancho El Aribabi. The collections include riparian species, such

asMexicangartersnake (Thamnophis eques),Sonoranmud turtle, and

Lowland leopard frog (Lithobates yavapaiensis); however, habitat

for these species is currently absent at 9 mi NNE of Imuris, and it

seems likely that Alcorn collected either at the Río Bambuto or the

Río Cocospera. Our efforts to locate Alcorn’s field notes or other

clarification of the collection locality have been unproductive. As a

result, for the purposes of tables 2 and 3we have assumed these collections were made near but not within Rancho El Aribabi. Species

not documented by us at El Aribabi or Los Fresnos, but which were

found in museum collections or were reported by others from localities in contiguous habitats near the two ranches are noted in table 3.

	 Bothrancheswere foundtosupportintroducedpredators.AtRancho

Los Fresnos, we found American bullfrog (Lithobates catesbeianus) to be abundant and widespread, as well as introduced crayfish

(Orconectes virilis), Green sunfish (Lepomis cyanellus), and Black

bullhead (Ameiurus melas), which were locally abundant. The native

Longfin dace (Agosia chrysogaster) was only abundant in Arroyo

Los Fresnos. American bullfrog was found in the Río Cocospera at

Rancho El Aribabi, but only appeared to be abundant in the pools at

the ciénega, which is also the only area where we found tadpoles of

that species. We found introduced Mosquitofish (Gambusia affinis)

and Green sunfish in the Río Cocospera at Rancho El Aribabi during

2006-08, and Largemouth bass (Micropterus salmoides) was found

in 2011. However, we also found native dace (Agosia sp.), Sonora

chub (Gila ditaenia), and topminnow (Poeciliopsis sp.).

Discussion

Faunal Differentiation and Causal Factors

	 The herpetofauna of Rancho Los Fresnos is in part a grassland

assemblage, as reflected in the presence of species such as Tiger

Table 1—Herpetological museums queried for specimens collected 

at or near Ranchos El Aribabi and Los Fresnos.

Museum 

acronym Full name

AMNH American Museum of Natural History

ASU Arizona State University Herpetology Collection

BMNH Bishop Museum of Natural History

BYU Brigham Young University

CAS California Academy of Science

CUMV Cornell University Museum of Vertebrates

KUNHM University of Kansas Natural History Museum

INHS Illinois Natural History Survey

LACM Los Angeles County Museum

LSUMNH Louisiana State University Museum of Natural History

MCZ Harvard University Museum of Comparative Zoology

MSUM Michigan State University Museum

MVZ Museum of Vertebrate Zoology

PSM Slater Museum of Natural History

ROM Royal Ontario Museum

SDNHM San Diego Natural History Museum

SU Stanford University collections (housed at CAS)

TNHC University of Texas at Austin

UAZ University of Arizona Herpetology Collection

UCM University of Colorado Museum at Boulder

UIMNH University of Illinois Museum of Natural History

UMMZ University of Michigan Museum of Zoology

UNAM Universidad Nacional Autónoma de México

USNM National Museum of Natural History

UTA University of Texas at Arlington

UTEP University of Texas at El Paso
106 USDA Forest Service Proceedings RMRS-P-67. 2013

Rorabaugh, Servoss, Boyarski, and others A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . .

Table 2—Amphibian and reptile species documented by vegetation community type at Ranchos Los Fresnos and El Aribabi. “UAZ” indicates the 

species is photo-vouchered in the University of Arizona Herpetological Collection. If no voucher exists, occurrence is shown with an “X”. All 

species documented by the authors unless indicated in the footnotes.

Species Rancho El Aribabi Rancho Los Fresnos

Amphibians Vegetation community type Vegetation community type

Caudata Sonoran 

Desert

Mesquite 

Grassland

Riparian/

Ciénega

Mesquite/

Oak Savanna

Oak 

Savanna

Plains

Grassland

Plains 

Grassland/

Scattered 

Oaks

Riparian Ciénega

Ambystoma mavortium Xa Xb

Ambystoma rosaceum UAZ UAZc

Anura

Anaxyrus punctatus X UAZ X X UAZ X X

Anaxyrus woodhousei UAZ X UAZ X X

Hyla arenicolor UAZ X UAZ UAZ

Hyla wrightorum UAZ UAZ

Lithobates catesbeianus UAZ X UAZ X

Lithobates yavapaiensis X X UAZ

Incilius alvarius UAZ X

Scaphiopus couchii X UAZ

Spea multiplicata UAZ X UAZ X

Reptiles

Testudines

Kinosternon sonoriense UAZ X UAZ

Terrapene ornata UAZ X UAZ

Squamata - Lizards

Aspidoscelis burti UAZ

Aspidoscelis sonorae X X X X X

Coleonyx variegatus UAZ

Crotaphytus collaris UAZ

Elgaria kingii UAZ X UAZ 

Heloderma suspectum X4 UAZ

Holbrookia elegans X X UAZ X UAZ

Plestiodon callicephalus X

Phrynosoma hernandesi UAZ X

Sceloporus clarkii X UAZ X X X X

Sceloporus jarrovii UAZ 

Sceloporus slevini UAZ X

Urosaurus ornatus X UAZ X UAZ X X

Squamata - Snakes

Coluber bilineatus X

Coluber flagellum X X X

Coluber mentovarius UAZ

Crotalus atrox UAZ X UAZ
USDA Forest Service Proceedings RMRS-P-67. 2013 107

A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . . Rorabaugh, Servoss, Boyarski, and others

Crotalus molossus UAZ UAZ X

Crotalus scutulatus Xe UAZ

Crotalus tigris UAZ

Diadophis punctatus UAZ

Heterodon kennerlyi Xf

Hypsiglena chlorophaea UAZ UAZ UAZ UAZ

Micruroides euryxanthus UAZ

Pituophis catenifer UAZ UAZ UAZ

Rhinocheilus lecontei UAZ Xf

Sonora semiannulata Xb

Salvadora grahamiae UAZ

Salvadora hexalepis Xd UAZ

Senticolis triaspis UAZ

Thamnophis cyrtopsis UAZ UAZ UAZ

Thamnophis eques Xg UAZ UAZ Xb

Trimorphodon lambda UAZ

a

In a cattle tank

b

Reported by S. Stefferud

c

Documented by R. Serraglio, S. Avila, and M. Quigley

d

Reported by D. Turner

e

Reported by T. Wood

f

Photographed by students from the Universidad de Sonora, Hermosillo

g

Reported by T. Hare

salamander (Ambystoma mavortium), Mexican hog-nosed snake

(Heterodon kennerlyi), and Slevin’s bunchgrass lizard (Sceloporus 

slevini); however, many of the upland species occur elsewhere in a

variety of biotic communities, and many of the riparian forms (e.g.

Canyontreefrog[Hyla arenicolor],Sonoranmudturtle,Black-necked

gartersnake [Thamnophis cyrtopsis], andMexican gartersnake) occur

over a wide elevational range in a variety of community types. The

presence of Yarrow’s spiny lizard (Sceloporus jarrovii), a montane

species, is due to the rocky outcrops and oak savanna in the northeastern portion of Rancho Los Fresnos, which is part of the bajada

of the Huachuca Mountains in Arizona.

	 Species documented at Rancho El Aribabi reflect a greater range

of community types, particularly emphasized by species more characteristic of lower elevations than are found at Rancho Los Fresnos

(e.g. Couch’s spadefoot [Scaphiopus couchii] and Gila monster

[Heloderma suspectum]). The influence of the Sonoran Desert and

associated foothillsthornscrub communitiesisin evidence at Rancho

ElAribabi in the presence of Western banded gecko (Coleonyx variegatus), and species found nearby to the west in contiguous habitats,

such as Sonoran green toad (Anaxyrus retiformis), Sinaloa toad

(Incilius mazatlanensis), and Lowland burrowing treefrog (Smilisca 

fodiens; table 3). Rancho ElAribabi also exhibits a subtropical faunal

influence with the presence of Tarahumara salamander (Ambystoma 

rosaceum), Neotropical whipsnake (Coluber mentovarius), and

Green ratsnake (Senticolis triaspis). If we had been able to survey

the higher elevations of the Sierra Azul more thoroughly, including

Table 2—Cont.

Table 3—Species present in museum collections or reported 

by others from localities close to Ranchos Los Fresnos or El 

Aribabi and in contiguous habitats. 

Species Locality and Museum

Rancho Los Fresnos

Lithobates chiricahuensis Bear Canyon, Huachuca 

Mountains, AZ, UAZ 24754

Lithobates yavapaiensis 2 mi NNE Sunnyside, AZ, 

UAZ 20294, 20273

Diadophis punctatus Southern end of San Rafael 

Valley, AZ, JCR 0023

Rancho El Aribabi

Anaxyrus retiformis 1 mi S Imuris, UAZ 31581-3

Incilius mazatlanenis 35 mi S Nogales, UIMNH 

24455

Smilisca fodiens 1.5 mi N Imuris on Hwy 15, 

UAZ 31555

Phrynosoma solare 9 mi NNE Imuris, KUNHM 

50729

Sceloporus virgatus Sierra Azul, MABA sontrv-1418

Crotalus willardi

Sierra Azul (Campbell and 

Lamar 2004, MABA sontrv-1300)
108 USDA Forest Service Proceedings RMRS-P-67. 2013

Rorabaugh, Servoss, Boyarski, and others A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . .

the pine-oak woodlands, we no doubt would have added species and

broadened the ecologicalrepresentation ofspecies upslope atRancho

El Aribabi. Despite the differences ecologically, our data show that

much overlap exists between the two ranches, as demonstrated by a

Jaccard Similarity Index of 0.48.

Significant Observations and Range 

Extensions

	 On 25August 1990,S.Stefferud,P.Warren, and others documented

a larval Tiger salamander at Rancho Los Fresnos (S. Stefferud’s

field notes). Based on its location, the former was almost certainly a

Sonora tigersalamander (Ambystoma mavortium stebbinsi), which is

widely distributed just to the north of Rancho Los Fresnos in the San

Rafael Valley in Arizona (Collins et al. 1988). The salamander was

found in ciénega pools in Portrero del Alamo, and assuming it was a

Sonora tiger salamander, this would be the only record of breeding

in that subspecies outside of artificial impoundments (U.S. Fish and

Wildlife Service 2002), and may represent a natural habitat used by

thesesalamandersbeforeEuropeansettlement.Wealsofoundbreeding

Tiger salamanders that we presumed to be Sonora tiger salamanders

in a cattle impoundment in the northeastern portion of the ranch and

at another cattle tank on the ranch just west of Rancho Los Fresnos.

But because morphologically similar introduced Barred tiger salamanders (A. t. mavortium) have been found in some cattle tanks in

the vicinity of Parker Canyon Lake, about 18 km north-northwest of

theLos Fresnos headquarters, we cannotrule out that the salamanders

at Los Fresnos are that subspecies. In 1990, S. Stefferud, P. Warren,

and others also found an Arizona treefrog (Hyla wrightorum). We 

too found this species at several localities at Los Fresnos, and within

Sonora those localities represent a range extension of approximately

235 km north-northwest from the closest record at El Chorro, 8 km

northeast of NacoriChico, Sonora (UAZ 45595), although the closest

out-of-country record is ca. 13 km north in Arizona (MaldonadoLeal et al. 2009). The Los Fresnos localities are also unusual in that

they are from aquatic sites in valley grasslands, whereas in Cochise

County, Arizona, and sites elsewhere in Sonora, the Arizona treefrog

is known as a montane or bajada species in woodland communities

or from meadows within woodlands.

	 Although we did not find them, leopard frogs (Lithobates chiricahuensis and/or L. yavapaiensis) were likely present in the streams

and ciénegas at Los Fresnos historically, as they occurred both in

drainages upstream in the nearby Huachuca Mountains (table 3), as

well as downstream in the San Pedro River (Clarkson and Rorabaugh

1989). No leopard frogs,American bullfrogs, or crayfish were noted

by S. Stefferud in her field notes during her visit to the Los Fresnos

in 1990, despite detailed observations on other amphibians, reptiles,

fishes, and invertebrates. The only non-native species noted by S.

Stefferud and others (Varela-Romero et al. 1992, S. Stefferud’s field

notes) were Mosquitofish (not found by us at Rancho Los Fresnos)

and Green sunfish. It islikely that leopard frogs,American bullfrogs,

and crayfish were scarce or absent in 1990. Sonoran mud turtles were

present and often abundant in all ofthe permanent waters we sampled

at Rancho Los Fresnos, and were also commonly found in the Río

Cocospera at Rancho ElAribabi, thus effects of introduced predators

on this species appeared to be minimal.

	 Mexican gartersnakes were found at both ranches.At Los Fresnos,

four were observed along or near Arroyo Los Fresnos, where nonnative predator densities appeared low and there was an abundance

of native Longfin dace. As evidenced by flood debris in the trees

and a severe flood we witnessed on the evening of 24 August 2006,

this arroyo is subject to periodic scouring, which tends to suppress

non-native fish (Minckley and Meffe 1987), American bullfrog, and

crayfish (Rorabaugh pers. obs.; also see Rosen and Schwalbe 2002)

populations.S.Stefferud(fieldnotes)observedaMexicangartersnake

in theLos FresnosCiénega in 1990.We observed several Thamnophis 

in that area, but we were unable to capture or identify them to species.

Mexican gartersnakes may be persisting in the ciénega despite the

presence of non-native predators. A single Mexican gartersnake was

observed and photographed by Trevor Hare (Sky Island Alliance)

along the Río Cocospera at Rancho El Aribabi. The only other gartersnakes observed atElAribabiwere twoBlack-necked gartersnakes

(T. cyrtopsis), also found along the Río Cocospera. The apparent low

densityofgartersnakeswasunexpected,becausepredatordensities are

relatively low outside of the ciénega and the habitat appearssuitable.

Eight Mexican gartersnakes collected by Alcorn at “9 miles NNE

Imuris” (table 3) may have been collected near or perhaps at the Río

Cocospera (see discussion above in the results section).

	 At Rancho El Aribabi, Sergio Avila and Carlos Robles Elías photo

documented Neotropical whipsnakes on the northwestern slopes of

the SierraAzul (e.g. UAZ 56736-PSV), which extends the range 144

km north-northwest of the closest known locality (21 km northeast

of Mazocahui on Highway 20, Sonora; Rorabaugh et al. 2009) and

placesthislargely tropicalspecies within 57 kmoftheArizona border

(Avila et al. 2008). Most records of this species in Sonora are from

the southeastern portion ofthe state,south ofHighway 16 (Rorabaugh

et al. 2009). We also documented Tarahumara salamandersfrom two

localities onthenorthwesternslope, aswell asthenorthwesternbajada

of the SierraAzul (table 2); these localities are near the northern edge

of the distribution of this species.

Species Likely to Be Found With Additional 

Effort

	 Figures 4 and 5 plot search effort (days) against cumulative

number of species found for Ranchos Los Fresnos and El Aribabi,

respectively. Both figures show that numbers of species found have

plateaued, suggesting that adding species to the list will be time

consuming and not very productive. Nonetheless, we believe several

species not yet observed remain extant on the ranches. Speciesfound

nearby in similar habitats are listed in table 3. Of those, we believe

all may occur on the ranches, with the exception of the Chiricahua

leopard frog (Lithobates chiricahuensis) and Lowland leopard frog at

Rancho Los Fresnos for the reasons discussed above. Other species

that could be found include, for Rancho El Aribabi: Western narrowmouthed toad (Gastrophryne olivacea), Barking frog (Craugastor 

augusti), Morafka’s desert tortoise (Gopherus morafkai), Thornscrub

hook-nosed snake (Gyalopion quadrangulare), Brown vinesnake

(Oxybelis aeneus), and Chihuahuan mountain kingsnake (Lampropeltis knoblochi); for Rancho Los Fresnos: Desert grassland whiptail

(Aspidoscelis uniparens),Ring-necked snake (Diadophis punctatus),

and Chihuahuan hook-nosed snake (Gyalopion canum), and for both

ranches:Black-headedsnakes(Tantillassp.), andCommonkingsnake

(Lampropeltis getula).

Acknowledgments

We gratefully thank staff of the natural history museums listed

in “Museum and Literature Searches” in the results section above

for sharing museum records and discussing specimens. Juan Carlos

Bravo and Gerardo Carreón of Naturalia provided access to and use

of facilities at Rancho Los Fresnos, and we thank the Robles family
USDA Forest Service Proceedings RMRS-P-67. 2013 109

A Comparison of the Herpetofaunas of Ranchos Los Fresnos . . . Rorabaugh, Servoss, Boyarski, and others

Figure 5—Species accumulation curve, Rancho El Aribabi.

for their hospitality at Rancho El Aribabi. Sergio Avila, students of

Kevin Bonine’s herpetology class at University of Arizona, students

from the Universidad de Sonora, Antonio Esquer, Trevor Hare,

Thomas R. Jones, Baruk Maldonado-Leal, Gricelda Meraz, Robert

Mesta, Mike Quigley, Scott Richardson, Eduardo López Saavedra,

Sally Stefferud, Robert Villa, Peter Warren, Tom Woods, and the

vaqueros at Rancho ElAribabishared their observations with us and/

or assisted with logistics in the field. Dale Turner and Don Swann

graciously reviewed a draft of this paper.

References 

Avila,Sergio,Robles-Elías,Carlos;andRorabaugh,JamesC.2008. Geographic

distribution:Coluber(=Masticophis) mentovarius (NeotropicalWhipsnake).

México: Sonora. Herpetological Review 39(3):370.

Baroni-Urbani, Cesare. 1980. A statistical table for the degree of coexistence

between two species. Oecologia 44:287-298.

Campbell,JonathanA.; and Lamar,WilliamW. 2004.The VenomousReptiles

of the Western Hemisphere, Volumes 1 and 2. Comstock Publishing Associates, Ithaca, NY.

Clarkson, Robert W.; and Rorabaugh, James C. 1989. Status of leopard frogs

(Rana pipiens Complex) in Arizona and southeastern California. The

Southwestern Naturalist 34(4):531-538.

Collins, James P.; Jones, Thomas R.; and Berna, Howard J. 1988. Conserving

genetically distinctive populations: the case ofthe Huachuca tigersalamander (Ambystoma tigrinum stebbinsi Lowe). Pp. 45-53 In R.C. Szaro, K.E.

Severson, and D.R. Patton (Tech. Coords.). Management of amphibians,

reptiles and small mammals in North America. Gen. Tech. Rep. RM-166.

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest

and Range Experiment Station. Fort Collins, CO.

Maldonado-Leal, Baruk G., Warren, Peter L.; Jones, Thomas R.; Boyarksi,

Valerie L.; and Rorabaugh, James C. 2009. Hyla wrightorum (Arizona

treefrog). México: Sonora. Herpetological Review 40(1):108.

Minckley, Wendell L., and Meffe, Gary K. 1987. Differential selection by

flooding in stream-fish communities of the arid American Southwest. Pp.

93-299 In W.J.Mathews andD.C.Heins(Eds.).Community andEvolutionary Ecology of North American Stream Fishes. University of Oklahoma

Press, Norman and London.

Rorabaugh, James C. 2008. An introduction to the herpetofauna of mainland

Sonora,México,with comments on conservation andmanagement.Journal

of the Arizona-Nevada Academy of Science 40(1):20-65.

Rorabaugh,James;Avila,Sergio;Robles-Elías,Carlos; andFerguson,George.

2009. An addition to the 100-MileCircle: Neotropical whipsnake (Coluber 

mentovarius). Sonoran Herpetologist 22(1):2-5.

Rosen, Philip C., and Schwalbe, Cecil R. 2002. Widespread effects of introduced species on reptiles and amphibians in the Sonoran Desert region.

Pp. 220-240 In B. Tellman (ed.). Invasive Exotic Species in the Sonoran

Region. University of Arizona Press, Tucson.

U.S. Fish and Wildlife Service. 2002. Sonora tiger salamander recovery plan.

U.S. Fish and Wildlife Service, Phoenix, AZ.

Varela-Romero, Alejandro; Galindo-Duarte, C.; Saucedo-Monarque, E.;

[and others]. 1992. Re-discovery of Gila intermedia and G. purpurea in 

northern Sonora, Mexico. Pp. 33 In D.A. Hendrickson (ed.). Proceedings

of the Desert Fishes Council. Volumes XXII and XXIII, 1990 and 1991

Annual Symposia, and Index for Volumes XVI through XXIII, Desert

Fishes Council, Bishop, CA.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Figure 4—Species accumulation curve, Rancho Los Fresnos.
110 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Comparison of Preliminary Herpetofaunas of the 

Sierras la Madera (Oposura) and Bacadéhuachi with the 

Mainland Sierra Madre Occidental in Sonora, Mexico

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Erik F. Enderson

Drylands Institute, Tucson, Arizona

Dale S. Turner

The Nature Conservancy, Tucson, Arizona

Roberto A. Villa

Tucson Herpetological Society, Tucson, Arizona

Stephen F. Hale

EcoPlan Associates, Inc., Mesa, Arizona

George M. Ferguson

The University of Arizona, Tucson, Arizona

Charles Hedgcock

Sky Island Alliance, Tucson, Arizona

Abstract—Amphibians and reptiles were observed in the Sierra La Madera (59 species), an isolated 

Sky Island mountain range, and the Sierra Bacadéhuachi (30 species), the westernmost mountain range 

in the Sierra Madre Occidental (SMO) range in east-central Sonora. These preliminary herpetofaunas 

were compared with the herpetofauna of the Yécora area in eastern Sonora in the main SMO, where 92 

species are known from the Río Yaqui to the Chihuahua border. The Yécora area, as we have defined 

it, extends from the Río Yaqui to the Chihuahua border along Mexico Federal Highway 16 and other 

areas accessible from it. Seven species in the Sierra la Madera are exclusive of the SMO fauna. Sky 

Island faunas are dominated by Madrean species, but also include species with tropical, desert, and 

northern temperate biotic affinities. Although the herpetofaunas of many Sky Island ranges in Sonora 

are not well documented, it is clear that the SMO fauna is much more diverse than any of them.

Introduction

	 The fauna of the southwestern United States is famous for its

diversity of amphibians and reptiles. The Madrean Archipelago is

the area of isolated Sky Island (SI) mountain ranges between the

northern Sierra Madre Occidental (SMO) in Sonora and Chihuahua

and the southern edge of the Colorado Plateau (Mogollon Mountains

and escarpment of the Mogollon Rim) in New Mexico and Arizona.

High species richness in the region and each Sky Island is related to

the convergence of five biotic provinces with diverse topography and

habitatsthat converge in the area: tropical temperate SMO, temperate

Rocky Mountains, Great Plains-Chihuahuan Desert, Sonoran Desert,

and the western Mexico low-lands tropics. The transition between

the New World tropics and the northern temperate zone is at about

29°N in east-central Sonora.

	 In this paper, we summarize the amphibian and reptile faunas of

the Sierra La Madera (SI) and the Sierra Bacadéhuachi (SMO) and

compare them with the extensive herpetofauna of the Yécora area in

east-central Sonora in the main Sierra Madre Occidental (Enderson

and Bezy, unpubl. data).

Study Areas

	 The three study areas are all in east-central Sonora and are either in

the RíoYaqui Basin or the adjacent Río Mayo Basin. Vegetation in all

three areasrangesfrom lowland tropicalskirts of foothillsthornscrub

(FTS) to oak woodland and pine-oak forest at higher elevations. The

Sierra Bacadéhuachi is unique due to the occurrence of desert grassland between FTS and oak woodland in some areas. The Yécora area

is substantially more tropical than the other two areas with tropical
USDA Forest Service Proceedings RMRS-P-67. 2013 111

Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . . Van Devender, Enderson, Turner and others

deciduous forest (TDF) between FTS and oak woodland. We follow

the classification used in Martin et al. (1998) for vegetation names.

Sierra la Madera

	 Sierra laMadera (=Oposura)is aSkyIslandmountainrange located

(ca. 29°51’N to 30°10’N, 109°25’W to 109°42’W; fig. 1) between

Óputo (= Villa Hidalgo) and Huásabas on the Río Bavispe to the

east and between Cumpas and Moctezuma on the Río Moctezuma

to the west (Yanes-Arvayo et al. 2011). Portions of the range are in

the Municipios of Cumpas, Granados, Huásabas, Moctezuma, and

Villa Hidalgo. Elevations range from 615 m at Huásabas and 660 m

at Moctezuma to over 2300 m on the highest peak, an elevational

range of about 1685 meters; Sierra la Madera is one of the higher

ranges in Sonora. Most of the area is privately owned; the higher

elevations are in Fracción V of the Área Natural Protegida AjosBavispe. Cottonwood-willow forest occurs along the Ríos Bavispe

and Moctezuma.

Sierra Bacadéhuachi

SierraBacadéhuachi,thewesternmostmountain range in theSierra

Madre Occidental, islocated east of Bacadéhuachi, Municipio Bacadéhuachi, 34 km east of the Chihuahua border, and 165 km south of

theArizona border (fig. 1). The southern portion of the Bacadéhuachi

is in the Ríos Nácori Chico and Riíto drainages, with the Río Bacadéhuachi to the west. Elevations range from 700 m at Bacadéhuachi

to about 2400 m on the highest peak, an elevational range of 1700

meters.

Yécora Area

Yécora is a town in the SMO in east-central Sonora 42 km east of

the Chihuahua border (fig. 1). Mexican Federal Highway 16 (MEX

16), one of the few highways that crosses the SMO, provides a

west-to-east elevational transect from Río Yaqui (180 m) east to the

Chihuahuaborder eastofYécora.Theherpetofaunaobservationswere

in the Municipios de Yécora, Ónavas, and Soyopa. The vegetation

gradient alongMEX 16 isfoothillsthornscrub (180-550melevation),

tropical deciduousforest (500-1,160 m), oak woodland (1,050-1,700

m), and pine-oak forest (1,220-2,140 m). Grassland occurs in high

valleys (1,200-1,700 m) within oak woodland or pine-oak forest.

Mixed-conifer forest (1,900-2,100 m) is present in Barranca El Salto

on Mesa del Campanero. The elevational range is 1960 m, with 1660

m in the Municipio de Yécora.

Inventories

Sierra La Madera

	 Early collections in Sonora were made along two routes from the

Arizona border that reach different sides of the Sierra la Madera; 1)

Agua Prieta to Fronteras and Esqueda, south through Nacozari de

García andCumpastoMoctezuma,mostly along theRíoMoctezuma,

and 2)Agua Prieta to Esqueda,southeast to the Sierra El Tigre, south

along the Río Bavispe through Angostura and Óputo to Huásabas.

Specimens of Aspidoscelis exsanguis from near Moctezuma were

collected by S. B. Benson in March 1936 and J. E. Simpson in May

1938 (Museum ofComparative Zoology, MCZ, Harvard University).

In the early 1950s, Paul S. Martin, University of Michigan, collected Bufo alvarius from west of Huásabas and Crotalus atrox and

Phrynosoma solare from south of Óputo. Lowe and Woodin (1954)

described Masticophis flagellum ssp. cingulum based on a specimen

from Moctezuma. Michael D. Robinson collected in the Huásabas

area for the University of Arizona Herpetological Collection (UAZ)

in July 1969. As part of his study of the systematics of Aspidoscelis,

John W. Wright collected in the Óputo-Huásabas area for UAZ, and

the LosAngeles County Museum of Natural History (LACM) in July

1963, April 1966, July 1969, July 1974, July 1975, June 1980, and

August 1982. Tod W. Reeder, San Diego State University, collected

in the area for the Museo de Zoología “Alfonso L. Herrera” Facultad

de Ciencias (MZFC, Universidad Autónoma Nacional de México).

Arthur H. Harris and T. Paul Maslin, collected for the University of

Colorado in the area in July 1970. Charles J. Cole collected in the

area for the American Museum of Natural History in June 1988.

	 The other observations and collections in this study were made by

the authors. Hale visited Arroyo la Sauceda in search of Rana tarahumarae on November 1981 and October 1982, the latter trip with

Cecil R. Schwalbe (Hale and Jarchow 1988), and in May 1998 with

Ferguson, Peter A. Holm, and Elizabeth B. Wirt (Hale et al. 1998).

Enderson, Turner, Hale, and Bezy visited the Sierra la Madera in

September 2003 and Enderson and Bezy returned in July 2009.Areas

visited included the microwave towers east-northeast of Cumpas and

RanchoMesaQuemadawestofHuásabas.VanDevender,Turner,Villa,

and Hedgcock visited the Sierra la Madera on a MadreanArchipelago

BiodiversityAssessment(MABA)ExpeditioninJuneandJuly-August

2010, when a group of 38 scientists,researchers,students, volunteers,

and agency biologists visited Ranchos Las Bateas, la Cieneguita,

San Francisco, and Mesa Quemada, and the antenna area.Additional

herpetologists in the group were R. Wayne Van Devender, Stephen

Figure 1—Sierra Bacadéhuachi, the westernmost mountain range in the L. Minter, Matt Nordgren, and Scott J. Trageser.

Sierra Madre Occidental
112 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Enderson, Turner and others Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . .

Sierra Bacadéhuachi

John W. Wright and James L. Patton in July 1967, Michael D.

Robinson in July 1969, and Hale and Schwalbe in October 1982

collected specimensin the Bacadéhuachi area for UAZ. Later Wright

collected in the area for LACM (July 1974) and with Allen E. Greer

for MCZ (July 1975). Van Devender visited the Bacadéhuachi area

in July 2008 and on MABA Expedition trips in June, August, and

September 2011, and March 2012. Turner, Villa, Hale, Ferguson,

and Hedgcock (also March 2012) were on the main expedition of 45

participantsinAugust 2011, alongwithWillLattea,Nordgren,Martín

Villa, and Eric Wallace. Robert Villa and David J. Bygott were on the

September trip. Our recent field activitiesin the Sierra Bacadéhuachi

were centered on Rincón de Guadalupe in Arroyo Campo los Padres

(29°50’40”N 108°58’37”W, 1680 m elevation).

Yécora Area

	 Between 1953 and 2005, amphibians and reptiles were collected

in the Yécora area for UAZ on 33 trips involving 26 people. Primary

collectors were Charles H. Lowe, Hale, Peter A. Holm, Darryl R.

Frost, Julia V. Salmon, Brent E. Martin, and Cecil R. Schwalbe. Van

Devender and Ana L. Reina-Guerrero made observations on 36 trips

to the area from 1995 to 2008. Enderson and Bezy spent 40 field days

investigating the Yécora area herpetofauna from 2004 to 2008.

Methods

	 Amphibians and reptiles were encountered during field searches

and driving roads at night. Most areas were visited at different times

of the day and in different seasons, but most effort was in the summer rainy season in July-September. Additional museum records

were found in online databases such as VertNet (http://vertnet.org/

index.php) and scientific publications, especially Rorabaugh (2008)

and Enderson et al. (2009, 2010). Most historical records and our

new observations and images are available in the MABA database

(Madrean.org). Photovouchers of selected taxa were deposited into

UAZ. Due to permit restrictions, animals were photographed but not

collected.

	 Nomenclature follows Enderson et al. (2010) with a few exceptions.We follow Pauly et al.(2009)in the use ofthe traditional genera

Bufo and Rana, rather than Anaxyrus, Incilius, or Lithobates. We use

Sonora aemula rather than Procinura aemula.

Results 

	 Atotal of 59 species of amphibians(11) and reptiles(48) are known

from Sierra la Madera (table 1). A total of 30 species of amphibians

(9) and reptiles(21) are known fromSierraBacadéhuachi.In contrast,

92 species of amphibians(21) and reptiles(71) are known from along

MEX 16 from the Río Yaqui at Tónichi through the Municipio de

Yécora to the Chihuahua border. None of the species observed in the

study areas are non-native.

Discussion and Conclusions

Biogeography

Several species observed in the Sierra Bacadéhuachi were range

extensions of central SMO animals, notably Bufo occidentalis,

Craugastor tarahumaraensis (Ferguson et al. in press), and Geophis 

dugesi (Lemos-Espinal and Smith 2007; Villa et al. in press). Our

observation of Trimorphodon lambda in pine-oak forest at Rincón

de Guadalupe was the first record in the SMO; T. tau is common in

various habitats in the Yécora area. A suite of typical SMO species,

including Crotalus lepidus, C. willardi, Elgaria kingii, Plestiodon 

callicephalus, Lampropeltis pyromelana, Sceloporus jarrovii, S. 

virgatus, and Senticolis triaspis, are widespread in oak woodland and

pine-oak woodland in Sky Island rangesin Sonora (Rorabaugh 2008;

Enderson et al. 2010) and Arizona (Brennan and Holycross 2006). 

	 A specimen from the Sierra la Madera was a typical L. p. ssp. pyromelana, buttwospecimensseenintheSierraBacadéhuachi appeared

to be L. p. ssp. knoblochi X L. p. ssp. pyromelana intergrades(images

with observations in the MABA database). Intermediate specimens

in thisintermediate site challenge recognition of L. knoblochi assuggested by Lemos-Espinal et al. (2003). A recent molecular study of

L. pyromelana by Burbrink et al. (2011) not only concluded that they

were separate species, but restricted the range of L. pyromelana to

north oftheMogollonRiminArizona.Basing species and geographic

distributions on non-selective molecular characters does not reflect

the existing evolutionary hypotheses (subspecies) in any meaningful

way. Extending the range of L. knoblochi into central Arizona would

mean that most individuals in Sky Island populations half of its geographic distribution would not have the pattern characters typical of

this subspecies.

	 Disjunct populations of a number of SMO amphibians and reptiles

are known in Sky Island mountain ranges in Sonora (table 2). Hale

collected Salvadora bairdi in Arroyo el Nogalito on the west side

of the Sierra la Madera (UAZ 53743) in November 1981. This is a

remarkable range extension of ca. 320 km north-northwest of Milpillas in the SMO on the Sonora-Chihuahua border (AMNH 102194).

Salvadora grahamiae would have been expected instead.

	 A few species in Sierra la Madera are not known from the SMO

(table 2). Aspidoscelis opatae, which is endemic to the Río Bavispe

Valley (Enderson et al. 2010), occurs just south of Óputo. Craugastor augusti and Oxybelis aeneus are expected to occur in the Sierra

la Madera. Bufo woodhousii and Thamnophis marcianus have been

collected along theRíoBavispe just north ofÓputo, andmay be found

farthersouth. Aspidoscelis exsanguis A. tigris, Cophosaurus texanus, 

Crotalus atrox, and Heloderma suspectum are desertscrub/desert

grassland species reaching their western/southern limits in foothills

thornscrub in central Sonora. Plestiodon obsoletus is a grassland

species that is widespread in the Great Plains from Wyoming south

to Texas and west to southeastern Arizona. Cliff (1953) reported it in

Sonora from west of Huásabas. We observed one in the same area at

Mesa Quemada in FTS in the southern Sierra la Madera. Quijada-M.

et al. (2007) reported it from 34 km east of Hermosillo in foothills

thornscrub/Sonoran desertscrub. In July 1975, Allan E. Greer collected one from near el Coyote (MVZ 137664), only 19 km north of

Bacadéhuachi and just west of the Sierra Bacadéhuachi, almost in

the SMO.

	 A few notably tropical animals reach their northern limits in the

Sierra la Madera. Leptophis diplotropis is a colorful snake found in

tropicalwesternMexicofromOaxacanorth(Lemos-Espinal andSmith

2009). Most Sonoran records of this showy, aggressive snake are in

TDF in the Álamos area, but McCoy (1964) reported it from along

the Río Moctezuma south of Cumpas. Thisis a substantial record of a

tropicalspecies evidently using a river corridorto disperse northward.

The nearest locality to Cumpas is Santa Rosa in the Municipio de

Yécora, ca.170 km to the south-southeast.

	 Until recently, Sonora aemula was known in Sonora only as a TDF

species with most records from around Álamos, and a single locality
USDA Forest Service Proceedings RMRS-P-67. 2013 113

Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . . Van Devender, Enderson, Turner and others
114 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Enderson, Turner and others Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . .
USDA Forest Service Proceedings RMRS-P-67. 2013 115

Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . . Van Devender, Enderson, Turner and others

east-northeastofÓnavas(28°29’N;UAZ45675).JulioLemos-Espinal

found one atTónichi on the RíoYaqui (28°36’N; CONABIO record).

In 2012Van Devender photographed another one east of San Pedro de

la Cueva in the Sierra Agua Verde (29°16’N), extending the known

distribution ca. 100 km north-northwest of the Ónavas locality and

ca. 80 km north-northwest of Tónichi. Finally Roberto HernándezJimenez photographed S. aemula at Tonibabi Spring in the Sierra la

Madera (29°51’N; Hugo Silva- Kurumiya, pers. comm., 2012), 150

km north of the Ónavas locality (new records and images in MABA

database).

Biodiversity

Several factorsinfluence biodiversity.Within a Sky Island,species

richnessisinfluenced by local factorssuch as area, elevational range,

vegetation diversity, habitats, bedrock types, and land use history.

Geographic locationisimportant because,in general,speciesrichness

in many groups’ biodiversity increases southward to more tropical

habitats. Although the northernmost tropical deciduous forest is in

the Sierra San Javier (28°35’N; 100 km south-southwest of Sierra

La Madera, 120 km southwest of Sierra Bacadéhuachi), foothills

thornscrubanditstropical elementsoccurinawide transitionbetween

Sonoran desertscrub and oak woodlandsin the Sky Island ranges and

the mainland Sierra Madre Occidental.Also the nearnessto the SMO

isimportant because biodiversity in general is higherin the SMO than

any Sky Island (see Reina-G. and Van Devender, 2005 for floristic

comparison).

	 The documentation of biodiversity isstrongly influenced by access

(roads), nearness of research centers, and the inventory effort (number of trips, field time, etc.) and the number and skills of observers.

Herpetofaunas are difficult to study for a variety of reasons. Many

species only emerge from underground retreatsto feed and breed during a short part of the warm season and especially during the summer

rainy season. Nocturnal, secretive species are rarely encountered.

	 The herpetofauna of the Sky Island Sierra la Madera with 59 taxa

is moderately well known. Its affinities are strongly Madrean with

88.1% of the fauna shared with the SMO. Except for the Río Bavispe

Valley endemic Aspidoscelis opatae, the species not known from the

SMO are desertscrub/desert grassland species that are Sky Island

faunas that are widespread in the Sonoran Desert to the west and/or

desert grassland to the north. Old records of several additional taxa

fromtheCumpas-Moctezuma area,includingAspidoscelis uniparens, 

Phrynosoma cornutum, P. modestum, Sceloporus cowlesi, and Terrapene ornata, are desert grassland taxa not known from the SMO.

These would be significant southern range extensions, but are not

included in the present study until validated.

	 The Sierra Bacadéhuachi is in the SMO, but the known herpetofauna is small despite four visits by many observers. Moreover,

the numbers of individuals, especially of typically common, easilyobserved species (Hyla arenicolor, Sceloporus jarrovii, Thamnophis 

cyrtopsis, etc.)seen in our 2011-2012 fieldwork were low,suggesting

the possibility thatsome recent catastrophemight have impacted their

populations— perhaps the severe freeze of February 2-3, 2011. All

of the Bacadéhuachi taxa occur in the Yécora area.

	 The Yécora herpetofauna is much richer than the Sierra la Madera

fauna (table 2), and likely any Sky Island in Sonora. One species, the

snail-eatingsnakeTropidodipsas repleta,is endemic totheYécora area

(Smith et al. 2005).Additional species known from the SMO may be

added to the Yécora area. Plestiodon brevirostis (Rorabaugh, 2008)

and Salvadora bairdi (AMNH 102194) are known in the SMO from

near Milpillas on the Chihuahua-Sonora border, 120 km south of the

Yécora area. Several species, including Coluber taeniatus, Crotalus 

scutulatus, Spea multiplicata, Thamnophis elegans, and T. sirtalis 

occur in the Yépomera area in Chihuahua to the east (Van Devender

and Lowe 1977). Sceloporus albiventris is known from the Álamos

and as far north as Zetasora on the Northern Jaguar Reserve north of

Sahuaripa in TDF Rorabaugh et al. 2011), but not yet in the Yécora

area. Another two amphibians and 12 reptiles (10 snakes) occur in

TDF near Álamos, but have not yet been found near Yécora. We

estimate the potential SMO herpetofauna in eastern Sonora at 113

species of amphibians (24) and reptiles (89). Colubrid snakes are

especially diverse with 45 species, which contrasts with 187 species

for Sonora as a whole, including 35 amphibian and 152 reptile species

(Enderson et al. 2009).

	 With the exception of the Northern Jaguar Reserve north of

Sahuaripa in FTS (Rorabaugh et al. 2011), there are no published

herpetofaunas for eastern and northeastern Sonora. The two local

herpetofaunas presented here and comparisons with the mainland

SMO herpetofauna in the Yécora area help understand the regional

biodiversity, but additional studies are needed in individual Sky Island ranges and other areas in the SMO. The fauna of the Tres Ríos

area, the wettest highlands on the Sonora-Chihuahua border in the

northernmost SMO, would be especially interesting.

Acknowledgments

	 We thank Ana Lilia Reina-Guerrero for help in the field. Careful

and thoughtful reviews by R. Wayne Van Devender and Thomas R.

Jones greatly improved the paper. Gertrudis Yanes-Arvayo provided

the map.

References

Brennan, Thomas C.; and Holycross, Andrew T. 2006. A field guide to 

amphibians and reptiles in Arizona. Phoenix: Arizona Game and

Fish Department. 152 p.

Burbrink, F. T.; Yao, H.; Ingrasci, M.; Bryson, Jr., R. W.; Guiher, T. J.; and

Ruane, S. 2011. Speciation at the Mogollon Rim in the Arizona Mountain

Kingsnake (Lampropeltis pyromelana). Molecular Phylogenetics and

Evolution. 60: 445–454.

Cliff,F.S.1953.SelectedrecordsoflizardsfromSonora.Copeia.1953(3):186-187.

Enderson, Erik F.; Quijada-M., Adrián; Turner, Dale S.; Rosen, Philip C.;

and Bezy, Robert L. 2009. The herpetofauna of Sonora, Mexico, with

comparisons to adjoining states. Check List. 5: 632–672.

Enderson, Erik. F.; Quijada- M., Adrián; Turner, Dale S.; Bezy, Robert L.;

and Rosen, Philip C. 2010. Una synopsis de la herpetofauna con comentarios sobre las prioridades en investicagión y conservación. Pg. 357-383

in Molina-Freaner, Francisco and Van Devender, Thomas R. eds., Diversidad Biológica de Sonora, Universidad Nacional Autónoma de México,

Hermosillo, Sonora.

Ferguson,GeorgeM.;Turner,Dale;Hale,StephenF.;Villa,Robert;Hedgcock,

Chip;Enderson,ErikF.Inpress.Craugastor tarahumaraensis(Tarahumara

Barking Frog). Geographic Distribution. Herpetological Review

Hale, Stephen F.; and Jarchow, James L. 1988. The Status of the Tarahumara Frog (Rana tarahumarae) in the United States and Mexico; Part 2.

Schwalbe, Cecil R. and Johnson, Terry B. eds. Report to Arizona Game

and Fish Department, Phoenix, and Office of Endangered Species, U.S.

Fish and Wildlife Service, Albuquerque, New Mexico.

Hale, Stephen F.; Ferguson, GeorgeM.; Holm, PeterA.; andElizabethB.Wirt.

1998. Re-surveyofselectedTarahumara frog(Rana tarahumarae)localities

in northern Sonora,Mexico, inMay 1998. Reportsubmitted toTheArizona

Zoological Society and The Tarahumara Frog Conservation Team. 24 p. 

Lemos-Espinal, Julio A.; and Smith, Hobart M. 2007. Anfibios y Reprtiles

del Estado de Chihuahua. CONABIO, México. 613 p.
116 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender, Enderson, Turner and others Comparison of Preliminary Herpetofaunas of the Sierras la Madera . . .

Lemos-Espinal, Julio A.; Chisar, David; and Smith, Hobart M. 2003. Knobloch’s kingsnake (Lampropeltis knoblochi) of México a species. Bulletin

of the Maryland Herpetological Society. 39: 53-58.

Lowe, Charles H.; and Woodin, William H. 1954. A new racer (genus Masticophis) from Arizona and Sonora, Mexico. Proceedings of the Biological

Society of Washington, 67: 247-250.

Martin, Paul S.; Yetman, David A.; Fishbein, Mark; Jenkins, Philip; Van

Devender, Thomas R.; and Wilson, Rebecca K. (eds.). 1998. Gentry’s

Río Mayo Plants. The tropical deciduous forest and environs of Northwest

Mexico. University of Arizona Press, Tucson. 558 p.

McCoy, Clarence J., Jr. 1964. Notes on snakes from northern Mexico. The

Southwestern Naturalist, 9: 46-48.

Pauly, Gregory B.; Hillis, David M.; and Cannatella, David C. 2009. Taxonomic freedom and the role of official lists ofspecies names. Herpetologia,

65: 115-128.

Quijada-M., Adrián; Enderson, Erik F.; Parra-S., Iván, and Bezy, Robert L.

2007. Plestiodon obsoletus (Great Plains Skink). Sonora, México. Herpetolgical Review, 38: 353.

Reina-G., Ana L.; and Van Devender, Thomas R. 2005. Floristic comparison

of an Arizona ‘sky island’ and the Sierra Madre Occidental in eastern

Sonora: the Huachuca Mountains and the Yécora area. Pp. 154-157 in 

Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; and Edminster,

C.B.(compilers). connectingmountainislands anddesertseas:Biodiversity

and management of the Madrean Archipelago II. Proc. RMRS-P-36, U.S.

Department of Agriculture, Forest Service, Rocky Mountain Research

Station, Fort Collins, CO.

Rorabaugh,J.C.2008.AnintroductiontotheherpetofaunaofmainlandSonora,

México, with comments on conservation and management. Journal of the

Arizona-Nevada Academy of Science. 40: 20-65.

Rorabaugh,James C.; Gómez-Ramírez, MiguelA.; Gutiérrez-González, Carmina E.; Wallace,J. Eric; and Van Devender Thomas R. 2011.Amphibians

and reptiles of the Northern Jaguar Reserve and vicinity, Sonora, Mexico:

A preliminary evaluation. Sonoran Herpetologist. 24: 123-131.

Smith, Hobart M.; Lemos-Espinal,JulioA.; Hartman, D.; and Chiszar, David.

2005.AnewspeciesofTropidodipsas (Serpentes:Colubridae)fromSonora,

Mexico. Bulletin of the Maryland Herpetological Society. 41: 39-41.

Yanes-Arvayo, Gertrudis; Montañez-Armenta, María de la Paz; SilvaKurumiya, Hugo; and Gil-Montaño, Enrique. 2011. Catálogo de Fauna de

Sierra La Madera. Fracción V de la Reserva Ajos Bavispe. Universidad de

la Sierra, Hermosillo, Sonora.

Van Devender, Thomas R., and Charles H. Lowe. 1977. Amphibians and reptiles of Yepómera, Chihuahua, Mexico. Journal of Herpetology. 11: 41-50.

Villa, Robert; Van Devender, Thomas R.; and Bygott, David G. In press.

Geophis dugesi aquilonaris (Chihuahuan Earth Snake). Sonora, México.

Herpetological Review,

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 117

Rivers and Streams
118 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 119

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G., Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Introduction

This paper examines the significance and conservation status of 

riparian vegetation along ephemeral streams in Pima County, with 

special emphasis on deciduous riparian forests. Despite the importance of riparian areas in the arid and semi-arid 

regions of the Western United States, the availability of water to 

riparian vegetation cannot be spatially evaluated over broad regions, 

in part because of inconsistent and incomplete information. U.S. 

Geological Survey (USGS) topographic maps differentiate perennial 

from non-perennial streams. Neither topographic maps nor National 

Wetland Inventory maps differentiate intermittent flow reaches from 

ephemeral stream reaches, a crucial distinction for many aquatic and 

riparian species of the Western United States. While discrete discharges 

of groundwater to the surface are sometimes mapped as springs on 

USGS maps, shallow groundwater systems that never discharge to the 

surface escape notice. Complicating the matter, effluent discharges 

today support riparian areas along streams that would otherwise be 

dry except during storm events (Fonseca and Regan 2001). The 2.3 

million-acre area within Pima County that was studied for the Sonoran 

Desert Conservation Plan (SDCP) is one region where stream inventory allows for a comparison of the distribution of riparian vegetation 

relative to water resources, across a diverse terrain. 

Distribution of Riparian Vegetation in Relation to 

Streamflow in Pima County, Arizona

Julia E. Fonseca

Pima County Office of Conservation and Sustainability, Tucson, Arizona

Mike List

Pima County Information Technology, Tucson, Arizona 

Abstract—We compared the distribution of riparian forest and woodlands relative to water resource availability for a 2.3 million-acre region for the Sonoran Desert Conservation Plan (SDCP). Most of Pima County’s 

riparian vegetation occurs along stream reaches that classify as ephemeral. Ninety percent or more of the 

mesquite woodlands, riparian scrub, and riparian strand classifications of the Brown, Lowe and Pase system 

are associated with ephemeral stream reaches in Pima County. Over half of the Sonoran cottonwood-willow 

vegetation is associated with stream reaches that classified as ephemeral. Conservation planning has often 

focused on protecting perennial stream habitat. Our results support the importance of identifying and 

protecting ephemeral and intermittent streams, particularly those having shallow groundwater tables, in the 

Southwestern United States.

Resource Inventory and Analysis

The water resource inventory for the Conservation Plan was initially 

conducted by Pima Association of Governments (PAG 2000) for all 

of Pima County outside the Tohono O’odham Nation. Surface flows 

were classified according to the United States Geological Survey 

(USGS) (Langbein and Iseri 1960) definitions to differentiate perennial, intermittent, and ephemeral reaches of streams. Where long-term 

stream flow extent monitoring (also known as “wet-dry” mapping) 

was available, the extent of flow during the driest conditions was 

used to define the perennial reach. 

Surface Flow Definitions

Perennial: one that flows continuously.

Intermittent: one that flows only at certain times of the year when 

it receives water from springs or from some surface source such as 

melting snow in mountainous areas.

Ephemeral: one that flows only in direct response to precipitation, 

and whose stream is at all times above the water table.

The topology of the ephemeral stream centerlines was based primarily on Pima County staff’s delineation of stream centerlines from 

orthophotographs at a scale of 1:12,000. In remote areas of western 

Pima County the 1:24,000 USGS digital line graphs were used.

Shallow groundwater systems were identified based on evidence 

of a water table within 50 feet of the land surface. This definition 

was based on evidence that groundwater at this depth could sustain 

mesquite bosques (ADWR 1994). Many of the larger zones were 

mapped using a combination of aerial photographic and topographic 

interpretation, and review and mapping of groundwater level information from various state and local agencies. The inventory of shallow 

groundwater systems was, and still is, incomplete, consisting only 

of the larger and better documented sites (PAG 2000).
120 USDA Forest Service Proceedings RMRS-P-67. 2013

Fonseca and List Distribution of Riparian Vegetation in Relation to Streamflow in Pima County, Arizona

The inventory identified 66 perennial stream reaches and 120 

intermittent stream reaches on 57 different streams (PAG 2000). 

Pima County later augmented the inventory with additional spatial 

coordinates for streams, springs, and shallow groundwater systems. 

The data layers are maintained as part of Pima County’s Geographic 

Information System and made available to the public at http://gis.

pima.gov/maps/sdcp/. 

To support conservation planning, Pima County used the Brown, 

Lowe, and Pase digital classification system (Brown et al. 1979) to 

classify the distribution and extent of vegetation along the region’s 

watercourses. The vegetation study area consisted of the 2.3 million 

acres of land within Pima County and excluded Wilderness, Monument, National Park, Refuge or Military Reservation, and areas under 

tribal control. 

Over 100 U.S. Geological Survey orthophotoquads at a scale of 

1:24,000 were used to delineate polygons of increased vegetation size 

and density or geomorphological evidence of hydrological patterns 

(Harris 2001). Biologists then assigned a vegetation classification to 

each polygon based upon field experience and existing data. 

Vegetation polygons were classified to the “biome” level, with the 

exception of the Sonoran Riparian Deciduous Forest and Woodlands 

biome (BLP code 224.5), which was further subdivided to distinguish 

mesquite from cottonwood-willow forest (Harris 2000). In this paper, 

“riparian’ vegetation refers to 200-series BLP biomes or associations.

The distribution of riparian vegetation along the watercourses 

was analyzed by intersecting watercourse centerlines with riparian 

vegetation polygons using ArcGIS 10. The miles of stream length 

was summed for the three types of flow and for the five categories 

of riparian vegetation. 

Watercourse centerlines came from three files. “Wash’ centerlines 

were based on delineations by County staff circa 1990, regardless of 

water resource availability. ‘Istreams’ and ‘pstreams’ were independently delineated by PAG in 2000 to identify intermittent and perennial 

stream reaches, respectively. To deal with overlapping centerlines 

in our analysis, we buffered ‘istreams’ and ‘pstreams’ and used the 

buffer zones to erase any adjoining centerlines from the ‘wash’ file. 

In areas where the thalweg of the channel had moved substantially 

over time, or where the topologies diverged, the two centerlines were 

retained as independent flow paths. This occurred along a few miles 

of the effluent-dependent Santa Cruz River.

Results

Over 320,000 acres of streamside vegetation was defined in 13 

naturally occurring biomes. The vast majority (86%) of vegetation 

along streams in Pima County is classified as upland vegetation 

(table 1). The remaining 14% consisted of four distinct riparian 

vegetation communities, with the Sonoran Riparian Deciduous Forest 

and Woodland biome being the most common, at over 25,000 acres. 

About 88% of this biome was mesquite “bosque” or woodland association. The remainder, some 3100 acres, was cottonwood-willow 

association (defined herein as 224.53 or 224.523).

The distribution of riparian vegetation communities is biased towards 

eastern Pima County (fig. 1). Interior Southwest riparian deciduous 

forest (223.2) is found primarily above 3000 feet, whereas mesquite 

bosques (224.52) are most common below 3000 feet. The majority 

of Sonoran cottonwood-willow forest (224.53) and riparian scrub 

(234.7) is found between 3000 and 5000 feet. 

Most of Pima County’s riparian vegetation (200-series Brown, Lowe 

and Pase classification) occurs along stream reaches that classify as 

ephemeral (table 2; fig. 2). Ninety percent or more of the mesquite 

woodlands, riparian scrub, and riparian strand plant communities are 

associated with ephemeral stream reaches in Pima County.

The Sonoran cottonwood-willow association of the Sonoran 

deciduous riparian forest and woodland biome is the type of plant 

community that is most closely, in the literature, tied to perennial or 

intermittent flow (e.g. Lite and Stromberg 2005). In Pima County, this 

association is more closely affiliated with perennial and intermittent 

flow than the other four riparian associations (table 2; fig 2). Still, 

over half of the Sonoran cottonwood-willow vegetation in our study 

area is associated with ephemeral stream reaches. Three-fourths of 

the cottonwood-willow vegetation was along a non-perennial reach.

Of the approximately 45 miles of Sonoran cottonwood-willow 

association found along ephemeral reaches, shallow groundwater 

conditions are known to occur along over 8 miles (18 percent). About 

76 of the 410 miles of ephemeral streams having Sonoran mesquite 

woodlands are mapped as intersecting a shallow groundwater polygon 

(19%). 

Discussion and Conclusions

Over 870 miles of ephemeral stream in Pima County possess some 

type of riparian vegetation. The majority of each of the five riparian 

vegetation associations was found along ephemeral, not perennial 

streams. Our results point to the importance of evaluating and protecting ephemeral streams for the significant habitat values they may 

provide to a wide array of wildlife. The degree of protection offered 

to ephemeral streams under the Clean Water Act is under debate 

(Levick and others 2008). Our results bolster evidence presented by 

Levick and others regarding the ecological functions of ephemeral 

streams. 

Shallow groundwater conditions, including perched aquifers along 

streams, could explain the persistence of some riparian vegetation 

found along ephemeral streams. Past groundwater pumping has contributed to a loss of year-round flowing streams and riparian forests 

in Pima County, mainly at lower elevations (Scalero and Fonseca 

2000). Even after the water table no longer discharges to the surface, 

Table 1—Acres of vegetation in Brown, Lowe and Pase biome classification. 

Brown, Lowe and 

Pase vegetation 

(biome) in acres

Upland 

biomes 

(sum of all 

100-series)

Interior 

Southwest 

riparian 

deciduous 

woodland 

(223.2)

Sonoran 

riparian 

deciduous 

forest and 

woodland 

(224.5)

Sonoran 

deciduous 

riparian scrub 

(234.7)

Sonoran 

interior strand 

(254.7)

269,281 5,833 25,437 7,752 5,337
USDA Forest Service Proceedings RMRS-P-67. 2013 121

Distribution of Riparian Vegetation in Relation to Streamflow in Pima County, Arizona Fonseca and List

Figure 1—Location of riparian vegetation in eastern Pima County.
122 USDA Forest Service Proceedings RMRS-P-67. 2013

Fonseca and List Distribution of Riparian Vegetation in Relation to Streamflow in Pima County, Arizona

shallow groundwater tables or moisture in the unsaturated zone below 

streambeds may remain accessible to streamside plants. 

This study did not analyze vegetation condition across the spectrum 

of flow intermittency. Along the San Pedro River in adjoining Cochise 

County, dense, multi-aged cottonwood-willow forests appear to be 

restricted to perennial or intermittent rivers where depth to the water 

table is less than 4 meters (Lite and Stromberg 2005). 

When analyzed for the Sonoran Desert Conservation Plan, only 

63% of the Sonoran cottonwood-willow forest type was located within 

an existing reserve, a lower degree of protection that was available to 

many other, more common vegetation communities in Pima County 

(Fonseca and Regan 2001). About 75% of the higher elevation Interior 

Southwestern forests were protected, although much of that was within 

National Forests subject to mining and other competing interests.

Pima County and Pima County Regional Flood Control District 

have adopted a number of provisions to conserve riparian areas, 

including floodplain management regulations that pertain to riparian 

habitats found along the County’s predominantly ephemeral streams. 

Figure 2—Vegetation by flow type.

A 2004 general obligation bond for implementing the SDCP resulted 

in Pima County acquiring cottonwood-willow forests and mesquite 

bosques formerly subject to private land development, and managing state grazing leases that include additional riparian areas. Pima 

County holds and manages water rights associated with the land 

acquisitions. However, Arizona law provides little protection for 

groundwater-dependent ecosystems and no direct protection for the 

water needs of riparian vegetation (Losi 2012). These “sub-irrigated” 

ephemeral and intermittent streams are critical wildlife resources in 

arid and semi-arid areas like Pima County, Arizona. 

Some 25 river miles of riparian vegetation along the Santa Cruz 

River is supported in part by effluent flows, supporting over 300 acres 

of riparian forest and woodland as well as riparian (Fonseca and 

Regan 2001). Continued discharge of effluent to the river depends 

largely on the actions of City of Tucson, which owns the majority 

of the effluent, and the Secretary of the Interior, who holds in trust 

28,300 acre-feet of effluent for the benefit of the Tohono O’odham 

Nation. 

Table 2—Distribution of vegetation associations (Brown and others 1979) by stream flow type. 

Interior Sonoran Sonoran Total 

Southwest	 mesquite	 Sonoran	 deciduous	 Sonoran	 flow

Flow type riparian woodland cottonwood- riparian interior length

(reach miles) deciduous (bosque) willow scrub strand in

(%) (223.2) (224.52) (224.53) (234.7) (254.7) miles

Ephemeral 129 (73%) 410 (92%) 45 (56%) 163 (90%) 129 (96%) 877

Intermittent 36 (20%) 30 (7%) 17 (21%) 16 (9%) 4 (3%) 104

Perennial 13 (7%) 5 (1%) 19 (23%) 1 (1%) 1 (1%) 39

Total miles 

intersecting 

vegetation biome 

or association 178 446 81 181 134 1020
USDA Forest Service Proceedings RMRS-P-67. 2013 123

Distribution of Riparian Vegetation in Relation to Streamflow in Pima County, Arizona Fonseca and List

Acknowledgments

The authors thank reviewers Eve Halper, Dale S. Turner, and Dr. 

Juliet Stromberg for their suggestions on earlier drafts. Vegetation 

mapping was supported by a federal grant for the Sonoran Desert 

Conservation Plan. Pima Association of Governments supported 

the initial water resource inventory for the Conservation Plan. Many 

citizens, scientists, developers and ranchers individually contributed 

their local knowledge and expertise. 

References

Arizona Department of Water Resources (ADWR). 1994. Arizona Riparian 

Protection Program Legislative Report. A report to the Governor, President 

of the Senate, and Speaker of the House. Phoenix: Arizona Department of 

Water Resources.

Brown, D. E., C. H. Lowe, and C. P. Pase. 1979. A digitized classification 

system for the biotic communities of North America, with community 

(series) and association examples for the Southwest. J. Ariz. Nev. Acad. 

Sci. 14 (Suppl. 1): 1-16.

Fonseca, J., and J. Regan. 2001. Suitability Analysis and Representation Goals 

for Cottonwood-Willow Forest Habitat in Pima County, Arizona. [Online] 

Sonoran Desert Conservation Plan County Administrator’s Office. Available: www.pima.gov/cmo/sdcp/reports/d10/019SUI.PDF [March 26, 2012]

Harris, L. K., J. A. Wennerlund, and R. B. Duncan. 2000. Riparian vegetation 

mapping and classification, Sonoran Desert Conservation Plan, Final Report. 

Pima County Administrator’s Office, Contract #07-30-H-127196. Available: 

pima.gov/cmo/sdcp/reports/d7/012SDC.PDF [Accessed October 18, 2010].

Langbein, W. B. and K. T. Iseri. 1960. Manual of Hydrology: Part 1. General 

surface-water techniques. General Introduction and Hydrologic Definitions. 

Geological Survey Water-Supply Paper 1541-A. Methods and Practices 

of the Geological Survey. United States Government Printing Office, 

Washington, DC.

Levick, L., J. Fonseca, D. Goodrich, [and others]. 2008. The Ecological and 

hydrological significance of ephemeral and intermittent streams in the arid 

and semiarid American Southwest. EPA/600/R-08/134; ARS/233046. U.S. 

Environmental Protection Agency and USDA/ARS Southwest Watershed 

Research Center.

Lite, S.J. and J.C. Stromberg. 2005. Surface water and ground-water thresholds 

for maintaining Populus-Salix forest, Sand Pedro River, Arizona. Biological 

Conservation. 125: 153-167.

Losi, C. 2012. Keeping dry streams green: Can landowners in Arizona and 

California use property rights to maintain groundwater-dependent riparian 

habitat along non-perennial watercourses? West-Northwest Journal of 

Environmental Law and Policy. 18:(1): 121-153.

Pima Association of Governments (PAG). 2000. GIS coverage of perennial 

streams, intermittent streams, and areas of shallow groundwater. Final 

Project Report, prepared for the Sonoran Desert Conservation Plan.

Scalero, D. and J.E. Fonseca. 2000. Historical occurrence of native fish in 

Pima County. Sonoran Desert Conservation Plan, County Administrator’s 

Office, Tucson, AZ. 

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
124 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Observations on the Seasonal Distribution of Native Fish 

in a 10-Kilometer Reach of San Bernardino Creek, Sonora, 

Mexico

C.O. Minckley

Cuenca los Ojos, Flagstaff, Arizona

Abstract—San Bernardino Creek is a northern tributary of the Río Yaqui that originates in the United States 

and crosses the International Border just east of Douglas, Arizona/Agua Prieta, Sonora and immediately 

south of San Bernardino/Leslie Canyon National Wildlife Refuge. Six of eight Río Yaqui native fishes occur 

in this reach:four minnows, a sucker, and a poeciliid. Information is presented on the annual and seasonal 

distribution of these fishes during 2008–2011.

Introduction

San Bernardino Creek is a small northern tributary of the Río Yaqui 

drainage originating in the United States and crossing into Mexico 

about 16 km east of Agua Prieta and Sonora/Douglas, Arizona. It 

flows south to join Río Bavispe of Sonora and Río Papigochic of 

Chihuahua and becomes Río Yaqui, entering the Sea of Cortez near 

Cuidad Alvero Obrego n (Hendrickson and others 1980).

Historically, San Bernardino Creek was a low-elevation, warmwater stream with alternating pools and riffles, undercut banks, and 

mud-gravel substrates. At one time, it also had considerable cienega 

habitat. This was before the San Bernardino Land Grant (~1820), after 

which, overgrazing and aquifer exploitation caused loss of aquatic 

habitats. These invasive management practices continued into the 

1970s until the property was acquired by The Nature Conservancy 

who transferred the land to U.S. Fish & Wildlife Service (USFWS) 

creating San Bernardino National Wildlife Refuge (SBNWR) (Cobble 

1995). However, by then, the impact of cattle and water mining had 

dramatically affected the watershed, leaving scars that can be seen 

today in the incised cienega deposits along San Bernardino Creek 

(Hendrickson and others 1980; Hendrickson and Minckley 1984; 

Cobble 1995). Impact on fish populations in the United States was 

extirpation of Yaqui sucker and near extirpation of Yaqui chub (Minckley and Marsh 2009). The status of fishes in Mexico during that time 

is unknown. Establishment of SBNWR included development of a 

management plan that focused on water conservation, aquatic habitat 

development, and native fishes restoration. These activities were augmented in Mexico with purchase of Rancho San Bernardino by Joe 

and Valer Austin in 1999, which includes San Bernardino Creek and 

the drainage to the south. Soon after its purchase, cattle were removed, 

ponds containing introduced fishes were drained, and a program of 

gabion construction began in San Bernardino Creek and associated 

drainages of Hay Hollow and Silver Creek. These activities removed 

most introduced fishes and slowed erosion and raised groundwater 

levels, which resulted in the once barren channel of San Bernardino 

Creek developing dense stands of cattail and bulrush.

History of Fish Collections

	 The San Bernardino Creek drainage has eight species of native

fishesincluding five minnows and one species each ofsucker, catfish,

and topminnow.The minnows are Mexican stoneroller,Campostoma 

ornatum; Mexican longfin dace, Agosia sp.; Mexican roundtail chub,

Gila minacae;beautifulshiner,Cyprinella formosus;Yaqui chub,Gila 

purpurea; Yaqui catfish, Ictalurus pricei; Yaqui sucker, Catostomus 

bernardini; andYaquitopminnow,Poeciliopsis sonoriensis round out

the fauna. Type locality for Yaqui catfish, Yaqui topminnow, Yaqui

chub, Yaqui sucker and beautiful shiner is San Bernardino Creek at

or near the International border (Girard 1857, 1859; E.A. Mearns

cited in Snyder 1915).

	 Fish collections in Mexico from 1880 to 1980 are summarized

in Hendrickson and others (1980) and include information on field

collections by Simon in 1943, Frost and Hendrickson in1944, and

Miller andWinn in 1950.Fishestaken in these surveysincludedYaqui

chub, Yaqui catfish, beautiful shiner, Yaqui topminnow, and Yaqui

sucker. Records also included carp, Cyprinus carpio; black bullhead,

Ameiurus melas; and largemouth bass, Micropterus salmoides. Later,

Varela and others (1990) reported rediscovery of Yaqui chub in the

creek justsouth ofthe InternationalBorder and also later, downstream

some 4 km from that border (Varela and Campoy-Favela 1996).

	 Surveys were initiated in San Bernardino Creek from the International Border to ~10 km downstream in May of 2008. This paper

reports the distribution of native fish of San Bernardino Creek and

some aspects oftheir biology betweenMay 2008 andDecember 2011.

Materials and Methods

Four stations were established along San Bernardino Creek: Station one is four, 25-m reaches in Black Draw, which runs from the

International Border south, 3 km; station two is downstream from
USDA Forest Service Proceedings RMRS-P-67. 2013 125

Observations on the Seasonal Distribution of Native Fish in a 10-Kilometer Reach . . . C. Minckley

the confluence of Black Draw and Silver Creek; station three is 3 km

downstream from station two, and station four is 3 km downstream

from station three. Stations two through four were 100 m in length.

Stations were sampled with 12, fine-mesh, baited minnow traps and 6

1-m baited Promar hoop nets. All traps were set overnight and run in

early morning. Fish collected were identified, counted, and released.

Results

Seasonal Distributions

	 Seasonal distributions are summarized in table 1. At Station one,

the most common species was Yaqui topminnow, which was most

abundant in fall and decreased in numbers into winter. Yaqui chub

was the second most abundant fish and was distributed inversely to

topminnow with highest percentages in winter, decreasing into fall.

Longfin dace and Mexican stoneroller were present in small numbers

throughout the year.

	 Yaqui topminnow was the most abundant fish overall at station

two followed by longfin dace, which was most abundant in summer.

Yaqui chub was present in small numbers in fall.

At station three, Yaqui topminnow was the most abundant species

with peak numbers in fall. It was also abundant in winter after which

decreased into spring and summer. Longfin dace wasthe second most

abundant fish, with largest numbers present in summer and spring. It

exhibited smallest numbers in winter and fall. Mexican stoneroller

was not abundant but found here more than any other station with

largest numbers occurring in summer and smallest numbersin winter.

Yaqui topminnow was the most abundant fish at station four and was

present in large numbers in fall, followed by summer. The smallest

number of topminnow was present in winter and spring. Mexican

stoneroller was represented in reduced numbers in all seasons with

higher numbers being found in winter and spring (table 1).

Within Species Distribution

	 Yaqui topminnow was more abundant at stations two through four

with largest numbers taken at station two and smallest at station one.

Thisreflects habitat differences between station one, which has dense

stands of cattail and bulrush, and other stations that are more open

and more shallow. It might also reflect some level of predation on

topminnow by Yaqui chub, which are abundant.

	 Yaqui chub was most abundant at Station one in winter when

large numbers were taken. Numbers decreased into summer/fall.

These numbers may reflect spawning, which occurs throughout

the year, but appears most common in winter/fall (Minckley 2009,

2010). This would concentrate the fish and also increase number of

young-of-year available for capture. During summer, the number of

Yaqui chub drops and may reflect a reduction in movements to avoid

higher water temperatures and anoxic conditions (Minckley 2010).

Currently, Yaqui chub is reported to be restricted to the 3-km reach

of station one (Minckley and Marsh 2009; Cobble 1995). This study

has extended its range ~6 km and documented a large population in

Los Ojitos Spring.

	 Mexicanchub,Gila minacae,wassynonymizedwithroundtailchub,

G. robusta (Miller 1976) but later resurrected based on molecular

differences discovered by Dowling (1998) (Norris and others 2003).

As discussed by Minckley (1973), “roundtail” were not thought to

occur in San Bernardino Creek, but later he concludes that Mexican

chub probably did occur historically in upper San Bernardino Creek

based on its connection with Cajon Bonito (Minckley and Marsh,

2009). The two Mexican chubs taken in this study represent the first

documented collections of this species from San Bernardino Creek.

	 Numbers of Mexican longfin dace and Mexican stoneroller are

much reduced in station one. This is not surprising as both species

are adapted to riffle/pool habitat, which is nonexistent in station one.

	 The last collection of beautiful shiner from the study area was in

1950 (Miller and Winn 1951). It was extirpated from San Bernardino

Creek in the United States in 1970 (Minckley 1973; Minckley and

Marsh 2005). In 1990,stock taken from Río Moctezuma, Chihuahua,

and cultured at Dexter National Fish Hatchery & Technology Center,

were repatriated into ponds on the SBNWR (Minckley and Marsh

2005). Today, populations on the refuge are large (B. Radke pers.

comm.).

	 Yaqui sucker was extirpated from San Bernardino Creek in the

United States in 1968 (DeMarais and Minckley 1993; McNatt 1974;

Minckley 1973). There are no historical records of this fish from the

study area although it is abundant in Cajon Bonito and is found in

other areas of Mexico (Miller and others 2005; Minckley and Marsh

2009). Thirty-six Yaqui sucker from Cajon Bonito were introduced

into San Bernardino Creek in August of 2008 on Station one. Adult

individuals of this stocking have been taken but no reproduction has

been documented (Minckley 2009; 2010).

Summary

	 San Bernardino Creek was surveyed for native fishes from the

International Border downstream 10 kilometers during 2008–2011.

Six native fish species were collected includingYaqui chub, Mexican

chub, Mexican longfin dace, Mexican stoneroller, Sonoran topminnow, and Yaqui sucker. Yaqui chub was most abundant in station one

and occurred downstream to station three. It was not taken further

downstreambutdocumentedinLosOjitosSpring.TwoMexicanchub

were takenfromStationone,representing thefirsttime thisspecieshas

been documented from this drainage. Mexican stoneroller occurred

throughout the study area. It was rare in station one but common in

remaining stations. Mexican longfin dace was also rarely found in

station one but was present in large numbersin otherstations. Sonora
126 USDA Forest Service Proceedings RMRS-P-67. 2013

C. Minckley Observations on the Seasonal Distribution of Native Fish in a 10-Kilometer Reach . . .

topminnow was throughout the study area and was more abundant

fromstation two downstream. Depending on season and year, it could

be represented by thousands of individuals. Yaquisucker occurred in

station one. Fish distributions were related to spawning, habitat types,

and effects of summer water temperature and anoxic conditions.

Acknowledgment

	 Joe and Valer Austin are thanked for allowing access to their

property in Mexico and supporting this research.

References

Cobble, K.S. 1995. Yaqui fishes recovery plan. Albuquerque, NM: U.S. Fish

and Wildlife Service, Region 2. 48 p.

DeMarais,B.D., andW.L.Minckley. 1993.Genetics andmorphology ofYaqui

chub,Gila purpurea, anendangeredcyprinidfishsubjecttorecoveryefforts.

Biological Conservation. 66(1): 195-206.

Dowling, T.E. 1998. Genetic diversity of the Gila complex as determined by

analysis of mitochondrial DNA. Rept. U.S. Bureau of Reclamation, Salt

LakeCity,UT.AwardNo4-FC-40-15700,ArizonaStateUniversity,Tempe.

Girard, C. 1857. Researches upon the cyprinoid fishes inhabiting the fresh

water of the United States ofAmerica, west of the Mississippi Valley, from

specimensin themuseumoftheSmithsonian Institution.Proceedings ofthe

Academy of Natural Sciences of Philadelphia. 8(1856):165-213.

Girard, C. 1859. Ichthyology of the boundary. Washington, DC: U.S. and

Mexican Boundary Survey. 85 p.

Hendrickson, D.A. and W.L. Minckley. 1984. Cienegas: Vanishing climax

communities of the American Southwest. Desert Plants. 6(3): 131-175.

Hendrickson,D.A.,W.L.Minckley,R.R.Miller,D.J.Siebert,andP.H.Minckley.

1980. Fishes of the Río Yaqui basin, Mexico and United States. Journal of

the Arizona-Nevada Academy of Science. 15: 65-106.

McNatt, R.M. 1974. Re-evaluation of the native fishes of the Río Yaqui in the

United States. Proceedings of the 54th Annual Conference of the Western

Association of State Game and Fish Commission: 273-279.

Miller, R.R. 1976.An Evaluation of Seth E. Meek’s Contributionsto Mexican

Ichthyology. 1976. Fieldiana Zoology. 69(1): 1-31.

Miller,R.R. andH.E.Winn.1951.AdditionstotheknownfishfaunaofMexico:

three species and one subspecies from Sonora. Journal of the Washington

Academy of Science. 41: 83-84.

Miller, R.R., W.L. Minckley, and S.M. Norris. 2005. Freshwater fishes of

Mexico., Chicago, IL: University of Chicago Press. 490 p.

Minckley, C.O. 2009. Observations on the fishes of San Bernardino Creek

Sonora, Mexico, 2008-2009. Final Report. Unpublished completion report

on file with the author.

Minckley, C.O. 2010. Observations on the fishes of San Bernardino Creek,

Sonora Mexico in 2010. Final Rept. Unpublished completion report on

file with the author.

Minckley, W.L. 1973. Fishes of Arizona. Phoenix, AZ: Arizona Game and

Fish Department.

Minckley, W.L., and P.C. Marsh. 2009. Inland fishes of the greater Southwest:

Chronicle of a vanishing biota. Tucson: University of Arizona Press.

Norris, S.M., J.M. Fischer, and W.L. Minckley. 2003. Gila brevicauda (Teleostei: Cyprinidae), a new species of fish from the Sierra Madre Occidental

of Mexico. Ichthyological Exploration of Freshwaters. 14: 19-30.

Snyder,J.O. 1915.Notes on a collection offishesmade byDr.EdgarA.Mearns

from rivers tributary to the Gulf of California. Proceedings, U.S. National

Museum. 49. No 2125: 573-586.

U.S. Fish and Wildlife Service. 2011. Endangered and threatened wildlife

and plants; review of native species that are candidates for listing as

endangered or threatened; annual notice of findings on resubmitted petitions; annual description of progress on listing actions. Federal Register

76(207): 66370-66439.

Varela-Romero,A., and J.R.Campoy-Favela. 1996.Evaluation of populations

ofCyprinodon macularius, Poeciliopsis occidentalis, Ictalurus pricei, Gila 

ditaenia and Gila intermedia in shared watersheds between Arizona and

Sonora. Tech. Rep. Submitted to Nongame Branch, Arizona Game and

Fish Dept., Phoenix, AZ.

Varela-Romero,A.,C.Galindo-Duarte,E.Saucedo-Monarque,L.S.Anderson,

P. Warren, S. Stefferud, J. Stefferud, S. Rutman, T. Tibbits, and J. Malusa.

1990. Re-discovery of Gila intermedia and G. purpurea. Proceedings,

Desert Fishes Council. 22: 33-34.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 127

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Biodiversity and Conservation of the Ciénega de Saracachi 

area, Sonora, Mexico

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Martín A. Villa-Andrade, Martín Reyes-Juárez, and Gonzálo Luna-Salazar

Comisión de Ecología Sustentable del Estado de Sonora, Hermosillo, Sonora

Martín Padrés-Contreras, Fernando Padrés

Rancho Agua Fría, Cucurpe, Sonora

Paul S. Martin

University of Arizona, Tucson, Arizona

Abstract—The Ciénega de Saracachi area, including Arroyo Santo Domingo and Cañón Quemado, is in the 

Municipio de Cucurpe in north-central Sonora (30°21’33”N 110°35’29”W), ca. 105 km south of the Arizona 

border. The vegetation is cottonwood-willow riparian forest in the Ciénega and rocky stream canyons with 

desert grassland on the slopes above. These upper tributaries of the Río San Miguel are natural corridors on 

the west side of the Sierras San Antonio and Azul. The flora contains 419 species in 99 families and the fauna 

301 species of animals in 83 families. Invertebrate animals with only 87 taxa in 17 families are very poorly 

known. Vertebrate animals with 215 species in 67 families are dominated by birds (79.5% of the species). 

Two plants and 19 species of animals are Federally listed in Mexico in the NOM-059-SEMARNAT-2010. The 

La Brisca talussnail (Sonorella aguafriensis) is endemic to Arroyo Santo Domingo. In 2010, the Saracachi 

area was nominated to be a Sonoran Área Natural Protegida to preserve its natural values and to develop 

ecotourism land use options.v

Introduction

During spring break at the University of Arizona in 1976, Paul S. 

Martin lead a group of students from the Geosciences Department, 

to participate in the Easter festivities in Cucurpe and to visit Ranchos 

Agua Fría and la Brisca. Postdoctoral student Tom Van Devender was 

amazed by the lush cottonwood forest at the Ciénega de Saracachi with 

its permanent pools inhabited by Sonoran mudturtles (Kinosternon 

sonoriense) and Yaqui sliders (Trachemys yaquia).

The seeds from this trip are still growing. Van Devender developed 

a life long interest in the flora of the area, and began a study of amazing vertebrate fossils in a sediment pocket in Arroyo Santo Domingo. 

Ornithologists Stephen Russell (University of Arizona) and Gale 

Monson (U.S. Fish & Wildlife Service) made the first of six trips to 

the area to make bird observations for their remarkable, groundbreaking book, The Birds of Sonora (Russell and Monson 1998). Martin’s 

legacy continues today as the Comisión de Ecología y Desarrollo 

Sustentable del Estado de Sonora (CEDES), the Sonoran state agency 

of ecology and sustainable development, is in the process of having 

the area declared a state protected natural area. In April 2011, Van 

Devender led an international group of 40 biologists to the area on a 

Madrean Archipelago Biodiversity Assessment (MABA) Expedition 

to observe and catalog animals and plants in the area, 9 months after 

Martin’s death at age 82. Here we summarize the biodiversity of this 

natural area in northern Sonora. 

Methods and Results

Study Area

The Ciénega de Saracachi is in north-central Sonora (30°22’42”N 

110°33’48”W), 12.3 km (by air) east-northeast of Cucurpe, Municipio 

de Cucurpe, ca. 105 km south of the Arizona border. The Ciénega is at 

942 m elevation in the Río Saracachi (30°21’33”N 110°35’29”W), a 

northern tributary of the Río San Miguel, which joins the Río Sonora 

just north of Hermosillo. The study area includes Rancho Agua Fría 

and the Ciénega de Saracachi in the Río Saracachi, Rancho La Brisca 

in Arroyo Santo Domingo, and Cañón Quemado in Arroyo la Palmita. 

It covers an area of ca. 7.9 km2 at 940-1000 m elevation centered on 

30°22’42”N 110°33’48”W. These upper tributaries of the Río San 

Miguel are natural wildlife corridors on the west side of the Sierras 

San Antonio and Azul.
128 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Biodiversity and Conservation of the Ciénega de Saracachi area, Sonora, Mexico

Inventories

Plants—The earliest plant collections in the general area were by 

Thomas Sheridan and Gary P. Nabhan near Cucurpe in October 1975 

and March 1976. The first collections in the Ciénega de Saracachi 

area were by Paul S. Martin, Mary K. O’Rourke, and Van Devender 

in April and August 1976. Martin returned to the area in June and 

September 1978, June 1981 (with Raymond M. Turner), May and 

September 1983 (with Jodie L. Duek), and April 1984. Van Devender 

has collected and observed plants in the Saracachi area on 17 trips 

between April 1976 and April 2012, accompanied by (in chronological order) Stephen F. Hale, Charles H. Miksicek, Niall F. McCarten, 

Kevin B. Moodie, Richard S. White, Kenneth L. Cole, Margaret C. 

Kearns, Vera Markgraf, Robert S. Thompson, Mary Erickson, Piet 

van de Mark, Philip D. Jenkins, George M. Ferguson, A. Lilia ReinaGuerrero, Martín Villa-Andrade, Gonzálo Luna-Salazar, Martín ReyesJuárez, Mark Fishbein, Anurag Agrawal, Barbara G. Phillips, Laura 

Moser, John L. Anderson, Gertrudes Yanes-Arvayo, María de la Paz 

Montañez-Armenta, F. Isaias Ochoa-Gutiérrez, and Martín PadrésContreras. Ferguson and Hale also visited the area with Fishbein and 

Michelle McMahon in May 2000 and March 2001. 

The flora of the Saracachi area contains 419 taxa in 301 genera and 

99 families. This is based on at least 393 collections in the University 

of Arizona Herbarium (ARIZ), and 682 observations on an October 

2010 and April-May 2011 Madrean Archipelago Biodiversity Assessment (MABA) scouting trip and Expedition. Most of these records 

are available in the Southwest Environment Information Network 

(SEINet)/MABA database (Madrean.org); Some of the older collections in ARIZ are not yet in the database.

Invertebrates—The few invertebrate observations from the study 

area were made on the MABA trips to the area. Van Devender photographed a small number of grasshoppers and dragonflies in 2010. John 

Palting made 100 observations of 77 species in April 2011. Robert 

A. Johnson observed six species of ants. Hugo Silva-Kurumiya from 

the Universidad de la Sierra collected a few terrestrial snails in the 

Ciénega de Saracachi in April 2011. Amy and Wayne Van Devender 

at Appalachian State University in Boone, North Carolina, identified 

them as Gastrocopta cristata and Hawaiia minuscula. 

Fishes—Alejandro Varela at the Universidad de Sonora inventoried 

the fish in the Ciénega de Saracahi in 1991, finding four species in 

three families. In April 2011, he found the same four species plus two 

non-native species: green sunfish/mojarra (Lepomis cyanellus) and 

largemouth bass/lobina (Micropterus salmoides). 

Herps—Van Devender has made casual observations of amphibians 

and reptiles on 16 trips to the study area between April 1976 and May 

2011, accompanied by (in chronological order) James A. Hudnall, 

Robert W. Van Devender, Villa-Andrade, Luna-Salazar, Reyes-Juárez, 

Ferguson, Hale, Eric Wallace, and Trevor Hare. The herpetofauna of 

the study area presently includes six species of amphibians in three 

families and twelve reptiles in seven families.

Birds—The primary bird surveys in the area were by Stephen M. 

Russell and Gale Monson who made 871 observations on six trips 

between April 1981 and April 1993. Villa-Andrade, Luna-Salazar, and 

Reyes-Juárez made casual observations as part of CEDES fieldwork. 

Carl Tomoff made 177 observations on the MABA Expedition in 

April-May 2011.

Mamvmals—Only casual observations of mammals have been 

made on trips to the study area.

Discussion

In 2010, the book Diversidad Biológica de Sonora edited by 

Molina-Freaner and Van Devender (2010) summarized the present 

knowledge of the abundance and distribution of plants and animals 

in Sonora. The Ciénega de Saracachi area is an important wetland 

natural area in north-central Sonora.

Flora Composition

The flora of the Saracachi area contains 419 taxa in 301 genera 

and 99 families. 

The families with the most species are Asteraceae (61 taxa, 14.6%), 

Poaceae (48 taxa, 11.5%), Fabaceae (29 taxa, 6.9%), Solanaceae (14 

taxa, 3.3%), Euphorbiaceae (13 taxa, 3.1%), Pteridaceae (12 taxa, 

2.9%), Convolvulaceae (11 taxa, 2.6%), Lamiaceae, Malvaceae, 

and Scrophulariaceae (10 taxa, 2.4% each). The genera with the 

most species are Bouteloua (8 taxa), Brickellia and Ipomoea (7 taxa 

each), Euphorbia and Muhlenbergia (6 taxa each), and Cheilanthes, 

Solanum, and Verbena (4 taxa each). 

Only Arizona walnut/nogal (Juglans major) and Santa Cruz 

striped agave/amole (Agave parviflora) have Amenazada (Threatened) Federal protection status protected in Mexico under NOM059-SEMARNAT-2010 (Diario Oficial de la Federación, 2010), but 

the NOM list does not consider infraspecific taxa. Agave parviflora

ssp. parviflora and the Santa Cruz beehive cactus/choyita cabezona

(Coryphantha recurvata) in the study area are of special concern in 

Arizona. The subspecies A. p. ssp. flexiflora and A. p. ssp. densiflora

are endemic to eastern Sonora (Starr and Van Devender, 2011). 

Lumholtz false cloak fern (Argyrochosma lumholtzii), collected 

by Fishbein, McMahon, and Ferguson in Arroyo Santo Domingo, 

is a very rare endemic species only known from a few localities in 

Sonora. Buddleja corrugata ssp. moranii in the same area is a variety 

of a Baja California butterfly bush that is a rare Sonoran endemic.

Plants with northern affinities in the Saracachi flora include Arizona 

dewberry/zarzamora (Rubus arizonensis), goldenrod (Solidago velutina ssp. sparsiflora), stinging nettle/ortiga (Urtica dioica ssp. gracilis), 

rice cutgrass (Leersia oryzoides), and smallflower desert-chicory

(Pyrrhopappus pauciflorus). The last two species are only known 

in Sonora from the Ciénega de Saracachi. Arroyo Santo Domingo 

has an unusual association of bigtooth maple (Acer grandidentatum, 

northern temperate), saguaro (Carnegiea gigantea, Sonoran Desert), 

and rock fig (Ficus petiolaris, tropical).

A number of tropical plants that are common in tropical deciduous forest and foothills thornscrub in southern Sonora are near their 

northern range limits in Arroyo Santo Domingo. These include a 

spurge/golondrina (Euphorbia subreniformis), rock fig, ortiguilla

(Gronovia scandens), and gatuño (Mimosa moniliformis). Additional 

tropical species that also occur in Sonoran desertscrub in central 

Sonora include tree ocotillo/ocotillo macho (Fouquieria macdougalii), fragrant elephant tree/torote blanco (Bursera fagaroides var. 

elongata), papelío (Jatropha cordata), and organpipe cactus/pitahaya

(Stenocereus thurberi). Chihuahua oak/cusi peludo (Quercus chihuahuensis) is a Sierra Madre Occidental and tropical species that was 

collected along the Río Saracachi just north of Cucurpe, but has not 

been found in the study area.

Twenty-four species were non-natives (5.7% of the flora, 8 grasses). 

Natalgrass/zacate rosado (Melinis repens), buffelgrass/zacate búfel 

(Pennisetum ciliare), and Johnsongrass/zacate juanón (Sorghum halepense) are invasive in some areas in Sonora, but not in the Saracachi 
USDA Forest Service Proceedings RMRS-P-67. 2013 129

VBiodiversity and Conservation of the Ciénega de Saracachi area, Sonora, Mexico Van Devender and others 

area. Watercress/berro (Nasturtium officinale) is potentially invasive 

in the stream in Arroyo Santo Domingo. Chinaberry tree/piocha 

(Melia azedarach), peach/durazno (Prunus persica), and pomegranate/granada (Punica granatum) are occasional survivors near old 

homestead sites in Arroyo Santo Domingo.

Vegetation

The vegetation in the Ciénega and in the rocky, incised stream canyon 

in Arroyo Santo Domingo is primarily cottonwood-willow riparian 

forest. Dominant species include Fremont cottonwood (Populus 

fremontii), Goodding willow (Salix gooddingii), and Bonpland willow (S. bonplandiana). Netleaf hackberry/cúmaro (Celtis reticulata) 

and yewleaf willow/tarais (S. taxifolia) are locally common in the 

Ciénega. Huérigo (P. monticola) is near its northern distributional 

limit in Arroyo Santo Domingo. Arizona walnut is occasional in both 

areas. The floodplain of the Río Saracachi is dominated by velvet 

mesquite (Prosopis velutina). The slopes above the arroyos are in 

general desert grassland. One area in Arroyo Salsipuedes (an upper 

portion of Arroyo Santo Domingo) is foothills thornscrub. 

Fauna

The insects are the largest and least studied group of animals. 

Additional inventories are needed in all areas in various seasons to 

better understand this fauna. The Mexican blue-winged grasshopper 

(Machaerocera mexicana (Behrstock and Sullivan 2011), the arroyo 

grasshopper (Heliastus benjamini), and a tropical katydid (Neobarrettia sinaloae) found in Arroyo Santo Domingo in October 2010 were 

new species for Sonora.

Terrestrial snails are reasonably diverse, but secretive and little 

studied in Sonora. The endemic La Brisca talussnail (Sonorella 

aguafriensis) is only known from the original November 1984 collection by Russell Duncan in Arroyo Santo Domingo (Naranjo-García 

and Miller 1986). Snail inventories would be excellent additions to 

the biodiversity of the area.

The fish fauna with four species is well known in the study area. 

The native fishes in the entire Río Sonora basin include desert chub/

charalito sonorense (Gila eremica), Gila topminnow/guatapote de 

Sonora (Poeciliopsis occidentalis), topminnow/charalito (Poeciliopsis 

sp. nov.), longfin dace/charalito aleta larga (Agosia chrysogaster), 

Mexican stoneroller/rodapiedras mexicano (Campostoma ornatum), 

and Opata sucker/matalote Ópata (Catostomus wigginsi; Alejandro 

Varela-Romero, pers. comm. 2009). The Opata sucker is endemic to 

Sonora. The Opata sucker and Gila topminnow have the Mexican 

Federal status Amenazada in the NOM-059-SEMARNAT-2010(Diario 

Oficial de la Federación 2010). Exotic green sunfish and largemouth 

bass should be monitored, and control programs are needed to protect 

native fishes. 

Herpetofaunas are difficult to study for a variety of reasons. Many 

species only emerge from underground retreats to feed and breed 

during a short part of the warm season. Nocturnal, secretive species 

are rarely encountered. Six species of amphibians are known from 

the study area, but additional species are expected during the summer 

rainy season. At times, the lowland leopard frogs (Rana yavapaiensis) 

can be abundant. This frog is of special concern to state and federal 

agencies in the United States and Mexico because a chytrid fungus 

has caused population declines in many areas. The Arizona 

toad (Bufo microscaphus), which occurs primarily in Arizona, 

was found in the Ciénega in September 1990. Reptiles, with 13 

species, are better known in the study area. Additional taxa, especially 

snakes, will be encountered with more fieldwork. The northernmost 

population of the Yaqui slider (Trachemys yaquia) is in the Ciénega, 

and is considered threatened. Boa constrictor/corua (Boa constrictor), 

which was found in the Palm Canyon southeast of Magdalena (24 

km northwest of the Ciénega) in September 1976, is expected to be 

in the study area. Six species of amphibians and reptiles (Callisaurus 

draconoides, Coluber flagellum, Crotalus molossus, Elgaria kingii, Rana 

yavapaiensis, and Thamnophis cyrtopsis) have the Mexican Federal 

status Amenazada or Protección Especial (Special Protection) in the 

NOM-059-SEMARNAT-2010(Diario Oficial de la Federación 2010). 

The birds, with 171 species in 40 families, are the best-studied group 

of animals in the study area (Russell and Monson 1998). Additional 

observations in spring and late summer would add information about 

Neotropical migratory species. Birds of special interest observed in 

the study area include Osprey (Pandion haliaetus) and Gray Hawk 

(Asturina nitida). The Yellow-billed Cuckoo (Coccyzus americanus) 

has been observed elsewhere in the Municipio de Cucurpe. Nine species 

are Federally protected in Mexico in the NOM-059-SEMARNAT-2010

(Table 1; Diario Oficial de la Federación 2010). 

Mammals have been casually observed on various trips to the study 

area. The mammal fauna is presently at 19 species in12 families. Camera 

traps would provide better documentation of what medium and large 

species occur in the area. Smaller species, especially bats and rodents, 

need to be inventoried to increase the biodiversity knowledge. Arizona 

gray squirrel (Sciurus arizonensis) in Arroyo Santo Domingo 

is a southern population of this regionally endemic species. 

Scat of American black bear/oso negro (Ursus americanus) 

was observed by Trevor Hare in Arroyo Santo Domingo in April 

2011. Both species are Federally protected in Mexico in the NOM059-SEMARNAT-2010 (Table 1; Diario Oficial de la Federación 2010). 

This arroyo is a potential corridor between the surrounding 

mountain ranges for dispersing animals, such as jaguar/tigre 

(Panthera onca) and ocelot (Leopardus pardalis).

Fossil Record 

Van Devender et al. (1985) reported 51 taxa of vertebrates from 

the late Pleistocene Rancho La Brea Local Fauna in Arroyo Santo 

Domingo. The presence of Bison and Mammuthus—recent immigrants 

into North America from Siberia— indicated a late Pleistocene age 

for the deposit. The abundance of Sonoran mudturtles and leopard 

frogs (Rana “pipiens”, includes R. yavapaiensis) and the presence 

of a slider (Pseudemys scripa, now Trachemys yaquia), Red-winged 

Blackbird (Agelaius phoeniceus), and fishes (Agosia, Catostomus, 

and Poeciliopsis) indicated a paleoenvironment similar to the modern 

Ciénega de Saracachi before Arroyo Santo Domingo was down cut. 

The extinct La Brea Owl (Strix brea) was in the fauna. Additional 

large mammals in the fauna were a burro-sized horse (Eques cf. tau), 

a large horse (Eques sp.), a camel (Camelops sp.), four-horned antelope (Tetrameryx sp.), and mule deer (Odocoileus hemionus). Today 

white-tailed deer (O. virginianus) is common in the area. A number 

of species indicate warmer and/or more tropical environments than 

today, probably in the last interglacial period. These included Arizona 

mudturtle (K. arizonense), barking frog (Craugastor augusti), Kellogg’s toad (Bufo cf. kelloggii), Neotropical whipsnake (Coluber cf. 

mentovarius), and sabinal frog (Leptodacylus melanonotus).

Conclusions

The species richness in the riparian wetland habitats in the Ciénega 

de Saracachi, Arroyo Santo Domingo and Cañón Quemado is very 
130 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Biodiversity and Conservation of the Ciénega de Saracachi area, Sonora, Mexico

diverse with 419 plant taxa and 215 vertebrate taxa. The invertebrate 

fauna has 87 taxa, but is only a sparse beginning of the true diversity. 

The endemic La Brisca talussnail and several other species are the 

only known Sonoran from the area.

A total of 21 species (2 plants, 19 animals) are Federally protected in 

Mexico under NOM-059-SEMARNAT-2010 (Table 1; Diario Oficial 

de la Federación 2010). Thirteen species have the status Amenazada, 

three Peligro (= Endangered), and three Protección Especial. The 

three Peligro species are American black bear (Ursus americanus), 

Bell’s Vireo (Vireo bellii), and Elf Owl (Micrathene whitneyi).

In 2010, CEDES submitted a proposal to the Governor of Sonora 

to declare The Ciénega Saracachi area a state protected natural area to 

conserve and restore structure and function of these important wetland 

habitats and to preserve the rich diversity of plants and animals.

Acknowledgments

We thank Ana Lilia Reina-Guerrero, Gertrudis Yanes-Arvayo, María 

de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya for help in 

the field. We thank Sky Island Alliance for supporting conservation 

in this unique area. We thank the Palafox family of Rancho la Brisca 

for more than 30 years of hospitality. Sky Jacobs’ careful review 

improved the manuscript. This paper is dedicated to the memory of 

Paul S. Martin (1928-2010), mentor, friend, scientist, natural historian, 

and explorer. 

References

Behrstock, Robert A.; and Sullivan, Patrick H. 2011. First records of the 

grasshopper Machaerocera mexicana Saussure, 1859 (Orthoptera: Acrididae) from the United States and Sonora, Mexico. Insecti Mundi. 199: 1-4.

Comisión de Ecología y Desarrollo Sustentable del Estado de Sonora. 2010. 

Etudio previo justificativo para proponer a la Ciénega de Saracachi, Municipio de Cucurpe, Sonora, como un Área Natural Protegida Estatal bajo 

la categoría de Reserva. Hermosillo, SON.

Diario Oficial de la Federación. 2010. Norma oficial Mexicana, NOM059-SEMARNAT-2010, Protección Ambiental-Especies nativas de México 

de flora y fauna silvestres-Categorías de riesgo y especificaciones para su 

inclusión, exclusión o cambio-Lista de especies en riesgo. Emitido por 

la Secretaría del Medio Ambiente y Recursos Naturales. 06 de marzo del 

2002. México D.F., p. 80.

Molina-Freaner, Francisco; and Van Devender, Thomas R., eds. 2010. 

Diversidad Biológica de Sonora. Universidad Nacional Autónoma de 

México, Hermosillo.

Naranjo-García, Edna; and Miller, Walter B. 1986. A new species of Sonorella 

(Gastropoda: Pulmonata: Helminthoglyptidae) from Sonora, Mexico. The 

Velager. 29: 166-168.

Russell, Stephen M.; and Monson, Gale. 1998. The Birds of Sonora. Arizona: 

Tucson. 360 p.

Starr, Greg; and Van Devender, Thomas R. 2011. Agave parviflora subspecies 

densiflora. A newly found treasure from the Sierra Madre in eastern Sonora. 

Cactus and Succulent Society Journal. 83: 224-231.

Van Devender, Thomas R.; Rea, Amadeo M.; and Smith, Michael L. 1985.

The Sangamon interglacial vertebrate fauna from Rancho la Brisca, Sonora 

(with A. M. Rea and M. L. Smith). Transactions of the San Diego Society 

of Natural History. 21: 23-55.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 131

Biodiversity
132 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 133

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Preliminary Assessment of Biogeographic Affinities of 

Selected Insect Taxa of the State of Sonora, Mexico

Robert W. Jones, Alejandro Obregón-Zuñiga, and Sandra Guzman-Rodriguez

Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Querétaro, Qro., México

Abstract—The biogeographic affinites of butterflies (Lepidoptera: Papilionoidea and Hesperidae), damsel and 

dragonflies (Odonata), and ants (Hymenoptera: Formicidae) reported from the State of Sonora, Mexico were 

analyzed using published species lists. The combined distribution of these taxa was proportionally greater 

(47.4%) for those species within the Mega-Mexico3 biogeographic category (Southwestern United States 

south to northern Central America). Formicidae was the most highly restricted taxon with greater proportions of Sonoran desert endemics. Butterflies had a greater proportion of tropical species (82.8%), whereas 

dragonflies and damselflies from Sonora were most widely distributed either north or south of Mega-Mexico. 

Differences in the biogeographic affinities of the three insect taxa are attributed to specificity of immature 

host plants and the distribution and persistence of species habitats. 

Introduction

	 Sonora is a state with high biodiversity (Molina-Freaner and

Van Devender 2010). The state is the second largest in Mexico and

traditionally considered a transition zone between the Nearctic and

Neotropical biogeographic realms. The state contains portions of

three biogeographic provinces within its boundaries (Morrone and

others 2002) including the major portion of the Sonoran province,

the most northern extension ofthe Sierra Madre Occidental Province,

and the extreme northwest portion of the Mexican Plateau. Marshall

and Liebherr (2000) report that the region containsimportant areas of

endemismandthese aredistinctfromendemic speciesfrommountains

(Sky Islands) of southeastern Arizona.

	 Althoughthe informationconcerningthe insectsofSonorahasbeen

enriched in recent years (Bailowitz and Palting 2010), the number

of insects reported from the state is still limited, given its area and

biological diversity with overrepresentation of several orders. Of

the records in the Madrean Archipelago Biodiversity Assessment

database, and from records of the Mexican National Commission

for the Knowledge and Use of Biodiversity (CONABIO), there are

reported a total of 10,341 insect specimen records. Of these records,

65% are Lepidoptera, 12% are Odonata and 9% are Hymenoptera.

Thisis a typical collection bias(butterflies and dragonflies) for many

relatively poorly collected areas. However, research efforts, some of

which have been promoted through CONABIO and other agencies

in Mexico, have led to the publication of many national, state, and

regional flora and faunistic studies such as Llorente-Bousquets and

others (1996), Rojas (2001), García-Mendoza and others (2004),

Garwood andLehman (2005),Upson and others(2007) andBailowitz

and Palting (2010), which now allows for comparisons of species

compositions of selected taxa between regions in Mexico.

	 In the present study, we analyzed previously compiled data from

various sources of the species reported from the Mexican state of

Sonora for three of the better known taxa: butterflies (Lepidoptera:

Papilionoidea and Hesperidae), damsel and dragonflies (Odonata),

and ants (Hyemnoptera: Formicidae). Our general hypothesis was

that the geographic affinities of the majority of species from these

three taxonomic groups reported from Sonora fit into a distribution

within the boundaries of “Mega-Mexico 3” (herein,Mega-Mexico) of

Rzedowski (1998), a term equivalent to “Mexican Transition Zone”

ofHalffter(1987).We further delineated biogeographic categoriesfor

species with distributions within and/or beyond Mega-Mexico, and

discuss species with unique distributions. A comparison of species

compositions of selected butterfly groups between other Mexican

states and Arizona is also presented.

Materials and Methods

	 Specieslistsofbutterflieswere recordedfromBailowitz andPalting

(2010), as well as Scott (1986), DeVries (1987), Llorente-Bousquets

and others (1996), Rivera Granados (1998), Garcia Jimenez and

Guevara Guerro (2005), Garwood and Lehman (2005), Warren and

others (2012) and Opler and others (2012). Species lists for Odonata

included Westfall and May (1996), Ramirez and others (2000), Upson

and others (2007), Bick and Mauffray (2011), and Paulson and

Gonzalez-Soriano (2012). Lists of ant species were compiled from

the list of the Ants of Sonora, Mexico, compiled by the ASU Social

Insect Search Group (2012), and Rojas (2001),

	 The geographic distributions ofspecies of butterflies, Odonata and

antsweredividedintofourgeneralbiogeographiccategories:(1)MegaMexico, (2) Tropical, (3) Temperate, and (4)American. The category

Mega-Mexico corresponded to “Mega-Mexico 3” of Rzedowski,

(1993), a distribution that includes the extreme southwestern United

States, Mexico, and northern Central America (to northern Nicaragua).

Tropical species were those that occur within Mega-Mexico, some
134 USDA Forest Service Proceedings RMRS-P-67. 2013

Jones, Obregón-Zuñiga and Guzman-Rodriguez Preliminary Assessment of Biogeographic Affinities of Selected Insect Taxa . . .

of which extended asfar north asthe southwestern United States, but

are recorded south beyond northern Central America into southern

CentralAmerica and/or into SouthAmerica. The species categorized

as temperate species had distributions extending north of the southwestern United States (north of the limits of Mega-Mexico 3), with

distributions extending south into Mexico and, rarely, into northern

Central America. American species were those with ranges from

the central to the northern United States, and/or Canada, into south

Central America and/or South America and/or the Caribbean.

	 Within each ofthe four categories,further categories were grouped

based on previously reported discontinuities reported by Halffter

(1987): the Transverse Volcanic Belt, and the Isthmus of Tehuantepec. Thus, species from Mega-Mexico were further subdivided into

(1)speciesfound throughout Mexico, and also south ofthe Isthmus of

Tehuantepec; (2)species with rangesin north and central Mexico but

found only north of the Isthmus of Tehuantepec; (3) Northern species

found north of the Trans volcanic axis; and (4) Sonoran endemics.

Temperate subcategories were based the separation of eastern and

western northAmerica, an important pattern noted by Noonan (1988)

and included (1) species from Sonora that extended to the western

Rocky Mountain States north of Mega-Mexico, sometimes into

Canada, but not into the central or eastern United States; (2) species

reported from Sonora but did not extend into the Rocky Mountains

and found in the southeast and/or central eastern United States; and (3)

species from Sonora found in both the eastern and western United

States, sometimes also into Canada. Subcategories of tropical species from Sonora were (1) species whose range extended south into

southern Central America (south of Nicaragua); (2) those reported

into southern Central America and the Caribbean; and (3) species

extending into South America and the Caribbean.

	 Comparison of proportions between biogeographic categories

among taxa were made using a χ2 pairwise analysis and species

composition between regions with a cluster analysis using Jaccard’s

index.The cluster analysiswas conductedusingrepresentative regions

(states) within Mexico (Veracruz, Tamaulipas, Chiapas, Querétaro,

and Sonora) and the State of Arizona. The butterfly families Pieridae

and Papilionidae were used as these taxa had the most complete and

comparable data. Both analyses were performed with the PAST®

statistical software program (Hammer and others 2001).

Results

	 The combined distribution of butterflies, dragonflies, damselflies

andantswereproportionallygreaterforthose specieswithintheMegaMexico 3 biogeographic category (47.4%), followed by species with

Tropical distributions (28.2%), Temperate (13.5%), and American

(10.8%). Ofthe species within Mega-Mexico, the greatest proportion

of species (27.3%) had distributions southward from Sonora, but not

extendingbeyondthe IsthmusofTehuantepecwith13.9%foundfrom

Sonora into south Mexico or beyond into northern Central America.

All other subcategories (species extending beyond MegaMexico to

the north,south, or both) had 11% orless ofthe speciestotals. Species

endemic to the Sonoran desert accounted for 6.3%, of which 76.9%

were species of ants.

	 The proportions ofspecies within these four general biogeographic

categories were not similar between the three taxa compared ( fig. 1)

and all pairwise comparisons were significantly different (χ2 > 40, df = 3,

p < 0.01). Formicidae was the most highly restricted taxon with 82

species (78.1%) found only within Mega-Mexico, and of these, 30

species 28.6% were classified as Sonoran desert endemics. Of the

speciesfrom Mega-Mexico, 50 species(47.7%) were found from the

Sonoran desert and into parts of central Mexico and only two species

(1.9%) extended beyond the Isthmus of Tehuantepec. Only 12 ant

species (11.4% ) found in Sonora had distributions extending northwards beyond the southwestern states ( fig. 1), nine (8.6%) extended

into Central America and beyond, and only two were found in both

temperate and tropical latitudes.

	 Butterflies also had notable proportions of species from MegaMexico ( fig. 1), but had a greater proportion than the other taxa of

tropical, wide ranging species (82.8 % either from Mega-Mexico or

further south). These extended primarily from the northern limits of

Mega-Mexico into southern Central America (32 species; 10.4%) or

beyond into South America and/or the Antilles (85 species; 27.6%).

Sonoran butterflies that had both temperate and tropical latitudes

accounted for less than 5 percent of the species.

Figure 1—Biogeographic affinities of species of butterflies (Lepidoptera), 

dragonflies and damselflies (Odonata) and ants (Hymenoptera: Formicidae) reported from the State of Sonora, Mexico. A. Species endemic 

to the Sonoran Desert region (United States and Mexico); B. Species 

of reported within the southwestern United States and/or north and 

central Mexico but found only north of the isthmus of Tehuantepec; C. 

Species of reported within the limits of Mega-Mexico including regions 

north isthmus of Tehuantepec, and also from southern Mexico and/or 

northern Central America; D. Species restricted to Mega-Mexico to the 

south and also found in the western United States; E. Species restricted 

to Mega-Mexico to the south and also found in the eastern and western 

United States; F. Species restricted to Mega-Mexico to the north and also 

found in the southern Central America; G. Species restricted to MegaMexico to the north with ranges extending into South America, and/or 

the Caribbean; H. Species with ranges extending both north and south 

of the limits of Mega-Mexico. 
USDA Forest Service Proceedings RMRS-P-67. 2013 135

Preliminary Assessment of Biogeographic Affinities of Selected Insect Taxa . . . Jones, Obregón-Zuñiga and Guzman-Rodriguez

	 Species of dragonflies and damselflies from Sonora were the most

widely distributed ofthe three taxa.Almost half(47.9%) ofthe species

found in Sonora were also reported from southern Central America,

the Antilles, and/or the South America, or had even greater distribution range from the eastern or western United Statessouth beyond the

southern limits of Mega-Mexico. The Odonata also had the greatest

number of species from Sonora that were also found in the central

and/or eastern United States (84; 35.7%).

	 Comparisons ofspecies compositions of Pieridae and Papilionidae

butterflies reported from Sonora with other regions indicated an expecteddifferentiationbetweensouthernandnorthernMexicanregions

with important differences between the adjacent States of Sonora

and Arizona ( fig. 2). For Papilionidae, there was greater similarity

between Sonora and the northeastern State of Tamaulipas, over 1000

km to the east, than to adjacent Arizona. The species composition of

Pieridae indicated that Sonora wasthe most different from all regions

with less than 40% shared species with Arizona and Tamaulipas,

with the southern states forming a cluster similar to that found with

Papilionidae. The central State of Querétaro did not align with the

northern regions, but with Veracruz and Chiapas; this despite its

connection with Tamaulipas through the Sierra Madre Oriental and

having regions ofChihuahuan desert “relicts” (Morafka 1978),which

share arid adapted plant species with northeastern Sonora (Morrone

and others 2002).

Discussion

	 Biogeographicdistributionsaretheproductofecologicalconstraints

and past geological events (Endler 1982; Casazza and others 2008;

Kalkman and others 2008). However, separating these factors and

specific methods, such as phylogenetic reconstructions, is difficult

(Endler 1982), and notwithin the scope ofthe presentstudy.However,

a comparative approachmaygivepreliminary insightsintoexplaining

the differences and similarities between the distributions of the three

taxa reported within the political boundaries of Sonora.

	 The key ecological factors determining most insect distributions

are those related to trophic relationships and microhabitat. In the case

of butterflies, trophic associations and distributions are clearly linked

becausebutterflies arewhere theirhostplants are, atleastduringlarval

stages. In the present data set used, all of the 15 most widespread

species (American distribution) are polyphagous and reported from

a wide range of plant species. For example, Achlyodes tamenund, is

found on the many species of the Rutaceae family including Citrus 

spp., Erynnis funeralis is found on 13 genera of Leguminosae, and

Panoquina ocola feeds on several genera of Poaceae, including rice

and sugarcane. The only exception to the polyphagous habit of widespread species is Copaeodes minima, which is restricted to feeding

aslarvae on Cynodon dactylon, the widely cultivated Bermuda grass. 

Ofthe two Sonoran endemic speciesrecorded here, their distribution,

as well, is determined by the range of their larval host plant. Both

are species of Hesperiidae and are specific to species of Agavaceae

of the Sonoran desert: Agathymus aryxna on Agave palmeri and A. 

chrysantha on Agave schotti (Scott1986). Although the presence of

adequate hosts plants is clearly not the only determinate of butterfly

distributions(Dingle andothers2000), especiallyforthehighlymobile

adult stage, the effect of historical factors and ecological limitations

on their host plants is the primary predictor of butterfly distribution.

	 Comparison of species composition of Papilionidae and Pieridae

(fig. 2) among selected states of Mexico andArizona had an expected

separation between southern Mexican States of Oaxaca,Chiapas, and

Veracruz with the northern states ofTamaulipas, Sonora andArizona.

The central State of Queretaro, although compiled from collections

in the arid regions of the state, has more affinities to the south than

north.The greatersimilarity of Papilionidae betweenTamaulipas and

Sonora, than Sonora with adjacentArizona, was notable and probably

reflectsthe increasing dominance oftropical plants as one goesfurther

south in Sonora, asisfound in southern Tamaulipas. The relative dissimilarity between the butterfly fauna of Sonora and Arizona ( fig. 2)

is also reported by Marshall and Liebherr (2000) between temperate

and tropical adapted species of other faunal groups and plants. In the

case ofbutterflies,thisdissimilarity isprobably becausemany tropical

butterflies and their host plants (Felger and others 2001) reach their

most northern distribution in the State of Sonora, particularly in the

Sierra Madre Occidental, and these butterfliesrarely reach the United

States' border.

	 The distribution of species of Odonata are determined by the

distribution of suitable aquatic habitats (Kalkman and others 2008),

althoughlikebutterflies,theycanbe foundfarfromimmaturehabitats.

Kalkmanandothers(2008)reportthattheSouthwesternUnitedStates

is a center of diversity and endemism of dragonflies but many species

are wide-ranging, a pattern also found for species from northwestern

Figure 2—Cluster analysis of species richness of Papilionidae and Pieridae (Lepidoptera) from various states of Mexico using Jaccard’s Index.
136 USDA Forest Service Proceedings RMRS-P-67. 2013

Jones, Obregón-Zuñiga and Guzman-Rodriguez Preliminary Assessment of Biogeographic Affinities of Selected Insect Taxa . . .

Mexico. Because of the highly developed flight capabilities of adults

and the fact that both adults and nymphs are predaceous and feed on

a wide variety of prey, it is easy to understand how many species

can maintain widespread distributions. However,species with highly

restricted distributions are more difficult to explain, partially because

there is much to learn about microhabitat requirements of Odonata,

and accurate population estimates are usually lacking (Abbott 2005).

Also, lacking is an understanding of specific adaptations of Odonata

to unpredictable, marked seasonality of aquatic habitats in tropical

regions such as Sonora.

	 Ant species had the most restricted distributions of the taxa examined, with greater numbers of species restricted to the Sonoran

Desert. The Sonora Desert has a unique climatic regime (Ives 1949;

Ehleringer 1985), which apparently has persisted as a desert refuge

throughout the Quartenary (Hewitt 2000). As foraging success and

colonyfunctionof ants are tightlylimitedbytemperature andhumidity

(Hölldobler andWilson.1990),the climatic conditions andpersistence

of the Sonoran desert ecosystem through time have apparently been

conducive to the speciation and survival of ant species. Close to half

of the ant species of Sonora had distributions throughout northern

Mexico, particularly in stateswithin theChihuahuan desert,including

Arizona and New Mexico, suggesting distributional limitations due

to specialization in arid environments.

	 Further collection of the insect groups discussed here, as well as

other taxa within the State of Sonora, will certainly improve our understanding of the complex and intriguing topic of the biogeographic

affinities of Sonoran insects. Increased international cooperation

between scientists in Mexico and the United States is necessary to

coordinate future collection efforts and share systematic knowledge

of the insects of Sonora, Mexico, and, specifically, their Sky Islands.

Many specimens collected from Sonora are already present in collections that also need to be incorporated into present data bases. These

efforts also need to be coordinated for all the surrounding states of

Sinaloa, Chihuahua, and Baja California, which will lead to a more

unified understanding of the natural history of the Northwestern

Mexican-Southwestern United States region.

Acknowledgments

	 We wish to thank Tom Van Devender and John Abbott for help in

acquiring references and comments and to Mahinda Martínez y Díaz

de Salas and Carlos López González for reviews of the manuscript.

References

Abbott,J.C. 2005. Dragonflies and damselflies of Texas and the south-central

United States. Princeton University Press. 344 pp.

Alatore C. and M. Vasquez. 2010. Lista comentada de hormigas

(Hymenoptera:Formicidae) del norte de Mexico. Dugesiana 17: 9-36.

ASUSocialInsectSearchGroup. 2012.Preliminary list of ants(Hymenoptera:

Formicidae)ofSonoraMexico,[Online]Available:http://www.asu.edu/clas/

sirgtools/Ants%20of%20Sonora,%20Mexico,%20list.pdf,[May20,2012]

Bailowitz , R.A. and J. Palting. 2010. Biodiversidad de los insectos con especial énfasis In Lepidoptera y Odonata. In: F.E. Molina-Freaner and T.R.

Van Devender, (eds.). Diversidad Biológica de Sonora. UNAM, Mexico,

pp. 315-337

Barrientos,A.,J.V. Correas, and J. Garcia, eds. 2005. Biodiversidad TamaulipecaVol1. Direccion General de Educacion superiorTecnologica-Instituto

Tecnologico de Cd. Victoria, Tamaulipas,. Mexico. pp 146-149.

Bick, G.H. and W. Mauffray. 2012. Distributin summary of North American

Anisoptera. [Online] Available: http://bellsouthpwp.net/b/i/billmauffray/

nalist.html [May 20, 2012].

Casazza, G.E. Zappa, M.G. Mariotti, F. Medail, and L. Minuto. 2008. Ecological and historical factors affecting distribution pattern and richness of

endemic plantspecies:The case ofthe Maritime and LigurianAlps hotspot.

Diversity and Distributions 14: 47-58.

DeVries, P.J. 1987. The butterflies of Costa Rica and their natural history:

Papilionidae, Pieridae, Nymphalidae. Princenton University Press, U.S.A.

Dingle, H., M.P. Zalucki, and W.R. Rochester. 2000. Relationships among

climate, latitude, and migration: Australian butterflies are not Temperate

Zone birds. Oecologia 124:196-207.

Ehleringer, J.R. 1985. Annuals and perennials of warm deserts. p. 162-180.

In: Chabot, B.F. and H.A. Mooney, eds. Physiological Ecology of North

American Plant Communities. Chapman and Hall, New York.

Endler, J.A. 1982. Problems in distinguishing historical from ecological factors in biogeography. Integrative and Comparative Biology 22: 441-452.

Felger, R.S., M.B. Johnson, and M.F. Wilson. 2001. The trees of Sonora,

Mexico. Oxford University Press, Oxford. pp. 3-28.

Garcia Jimenez, J. and Gonzalo Guevara Guerro. 2005. Mariposas. In: Barrientos, L.L., A. Correa S., J.V. Horta, and J. García J., eds. Biodiversidad

Tamaulipeca. Vol. 1. D.G.E.S.T- Instituto Tecnológico de Cd. Victoria,

Tamaulipas, México. pp. 125-137.

García-Mendoza A. J., M. de J. Ordóñez Díaz, M. Briones-Salas, Abisaí, J.

García-Mendoza. 2004. Mariposas del Estado de Tamauliapas, Redacta S.

A. de C. V., Mexico, D. F. 603 pp.

Garwood,K. andR.Lehman.2005.ButterfliesofNortheasternMexico.Nuevo

Leon, SanLuis Potosi,Tamaulipas.APhotographicChecklist . 2ndEdition.

Eye Scry Publishing, McAllen, Texas. 194 pp.

Halffter, G. 1987. Biogeography of the montane entomofauna of Mexico and

Central America. Annual Review of Entomology 32: 95-114.

Hammer, O., Harper, D.A.T., and P.D. Ryan, 2001. PAST: Paleontological

StatisticsSoftwarePackage forEducationandDataAnalysis.Palaeontologia

Electronica 4(1): 9 pp.

Hewitt, G. 2000. The genetic legacy of the Quartenary ice ages. Nature 405:

907-913.

Hölldobler, B. and E.O. Wilson. 1990. The Ants. Belkap Press of Harvard,

Harvard University Press, Cambridge, MA.

Ives, R. 1949. Climate of the Sonoran desert region.Annals of theAssociation

of American Geographers. 39(3): 143-187.

Llorente,J.E.,A. Luis Martínez,I.Vargas Fernández, and J. Soberón Mainero.

1996. Papilionoidea (Lepidoptera). In: J. Llorente Bousquets,A.N. García

Adrete and E. González Soriano, eds. Biodiversidad, taxonomía y biogeografía de artrópodos de México: Hacia una síntesis de su conocimiento.

Instituto de Biología, UNAM, México, D. F.

Llorente, J.E., L.O. Oñate, A.M. Luis, and I.F. Vargas. 1997. Papilionidae

y Pieridae de México: Distribución Geográfica e Ilustración. Comisión

Nacional Para el Conocimiento y Uso de la Biodiversidad (CONABIO) y

Facultad deCiencias,UniversidadNacionalAutónomadeMéxico,México,

D.F. viii + 227pp.

Marshall, C.J. and J.K. Lieberr. 2000. Cladistic biogeography of the Mexican

transition zone. Journal of Biogeography 27: 203-216.

Molina-Freaner, F.E. and TR. Van Devender. 2010. Diversidad biológica de

Sonora. In: F.E. Molina-Freaner, and T.R. Van Devender, eds. Diversidad

Biológica de Sonora. UNAM, Mexico, pp. 315-337.

Morafka, D.J. 1978. Is there a Chihuahuan Desert? A quantitative evaluation 

through the herpetofaunal perspective. In: Transactions of the Symposium 

of the Biological Resources of the Chihuahuan Desert Region, United States 

and Mexico, U.S. National Park Service Transactions and Proceedings 

Series, 437-454. Washington, DC: Government Printing Office. 

Morrone, J.J. 2002. Mexican biogeographic provinces: preliminary scheme,

general characterizations and synonymies. Acta Zoologica Mexicana 85:

83-108.

Morrone,J.J. and J. Llorente-Bosquets. 2006. Conclusiones. In: Morrone,J.J.

andJ.Llorente-Bosquets, eds.,ComponentesPrincipalesde laEntomofauna

Mexicana. Facultad de Ciencas, UNAM. Mexico, D. F. 1020 pp.

Noonan G.R. 1988. Biogeography of North. American and Mexican insects,

and a critique of vicariance biogeography. Syst. Zool. 37:366–84.

Opler, Paul A., Kelly Lotts, and Thomas Naberhaus, coordinators. 2012.

Butterflies and Moths of North America. [Online] Available: http://www.

butterfliesandmoths.org/. [May 20, 2012].
USDA Forest Service Proceedings RMRS-P-67. 2013 137

Preliminary Assessment of Biogeographic Affinities of Selected Insect Taxa . . . Jones, Obregón-Zuñiga and Guzman-Rodriguez

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Paulson, D.R. and E. González-Soriano. 2012. Mexican Odonata. [Online]

Available: http://www.pugetsound.edu/academics/academic-resources/

slater-museum/biodiversity-resources/dragonflies/mexican-odonata/[May

23, 2012].

Rivera Granados, M. L. 1998. Tesis: Diversidad, distribución y fenología

de los Rhopalocera (Lepidoptera) en algunos municipios del estado de

Querétaro, Licenciatura en Biología, UniversidadAutónoma de Querétaro

Rojas, P. 2001. Las hormigas del suelo en México: diversidad, distribución e

importancia (Hymenoptera: Formicidae).Acta Zoológica Mexicana (n.s.),

Número especial 1: 189-238.

Rzedowski, J. 1978. Vegetación de México. Limusa, México. Dingle, H.,

W. A. Rochester and M. P. Zalucki. 2000. Relationships among climate,

latitude and migration: Australian butterflies are not temperate-zone birds.

Oecologia 124: 196-207

Rzedowski, J. 1993. Diversity and origins of the Phanerogamic flora of

Mexico. In: T. P. Ramamoorthy, R. Bye, A. Lot, and J. Fa, eds. Biological

diversity of Mexico. Origins and distributions. Oxford Univ. Press. New

York, pp. 129-146.

Scott, James A. 1986. The Butterflies of North America, a natural history and

field guide. Stanford University Press. Stanford, CA.

Upson, S. D. Danforth, E. Gonzalez-Soriano, R. A. Behrstock, and R. A.

Balowitz. 2007. Bulletin of American Odonatology 10: 23-51.

VincentJ.Kalkman,ViolaClausnitzer,Klaas-DouweB.Dijkstra,AlbertG.Orr,

Dennis R. Paulson, and Jan van Tol. 2008. Global diversity of dragonflies

(Odonata) in freshwater. Hydrobiologia 595: 351-363.

Warren, A.D., K.J. Davis, E.M. Stangeland, J.P. Pelham, and N. V. Grishin.

2012. Illustrated Lists of American Butterflies. [Online] Available: http://

www.butterfliesofamerica.com/ [May 23, 2012].

Westfall, M.J. and M.L. May. 1996. Damselflies of North America. Scientific

Publishers, Gainesville, Florida.
138 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Preliminary Survey of Bee (Hymenoptera: Anthophila) 

Richness in the Northwestern Chihuahuan Desert

Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

John S. Ascher

Department of Entomology, American Museum of Natural History, New York, New York

Abstract—Museum records indicate that the peak number of bee species occurs around the Mediterranean 

Sea and in the warm desert areas of North America, whereas flowering plants are most diverse in the tropics. We examine this biogeographic pattern for the bee species known from a limited area of northeastern 

Chihuahuan Desert, Mexico/United States. This topographically complex area has been studied for more than 

50 years for bees, which allows us to compare faunas in nearby areas that vary from low elevation desert 

scrub to high elevation montane forest. Our analysis indicates that bee diversity in this area is unusually high, 

and also that there is a poorly documented unique montane fauna.

Introduction

	 Bees (Hymenoptera, Anthophila) are a group of ca. 20,000 species worldwide (Ascher and Pickering 2012; Michener 2007), and

the predominant pollinators in most terrestrial ecosystems. Yet,

paradoxically, given their reliance on flowering plants, the literature

and museum recordssuggest that bee speciesrichness peaksin xeric,

Mediterranean-climate areas far from the wet tropical areas where

most groups, including flowering plants, reach their highest known

species richness. Worldwide diversity of bee species is thought to be

highest in xeric warm-temperate areas of the Western Hemisphere

and around the eastern Mediterranean Sea in the Eastern Hemisphere

(Grace, 2010), and to decrease inmesic environments and towardsthe

tropics (Michener 1979, 2007). Local bee faunas in North America

can be extremely diverse but appear to be particularly so in deserts

of the southwestern United States and northwest Mexico (Ayala and

others 1993; Michener 1979 table 1; Moldenke 1976). Of the roughly

3500 described bee speciesin NorthAmerica north of Mexico, about

75% occur in the topographically diverse western United States. The

most diverse bee faunas are reported forseasonally dry areas, including chaparral (and adjacent vegetation types) in Riverside California

(439 species, Timberlake in Linsley 1958), sparse Sonoran Desert

scrub in and around Palm Springs, California (more than 500 species,

Timberlake, in Michener 1979), chaparral and other habitats in the

inner Coast Ranges of central California (393 species, Messinger &

Griswold2002) andGreatBasinDesertinUtah(334species,Griswold

and others 1997).The bee fauna in lowland subtropical deciduousforest in Chamela, Jalisco in western Mexico has intermediate diversity

(228species,Ayala1988).Incontrast,numbersofbee speciesreported

from mesic areas of eastern and central United States are 64 species

in Miami, Florida (Graenicher 1930), 103 in Hattiesburg, Mississippi

(Michener 1947), 169 in the Chicago, Illinois, area (Pearson 1933),

297 in Carlinville, Illinois(Robertson 1929), and 552 in the large and

topographically extremeBoulderCounty,Colorado (Scott and others

2011). 

	 Although the bee studies listed above suggest that there is high

bee diversity in the xeric areas of western North America, these

studies differ tremendously in the area sampled and duration over

which collections were made. Large or topographically complex

areas can have increased species richness due to greater numbers of

habitats, greater area, or both (Rosenzweig 1995). Sampling that is

limited temporally, for example from one part of the activity season

orfromonly a fewyears, can greatly underestimate local bee diversity

(Williams and others 2001). The purpose of this study is to examine

known bee speciesrichnessfor an unusually well-sampled area ofthe

northwestern Chihuahuan Desert. This is a region of North America

where bee diversity has been thought to be unusually high. One set

of specimen records is associated with activity associated with the

Southwestern Research Station of the American Museum of Natural

History (AMNH). Numerous researchers, including J. G. Rozen, Jr.

and others based at this station have collected and studied bees most

years since the station was founded in 1955. For the past 13 years,

massive collections from individuals based there have been made in

association with theBeeCourse on bee identification and biology that

has been held at the Southwestern Research Station. A second set of

specimens is from an ecological study of bee diversity that has been

underway since 2000 south of the Southwestern Research Station

along the United States-Mexico border in the San Bernardino Valley

(Minckley 2008). Collections in this area were started initially to

estimate bee speciesrichnessin this area using standardizedmethods,

but have broadened in association with other projects (Minckley and

Roulston 2006).

	 Lists of bee species from the two efforts described above are still

increasing as additionalspecies are collected and identified (orrecognized asnew).Despite the preliminary statusofthedata, a compilation
USDA Forest Service Proceedings RMRS-P-67. 2013 139

Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern Chihuahuan Desert Minckley and Ascher

of records assembled to date is useful because sampling from the

region is now unusually large in scope (duration and intensity) and

very large numbers of specimens have been identified to species and

databased. Furthermore, many bee species reach the extreme limits

of their distribution in this area (Burquez 1997; Minckley and Reyes

1996). Here we compile and compare lists of known species from

six well-sampled localities to gain some insight into (1) the number

of species in this area, (2) the differences in the faunas in desert, mid

elevation and upper montane areas, and (3) the future work needed

for a fuller understanding of bee species richness in this region and

its implications for assessing larger-scale patterns.

Methods 

	 Atotalof65,485bee specimenswere includedinthisstudy; allfrom

Cochise County in southeastern Arizona, United States and extreme

northeastern Sonora, Mexico. We limited the species included to

specimensfromsixlocalities(five inCochiseCountywellrepresented

in the AMNH database and the San Bernardino Valley that have

been unusually well-sampled. Thus, other bee species known from

this area (i.e. Cochise County, Arizona, and adjacent northernmost

Mexico) are not considered in this study because they either do not

occur or have yet to be recorded from these localities.These localities

demarcate an area approximately 5700 km2 that is topographically

complex, ranging in elevation from 1100 to 2895 m. Vegetation is

mainly Chihuahuan Desert scrub at the lowest elevations and shifts

through desert grassland, oak woodland, pine-oak, and coniferous

forest at progressively higher elevations (Brown 1994). Collections

were alsomade atunusual,more localizedhabitatsincludinglimestone

outcrops in the San Bernardino Valley and sand dunes near Willcox,

Arizona.

	 For brevity, hereafter we refer to those specimen records that are

being assembled as part of a larger collaborative effort to document

information from specimens housed in a number of North American

museums astheAMNHspecimens or database (see acknowledgements)

Localities

	 Following is a brief description of each of the six localities.

San Bernardino Valley, Arizona/Sonora (31o 20’ N, 109o 15’ W,

1134m)—This is the headwaters of the Rio Yaqui in southeastern

Arizona,UnitedStatesthatrunssouthacrosstheMexico-UnitedStates

borderinto northeasternSonora,Mexico (31o 20’N, 109o 15’W).Most

collections in the United States were made in the San Bernardino

National Wildlife Refuge, a 900-hectare preserve in southeastern

Cochise County, Arizona. In Mexico, most collections were made

at Rancho San Bernardino, 30 km east of Agua Prieta, Municipio of

Agua Prieta, Sonora, Mexico. Elevation rangesfrom 1100 to 1340 m.

Minckley (2008) provides a detailed description of the climate and

area. Vegetation is a mix of creosote bush (Larrea tridentata (Sessé

& Moc. ex DC.), Coville and velvet mesquite (Prosopis velutina

Wooton) dominated Chihuahuan Desert scrub in the lowest elevations, and Chihuahuan Desert grasslands (sensu Brown 1994) at the

higher elevations (see also Minckley, Trajectory and Rate of Desert 

Vegetation Response Following Cattle Removal, this volume).

	 Unlike all other localities, the San Bernardino Valley study extensively used pan traps in addition to net sampling, the method used to

obtain the vast majority of collections at other localities.

Douglas, Cochise County, Arizona(31o 21’N,109o 27’W,1220m)—

Most collectionsin this area are from“1mile east of Douglas” at 1300

m elevation. Specimens with label data from within 6 km of Douglas

in any direction were included in this study because vegetation in

this area is desert scrub similar to that described above for the lower

elevations of the San Bernardino Valley. Although only 25 km due

west of the San Bernardino Valley, this area is in the Sulfur Springs

Valley and separated from the San Bernardino Valley by the southern

end of the Pedregosa Mountains.

	 Cazier and Linsley (1963) provide a description of the site “1 mile

east of Douglas” and the late-summer vegetation. Since that description, some of this area has been developed. Collections span from

1943 to the present. 

Apache, Cochise County, Arizona (31o 41’ N, 109o 08’W, 1336 m)—

Collections included in the area are along Highway 80 that runs

along the north-flowing San Simon River in the San Simon Valley.

Specimens were included if label data indicated they were between

2 km south ofApache to 2 km south of Rodeo, Hidalgo County, New

Mexico (23 km in total). This area is primarily desertscrub similar to

that described for the lower elevations of the San Bernardino Valley

(see above). Apache, Arizona, is 60 km north of the San Bernardino

Valley. They are separated by grasslands on the San Bernardino

volcanic field.

	 This locality along Highway 80 is a primary corridor between the

Southwestern Research Station and Douglas, the nearest city, and

numerous biological studies of bees describe this area (Danforth

1989, 1991, 1999; Hurdand and Linsley 1975; Hurd, LaBerge and

Linsley 1980. Collections span 1956-2011.

Willcox, Cochise County, Arizona(32o 11’N,109o 44’W,1290m)—This

area is along the northeast boundary of the Willcox playa, a closed

basin (i.e. lacking external drainage) and remnant of the Pluvial Lake

Cochise that had been a much larger and deeper lake at its maximum

Table 1—Number of bee species, number (and proportion) of species and collection phenology recorded from six 

Chihuahuan Desert localities. Also shown is the year each locality was first sampled and the numbers of years 

and days bees were made. 

Number and

Number proportion First year Total years Total days

species unique species sampled sampled sampled

San Bernardino Valley 435 223 (0.51) 2000 11 516

Douglas 141 3 (0.02) 1943 41 116

Apache to Rodeo 144 12 (0.08) 1956 39 111

Willcox 155 15 (0.09) 1952 46 150

Apache southwest 129 6 (0.05) 1956 37 134

Upper montane 104 15 (0.14) 1950 43 116
140 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley and Ascher Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern Chihuahuan Desert

13,750-13,400 yr B.P (Waters 1989). The area has typical vegetation

ofthe desertscrub community in addition to other, less common plant

species associatedwiththe sanddunes(Oenothera spp., Psorothamnus 

scoparius (A.Gray)Rydb., Euphorbia spp.,Nama demissum A.Gray,

and Wislizenia refracta Engelm.), and many notable bees (Rozen

1987, 1992; Rozen and Rozen 1986). 

	 The most common locality on labels from this area references “4

mi east of Willcox.” For the species list included here we also included sites in the vicinity of Willcox that are bounded by Interstate

10 on the north and by the Willcox Playa on the south. This locality

is the furthest north of any other in this study. It is ca. 110 km NNW

of the San Bernardino Valley on the opposite side of the Chiricahua

Mountains.

Southwest of Apache, Cochise County, Arizona: (31o 34’ N, 109o

15’W, 1400m)—Thislocality is also along Hwy 80 and wasincluded

because itisthebest-collectedarea intheChihuahuanDesertgrassland

(i.e. at lower elevations than oak and pine woodland) and because it

has a rich and distinctive fauna (Rozen 1989, 1992). Most labelsrefer

to “[13, 13.5, or] 14 milessouthwest ofApache,” a roadside site along

a railway line among cinder cones. It is just north of a riparian area

along Silver Creek, a stream that drains a large area of the southern

Chiricahua Mountains. Higher water availability and cold air drainage in habitats such as these enables some plant species to occur at

lower elevations than they do normally (Shreve 1922), which may

influence where bees occur.

Upper Chiricahua Mountains, Cochise County, Arizona (31o 51’N,

109o 17’ W, 2700m)—Several roads climb to the oak-pine and pine

forestfound on the upper elevations oftheChiricahua Mountains.We

includedanybee speciesinthismountainrange thathadbeencollected

over 1580 m, where oak, oak-pine, or coniferous are the dominant

vegetation type. By using this elevation as a cutoff we excluded a

very large set of specimens collected at the Southwestern Research

Station where disturbance associated with building construction has

createdanunusuallyopenandmorexerichabitat allowingbeestypical

of desert scrub to occur above their usual range.

Bee Species Lists

	 Lists were limited to specimens that were identified to genus and

species. This excluded a number of species from the San Bernardino

Valley study that are now designated as morphospecies. 

	 A complete list of the bee species at each of the six localities will

be published in the journal Checklist (www.checklist.org.br/) and

made available as a Research Species list in Madrean Archipelago

BiodiversityAssessment(MABA)/SouthwestEnvironmentalInformationNetwork (SEINet) online database (Madrean.org).Allspecimens

records in the AMNH Bee Database and associated databases are

available at Discover Life (Schuh and others 2010).

	 We compared the similarities of bee species lists rather than using

statistical approaches based on incidence that species were collected

or species abundance (Gotelli and Colwell 2001) because of the differences in how species lists were assembled at the AMNH and in

the San Bernardino Valley study. Records of species from five sites

(Douglas, Willcox, Apache-Rodeo, 14 miles southwest of Apache,

andUpperChiricahuaMountains)representedin theAMNHdatabase

is ongoing and does not presently include comprehensive records of

all individuals of each species, or the number of times they were collected at each site. Also, retrospective data capture from information

on specimen labels is from identified material, mostly housed in the

AMNH and isin progress. In contrast, all material from every collection made in the San Bernardino Valley is recorded in a database but

species-level identifications ofsome specimensremain to be verified.

Results

Localities

	 The number of bee species at the six localities differed from 435

in the San Bernardino Valley to 104 species in the high elevations

of the Chiricahua Mountains (table 1, fig. 1). The five localities in

the AMNH database have much closer numbers of species among

themselves than any have to the San Bernardino Valley.

	 Approximately55%of allspecies(300of540species),were recorded

at one locality and very few were extremely widespread (fig. 2),

consistent with high bee richness in our study area. The proportion

of unique bee species reported only from the San Bernardino Valley

(52%) was far greater than found in the other localities (fig. 1). In

Figure 1—Bee species recorded from six localities in the northwestern 

Chihuahuan Desert. Locality designations used here are the same as in 

the text except for SBV = San Bernardino Valley. Numbers of species that 

are found only at the locality are indicated on the bar in black.

Figure 2—Number of localities where bee species were recorded. Most 

species occur at one locality and very few are extremely widespread.
USDA Forest Service Proceedings RMRS-P-67. 2013 141

Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern Chihuahuan Desert Minckley and Ascher

descending order, the proportion of unique species at the other localities were montane (30%, 32 of 105 species); Willcox (19%, 30 of

156 species); between Rodeo, New Mexico, andApache,Arizona, on

Highway 80 (8%, 15 of 144 species); southwest of Apache, Arizona

(6%, 8 of 128 species); and Douglas,Arizona (3%, 4 of 120 species).

Vegetation Associations

	 The three major vegetation typesrepresented in the localities were

desertscrub, oak-grassland, and oak pine or coniferousforest. By far,

most bee species were found in one vegetation type (N = 386), none

were reported from two, and few were found in all three (N = 44).

	 The number of unique species recorded from the desert scrub

localities was far greater (N = 390) than reported from grassland (N

= 34) or upper montane (N = 57) (fig. 3). However, the proportion

of unique species was high and similar in the desert (78%) and the

upper montane (72%) habitats, and wasintermediate in the transition

zone of Chihuahuan grassland and oak (41%).

Collecting Effort

	 Collections in the San Bernardino Valley span the fewest years of

any locality but amount to four to five times more collection days

than at other localities (table 1). The other five localities range from

111 to 150 collection days (table 1). The proportion of all collections

represented by the AMNH specimens included in this study and the

San Bernardino Valley are shown in figure 4. AMNH collections are

biased towards August (particularly mid-late August), when the Bee

Course is offered (see Introduction) and fewer collections are in early

summer and spring. In contrast, San Bernardino Valley collections

were made in most months, except for a deficit towards the end of

the bee activity season (late September and October).

Discussion

	 In total, 540 species are represented in this study. In the western

UnitedStates, a four-yearstudy reported 656 bee species andmorphospecies in Grand Staircase National Monument, Utah (Griswold and

others 1997). The Utah study included collections made throughout

the monument. The similar number of bee speciesin thatstudy to just

six localitiesincluded here from the northeastern Chihuahuan Desert

hints at high bee diversity. The San Bernardino Valley, where the

greatest number of bee species was recorded from the six localities,

was where the most collections were made across a wider date range

and where pan traps were used extensively in addition to nets. Pan

traps are known to sample a different portion of the bee fauna than

other methods (Droege and others 2009; Rouston and others 2007).

Thus, more data are needed to establish if the bee diversity in the San

Bernardino Valley is unusual or collecting bias.

	 Few bee species are widespread and many are localized (N =

44 in three vegetation types, none in two, and 386 in one). The

variation in species number among localities reflects both statistical

issues with these data and underlying biological pattern. Statistical

biases arise from the way the species lists are being assembled at

the AMNH. First, database information on AMNH specimens is

ongoing making statistical estimates of species richness or species

turnover not possible. Second, some differences can be attributed to

where taxonomic expertise of specific bee groups has been focused.

Triepeolus, Lasioglossum (Dialictus), and Osmia have been curated

more thoroughly from the San Bernardino Valley than in the AMNH

specimens whereas Andrena, Pseudopanurgus, and Protandrenini 

were more thoroughly curated at the AMNH than those same taxa

from the San Bernardino Valley. Finally, biases in the timing of collections also confound these types of analyses. The proportion of

spring collections is far greater in the San Bernardino Valley than in

collections made at the other localities (fig. 4). Most bee species in

the warm deserts of North America have short flight seasons and are

solitary, particularly those oligolectic species that visit one or a few

host plants for pollen. Approximately 35% of pollen-collecting bee

species(excluding cleptoparasitic species) are oligolectic (Minckley

2008). In solitary bee faunas, the effect of sampling effort should be

pronounced, as will be the presence of host plants required by those

species that are oligolectic. Many of the spring-active bee species

found in the San Bernardino Valley samples are not represented in

the AMNH database.

	 There are many biological reasons why bee species composition

differed among sites. First, rarity in desert beesis pervasive; collections

Figure 3—Bee species recorded from three primary vegetation habitats 

in the northwestern Chihuahuan Desert. Locality designations used 

here are the same as in the text except for Montane = Upper Montane. 

Numbers of unique species (recorded from one locality) are indicated 

on the bar in black.

Figure 4—The phenology of collections made at San Bernardino Valley 

(SBV) and the other five localities (AMNH specimens) expressed as a proportion. Of note, is the difference in collection effort in March and April.
142 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley and Ascher Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern Chihuahuan Desert

that differ in distance and time have all found the proportion of rare

(i.e. rarely-detected) species in deserts is high (Minckley and others

1994; Williams and others 2001). Second, in some cases, rare bees

or those with limited local distributions only occur where their host

plantisfound.Twopollenspecialistbee species,Calliopsis macswaini

(Rozen 1958) andPerdita wislizeniae (Timberlake 1964), were found

only at the locality their host plant, Wislenzia refracta, occurs in the

sanddunesnearWillcox, andthe formeris absentfromrecentsamples.

The extremely rare bee, Macrotera parkeri (Timberlake 1980), is a

cactus specialist species known previously only from Austin, Texas,

(two individuals) and Puebla, Mexico (one specimen) (Danforth

1996). In the San Bernardino Valley it visitsthe cactus, Coryphantha 

robbisorum (W. Earle) A.D. Zimmerman, a limestone soil endemic.

Finally, beta-diversity is expected to be exceptional given that this

study is focused at an intersection of major North American biomes

where many bee species reach their range limits (Minckley and

Reyes 1996). Two notable examples of species in this dataset at the

northernmost point of collection are Eulonchopria punctatissima

Michener, the northernmost-occurring species of Neopasiphaeinae

(formerly included in Paracolletinae or Colletinae sensu lato) that

visits species of Acacia with white flowers (Acacia angustissima

(Mill.) Kuntze and A. millefolia S. Watson) and wasfound in the past

but not in recent years along Highway 80 in Cochise County, and

what is likely to be a transient individual of Agapostemon nasutus

Smith, 1853, a widespread Neotropical species known in the San

Bernardino Valley from only a single female (Mexico: Sonora, 30

km E. Agua Prieta, 29 May 2007, on Sphaeralcea angustifolia, RL

Minckley [coll.]).

	 Identifying why most bee species in this study are geographically

limited or rare is not easily explained. Many species are similar to

the pollen specialist bee Lasioglossum lusorium (Cresson 1872) that

has been recorded in this study only near Willcox, Arizona, but has

a much broader distribution (southern Canada, California, central

Mexico Kansas) (McGinley 2003). 

	 The most notable findings of this preliminary study are the low

species overlap in the desert and uppermontane habitatsin thislimited

area, and the high proportion of unique bee species found in these

same habitats. The Sky Island region above 2300 m elevation are the

northernmost knownlocalitiesfortheNeotropical genus(Mexalictus)

(Eickwort 1978), known in the Chiricahua Mountains only from a

recent collection by JSA (new information) and the southernmost

knownlocalitiesoftwospecieswithHolarctic affinities;Lasioglossum 

boreale (Svensson and others 1977) and L. dasiphorae (Cockerell 1907)

(Packer and Taylor 1997). In Mexico, in the northern Sierra Madre

Occidental at Yécora, one specimen of a stingless bee (Partamona 

bilineata)(Say1837)wascollectedinpine-oakforest,whichrepresents

the most northern record of the tribe Meliponini in North America

(Minckley and Reyes 1996). Ten tropical bee species have been

rarely collected from this area and all but one of these occurred in

oak woodlands or pine-oak forests of Sky Islands along the ArizonaMexico border (Minckley and Reyes 1996). Recent collections from

the oak woodland in the Sierra San Luis, just 3 kmsouth ofthe United

States-Mexico border (31o 17’ N, 108o 47’ W), have yielded two new

species of spring-active species of Osmia (Rightmyer and Griswold

2010).Anewspecies ofLasioglossum (L. viridipetrellumGibbs2009)

is known only from Sky Islands above 2200 m in southeast Arizona

(Gibbs 2009), asis an undescribed species allied to L. (D.) ruidosense

(J. Gibbs, pers. comm.). These recent descriptions, in addition to

the study reported here, suggest there is greater bee endemism in

the upper montane habitats of these areas than has been appreciated.

Given the distinctiveness of this bee fauna, the predicted increasing

frequency of fires(Westerling and others 2007), and effects of climate

change (International Panel on Climate Change 2007) on these upper

elevation habitats, further documentation and sustained monitoring

of this bee fauna is of particular and pressing interest.

Acknowledgments 

	 D. Scantlebury (University of Rochester) and J. Gibbs (Cornell

University) kindly reviewed the manuscript. R.L. Minckley was

supported by grants from DEB0418871from the National Science

Foundation, Southwest Consortium for Environmental Research and

Policy,WorldWildlife Foundation,Chihuahuan Desert Program.J.S.

Ascher wassupported by a grant DBI0956388 from the National Science Foundation. Jerome G. Rozen, Jr. made the largest contribution

to collecting bees for the AMNH dataset.

References

Ascher, J.S.; Pickering, J. 2012. Discover Life bee species guide and

world checklist (Hymenoptera: Apoidea: Anthophila). Draft 33.[online]

Bee species guide: http://www.discoverlife.org/mp/20q?guide=Apoidea_

species &flags=HAS:

Ayala, R.L. 1988. Abejas silvestres (Hymenoptera: Apoidea) de Chamela,

Jalisco, México. Folia Entomologica Mexicana 77: 395-493.

Ayala, R.L.; Griswold, T.L.; Bullock, S.H. 1993. The native bees of Mexico.

In T. P. Ramamoorthy, R. Bye, A. Lot, and J. Fa (eds.), Biological Diversity of Mexico: Origins and Distribution. Oxford University Press, New

York and Oxford.

Brown, D.E. 1994. Biotic communities: southwestern United States and

northwestern Mexico. University of Utah Press, Salt Lake City.

Burquez, A. 1997. Distributional limits of Euglossine and Meliponine bees

(Hymenoptera: Apidae) in northwestern Mexico. Pan-Pacific Entomologist 73:137-140.

Cazier,M.A.;Linsley,E.G.1963.Territorialbehavior amongmalesofProtoxea 

gloriosa (Fox) (Hymenoptera: Andrenidae). The Canadian Entomologist

95: 547-556.

Cockerell, T.D.A. 1907. The bees of Boulder County, Colorado. University

of Colorado Studies 4: 239-259.

Danforth, B.N. 1989. Nesting behavior of four species of Perdita (Hymenoptera:Andrenidae).Journal of the Kansas Entomological Society 62: 59-79.

Danforth, B.N. 1991. Female foraging and intranest behavior of a communal

bee, Perdita portalis (Hymenoptera: Andrenidae). Annals of the Entomological Society of America 84: 537-548.

Danforth, B.N. 1996. Phylogenetic analysis and taxonomic revision of the

Perdita subgenera Macrotera, Macroteropsis, Macroterella and Cockerellula (Hymenoptera: Andrenidae). Kansas Science Bulletin 55:635-692.

Danforth, B.N. 1999. Emergence dynamics and bet hedging in a desert bee,

Perdita portalis. Proceedings of the Royal Society of London B 266:

1985-1994.

Droege, S.; Tepedino, V.; Griswold, T.G.; [and others]. 2009. Spatial patterns

of bee capturesin NorthAmerican bowl trapping surveys.InsectConservation and Diversity 3: 15-23.

Eickwort, G.C. 1978. Mexalictus, a new genus of sweat bees from North

America (Hymenoptera: Halictidae, Halictinae). Journal of the Kansas

Entomological Society 51:567-580.

Gibbs, J. 2009. Integrative taxonomy identifies new (and old) species in the 

Lasioglossum (Dialictus) tegulare (Robertson) species group. Zootaxa 

2032: 1-38.

Giles, V.; Ascher, J.S. 2006. A survey of the Black Rock Forest Preserve,

New York (Hymenoptera: Apoidea). Journal of Hymenoptera Research

15: 208-231.

Gotelli, N.J.; Colwell, R.K. 2001. Quantifying biodiversity: procedures and

pitfallsin themeasurement ofspeciesrichness.EcologyLetters 4: 379-391.

Grace, A. 2010. Introductary biogeography to bees of the eastern Mediterranean and Near East. Bexhill Museum. Sussex, United Kingdom. 284 pp.
USDA Forest Service Proceedings RMRS-P-67. 2013 143

Preliminary Survey of Bee (Hymenoptera: Anthophila) Richness in the Northwestern Chihuahuan Desert Minckley and Ascher

Graenicher, S. 1930. Bee fauna and vegetation of the Miami region of Florida.

Annals of the Entomological Society of America 23:153-174.

Griswold,T.;Parker,F.D.; andTepedino,V.G. 1997.The bees oftheSanRafael

Desert: implications for the bee fauna of the Grand Staircase-Escalante

National Monument. Pp. 175-186 in L. M. Hill (ed.) Proceedings of the

Grande Staircase-Escalante National Monument Science Symposium, vol

BLM/UT/GI-98/006+1200. U.S. Department of the Interior, Bureau of

Land Management, Salt Lake City, UT.

Hurd, Jr., P.D.; LaBerge, W.E.; and Linsley, E.G. 1980. Principal sunflower

bees of North America with emphasis on the southwestern United States

(Hymenoptera:Apoidea).SmithsonianContributionstoZoology310:1-158.

Hurd, Jr., P.D.; Linsley, E.G. 1975. The principal Larrea bees of the southwesternUnitedStates(Hymenoptera:Apoidea).SmithsonianContributions

to Zoology 193: 1-74.

International Panel on Climate Change. 2007. IPCC website http://ippc-ddc.

cru.ued.ac.uk [online]

Linsley, E.G. 1958. The ecology of solitary bees. Hilgardia 27: 543-599.

Messinger, O.; Griswold, T. 2002. A pinnacle of bees. Fremontia 30: 32-40

Michener,C.D. 1947.Bees of a limited area in southernMississippi.American

Midland Naturalist 38:443-455.

Michener, C.D. 1979. Biogeography of the bees. Annals of the Missouri

Botanical Garden 66:277-347.

Michener, C.D. 2007. The Bees of the World.John Hopkins University Press,

Baltimore, MD.

Minckley, R.L. 2008. Faunal composition and species richness differences

of bees (Hymenoptera: Apiformes) from two North American regions.

Apidologie 39: 176-188.

Minckley R.L.; Cane, J.H.; Kervin, L.; and Roulston, T.H. 1999. Spatial predictability and resource specialization of bees (Hymenoptera : Apoidea) at

a superabundant, widespread resource. Biological Journal of the Linnaen

Society, 67:119-147.

Minckley, R.L.; Roulston,T.H. 2006. Incidental mutualisms and pollen specialization among bees. Pp. 69-98. in N. M. Waser and J. Ollerton (eds.),

Plant-Pollinator Interactions: From Specialization to Generalization.,

University of Chicago Press, Chicago, IL.

Minckley, R.L.; Reyes, S.G. 1996. Capture of the orchid bee, Eulaema polychroma, (Friese) (Apidae: Euglossini) in Arizona, with notes on northern

distributions of other Mesoamerican bees. Journal of the Kansas Entomological Society 62:102-104.

Minckley, R.L.;Wcislo,W.T.;Yanega, D.A.; Buchmann, S.L. 1994. Behavior

and phenology of a specialist bee (Dieunomia) and sunflower (Helianthus)

pollen availability. Ecology. 75: 1406-1419.

Mitchell, T.B. 1960. Bees of the eastern United States. I. Tech. Bull. 141.

North Carolina Agricultural Experiment Station. 538 p.

Mitchell, T.B. 1962. Bees of the eastern United States. II. Tech. Bull. 152.

North Carolina Agricultural Experiment Station. 557 p.

Moldenke, A.R. 1976. California pollination ecology and vegetation types.

Phytologia 34:305-361.

Packer,L.;TaylorJ.S.1997.Howmanyhiddenspecies are there?Anapplication

of the phylogenetic species concept to genetic data forsome comparatively

well known bee “species.” Canadian Entomologist 129: 587-594.

Pearson, J. F. W. 1933. Studies on the ecological relations of bees in the

Chicago region.Ecological Monographs 3: 373-441.

Rightmyer Gee, M. and T. Griswold. 2010. Descriptions of two new species

ofOsmia(Hymenoptera:Megachilidae)fromsouthwesternNorthAmerica.

Zootaxa 2512: 26-46.

Robertson, C. 1929. Flowers and Insects: Lists of Visitors to Four Hundred

and Fifty-three Flowers. Publ. by author, Carlinville, Illinois.

Roulston,T.H.;Smith,S.A.; andBrewster,A.L.2007.Acomparisonofpantrap

andintensivenetsamplingtechniquesfordocumentingabee (Hymenoptera:

Apiformes)fauna.JournaloftheKansasEntomologicalSociety80:179-181.

Rosenzweig, M.L. 1995. Species diversity in space and time. University

Press, Cambridge.

Rozen, J.G., Jr. 1987. Nesting biology and immature stages of a new species

in the bee genus Hesperapis (Hymenoptera: Apoidea: Melittidae: Dasypodinae). American Museum Novitates 2287: 1-13.

Rozen,J.G.,Jr.1989.Twonewspeciesandtheredescriptionofanotherspeciesof

the cleptoparasitic bee genusTriepeoluswith notes on theirimmature stages

(Anthophoridae: Nomadinae). American Museum Novitates 2956: 1-18.

Rozen, J.G., Jr. 1992. Systematics and host relationships of the cuckoo bee

genus Oreopasites(Hymenoptera:Anthophoridae:Nomadinae).American

Museum Novitates 3046: 1-56.

Rozen, J.G., Jr.; Rozen, B.L. 1986. Bionomics of crepuscular bees associated

with the plant Psorothamnus scoparius (Hymenoptera: Apoidea). Journal

of the New York Entomological Society 94: 472-479.

Scott, V.L.; Ascher, J.S.; Griswold, T.; Nufio, C.R. 2011. The bees of Colorado (Hymenoptera: Apoidea: Anthophila). Natural History Inventory

of Colorado No. 23. University of Colorado Museum of Natural History.

Boulder, CO. 100 p.

Schuh, R. T., Hewson-Smith, S., and J. S. Ascher. 2010. Specimen databases:

a case studyinentomologyusingweb-basedsoftware.AmericanEntomologist 56(4): 206-216.

Shreve, F. 1922. Conditionsindirectly affecting vertical distribution on desert

mountains. Ecology 3: 269-274.

Svensson,B.G.;Ebmer,A.W.; Sakagami, S.F. 1977.Lasioglossum(Evylaeus)

boreale, anewHalictinae (Hymenoptera:Apoidea)speciesfoundinnorthern

Sweden and on Hokkaido, Japan, with notes on its biology. Entomologia

Scandinavica. 8: 219-229.

Tarnocai, C. 1984. Characteristics of soil temperature regimes in the Inuvik

area. In: Olson, R.; Hastings, R.; Geddes, F., eds. Northern ecology and

resource management. University of Alberta Press: 19-37.

Waters, M.R. 1989. Late Quaternary lacustrine history and paleoclimatic

significance of Pluvial Lake Cochise, southeastern Arizona. Quaternary

Research 32: 1-11.

Westerling, A.L.; Hidalgo, H.G.; Cayan, D.R.; and Swetnam T.R. 2006.

Warming and Earlier Spring Increase Western U.S. Forest Wildfire Activity. Science 313: 940-943

Wilson,J.S.;Griswold,T.;andMessinger,O.J.2008.Samplingbeecommunities

(Hymenoptera: Apiformes) in a desert landscape: Are pan traps sufficient?

Journal of the Kansas Entomological Society 81:288-300.

Williams, N. M.; Minckley, R.L.; and Silveira, F.A. 2001. Variation in native bee faunas and its implications for detecting community changes.

Conservation Ecology 5. Online: http://www.consecol.org/vol5/iss1/art7.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
144 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Introduction to the Arizona Sky Island Arthropod 

Project (ASAP): Systematics, Biogeography, Ecology, and 

Population Genetics of Arthropods of the Madrean Sky 

Islands

Wendy Moore, Wallace M. Meyer, III, Jeffrey A. Eble, and Kimberly Franklin

Department of Entomology, University of Arizona, Tucson, Arizona

John F. Wiens and Richard C. Brusca

Arizona-Sonora Desert Museum, Tucson, Arizona

Abstract—The Arizona Sky Island Arthropod Project (ASAP) is a new multi-disciplinary research program 

at the University of Arizona that combines systematics, biogeography, ecology, and population genetics to 

study origins and patterns of arthropod diversity along elevation gradients and among mountain ranges in 

the Madrean Sky Island Region. Arthropods represent taxonomically and ecologically diverse organisms 

that drive key ecosystem processes in this mountain archipelago. Using data from museum specimens and 

specimens we obtain during long-term collecting and monitoring programs, ASAP will document arthropod 

species across Arizona’s Sky Islands to address a number of fundamental questions about arthropods of this 

region. Baseline data will be used to determine climatic boundaries for target species, which will then be 

integrated with climatological models to predict future changes in arthropod communities and distributions 

in the wake of rapid climate change. ASAP also makes use of the natural laboratory provided by the Sky 

Islands to investigate ecological and genetic factors that influence diversification and patterns of community 

assembly. Here, we introduce the project, outline overarching goals, and describe preliminary data from the 

first year of sampling ground-dwelling beetles and ants in the Santa Catalina Mountains.

Introduction

	 The outcome of millions of years of mountain building, the 7250

km long North American Cordillera or “Western Cordillera” runs

from northern Alaska to southern Mexico. This great cordillera, the

spine of the North American continent, has but one break, a low

saddle betweentheRockyMountains/ColoradoPlateau and theSierra

Madre Occidental, which forms a biogeographic barrier between the

montane biotas oftemperate and tropicalNorthAmerica (Heald 1951;

Marshall 1957; McLaughlin 1986, 1995; Warshall 1995; Bowers

and McLaughlin 1996). The Madrean Sky Islands are the isolated

mountain ranges that span this Cordilleran Gap. Sometimes called

the “Madrean Province” or “Madrean Archipelago,” these mountain

ranges are a unique subset oftheBasin andRange Province and cover

about 168,400 sq. km. (~77,700 sq. km. in the United States).

	 Although many of the plants and some of the animals of the Sky

Island Region have been well studied, little is known about the

arthropods(e.g.,insects,isopods,millipedes,mites,spiders,scorpions,

etc.). Yet, these ecologically diverse organisms drive key ecosystem

processessuchaspollination,litterdecomposition,nutrientrecycling,

and soil aeration, and they are important food for reptiles, birds, and

small mammals. Despite their ecological importance, arthropods are

poorly known because most are small, live in opaque habitats where

observation is difficult, and few taxonomists have specialized on

these groups in the Sky Island Region (Behan-Pelletier and Newton

1999). Using data from museum specimens and specimens we obtain

during long-term collecting and monitoring programs, ASAP will

document arthropod species acrossArizona’s Sky Islandsto address a

number of fundamental questions about the arthropods of thisregion,

including the following:

1. What arthropods inhabit Arizona’s Sky Islands? How are these

species distributed within the region?

	 One of the greatest resources for this project is the University of

Arizona Insect Collection (UAIC), which contains over two million

researchspecimens,83%ofwhichareidentifiedtospecieslevel,mostly

fromthe Sonoran DesertRegion, its Sky Islands, and adjacent biomes

of northwestern Mexico. In the last few decades, UAIC researchers

have participated in long-term surveys in the Santa Catalina, Baboquivari, Mule, Huachuca, Chiricahua, andWaterman Mountains, and

other montane areas; as well as in impact studies of the Mt. Graham

telescope site in the Pinaleño Mountains and the Rosemont Mine site

in the SantaRitaMountains.Locality data fromUAICspecimens will
USDA Forest Service Proceedings RMRS-P-67. 2013 145

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

be incorporated into a specimen-level database for theASAPProject;

this database also will be networked with other arthropod museums

in the southwestern United States.

	 SpeciesdiversityoftheSkyIslandRegionis alsobeingdocumented

through a series of collecting expeditions. Throughout the project,

we will conduct surveys of the arthropod fauna using a variety of

collecting methods including light traps, malaise traps, pitfall traps,

and hand collecting methods. All specimens from the ASAP project

will be accessioned into the UAIC. They will be identified to species,

and specimen-level data will be maintained in the UAIC specimenlevel database. Representatives of many species will be preserved in

a frozen tissue collection in -20 °C freezers and all other specimens

collected will be mounted on points/pins and deposited in the UAIC

pinned collection or preserved in 80% ethyl alcohol and archived for

future work.

2. What are the biogeographic affiliations and phylogeographic relationships of the Sky Island arthropods? Have arthropods diversified

or radiated within the Sky Island Region?

	 Thedistributionand/or elevationofMadreanSkyIslandspecieshas

repeatedly shifted as a result of cyclical climate changes during the

Quaternary (Martin 1963;Betancourt and others 1990;Van Devender

and Spaulding 1990; Davis and Brown 1988; Van Devender 2002).

Several studies have focused on the biogeography of species in the

Arizona Sky Island Region including plants, arthropods, birds, lizards, and mammals(Brown 1971; Downie 2004; Linhart and Permoli

1994; McCord 1995; Sullivan 1994; Slentz and others 1999; Barber

1999a,b; Maddison and McMahon 2000; Masta 2000; Boyd 2002;

Smith and Farrell 2005a,b; McCormack and others 2008; Tennessen

andZamudio 2008; Ober and others 2011).Most ofthese studies have

documented significant morphological variation or genetic structure

among separate subspecies or populations endemic to different

mountain ranges within the Sky Island Region. However, none of

these studies have used broad taxonomic sampling in the context of

phylogeographic analyses to look at the geographic origin of the Sky

Island fauna. In ASAP, we will infer the evolutionary relationships

and estimate divergence times using molecular sequence data from

fast-evolving mitochondrial and nuclear genes. From these data, we

will reconstruct the evolutionary and biogeographic history of the

Sky Island arthropod species, and look for emergent patterns that

may reveal phylogeographic origins of the fauna and radiations of

species groups among the mountain ranges.

3. What are the over-arching drivers that structure arthropod communities in the Sky Islands?

	 The spatial arrangement of similar, isolated habitats repeated on

numerous mountain rangesin the Sky Island Region provides natural

replication for ecological studies. We will utilize the natural laboratory provided by the Sky Islandsto investigate ecological and genetic

factors that influence diversification and patterns of community assembly. In particular we will address the following questions: (1)

How are species distributed along elevational gradients? (2) Are the

patterns correlatedwithplant communities/biomes,soilpH,precipitation, temperature, and/or humidity? (3) Do high elevation arthropod

communitiesdifferinspecies compositionbetweenisolatedmountain

ranges? (4) Do larger tracks of woodland or forest harbor a greater

diversity of arthropods? To address these questions, ASAP makes

the first effort to extensively and quantitatively sample the arthropod

fauna for a broad swath of the Madrean Sky Islands. It will provide

baseline data on the abundance, diversity, and community structure

alongelevationgradientsthroughoutthe regionforthistaxonomically

and ecologically diverse group of organisms.

4. How might montane arthropod communities respond to rapidly

changing climate conditions?

	 The Madrean Sky Islands provide amodelsystemforinvestigating

the effects of rapidly changing climate on the biodiversity of isolated

landscapes. Sky Islandsshow a stronger effect of isolation than other

types of isolated habitats (Watling and Donnelly 2006). Effects of

climate change are already evident here (increased fire, drought, pest

outbreaks, and invasive species).Additional temperature increases of

as little as a few degrees could push Sky Island plant and arthropod

species/communities to higher elevations, reducing their habitable

area and potentially causing local extinctions of endemic taxa and

evolutionarilyuniquelineages.Projectionsfromglobalclimatemodels

suggest that temperatures in the Southwest will increase by 3-6 °C

over the next century (Kupfer and others 2005; Gutzler and Robbins

2010; Overpeck and Udall 2010; Dominguez and others 2010; IPPC

2001,2007;CLIMAS2012).Predictions ofwhetherprecipitationwill

increase or decrease are still in conflict, and confidence in precipitation estimates is low at this time. Models used to examine how plant

communities may respond to projected climate change indicate that

increasesin temperature will lead to upslope movement of communities,leading to an increased area ofDesertscrub (westernSky Islands)

or Desert Grassland (eastern Sky Islands) and a decrease in the area

occupied by Mixed Conifer Forest, the biome situated at the top of

the highest mountains(Kupfer and others 2005). However, increased

precipitation might slow upslope movements by lessening growth

constraints imposed by arid conditions. If climates continue to warm

as projected, species will need to shift their present distributions or

adapt to warmer conditions rapidly.

	 Anthropogenic activities are contributing to increased levels of

atmospheric CO2 and these higher levels are leading to increased

global temperatures and changesin the hydrologic cycle, thus causing

or contributing to climate change (IPCC2001, 2007).Severalreviews

have highlighted that little is known about the current and predicted

responses to climate change for most arthropod groups (Coviella

and Trumble 1999; Hughes 2003). Because important ecosystem

processes, such as litter decomposition and nutrient cycling, are

driven by ground-dwelling arthropods(Powers and others 2009;Yang

and Chen 2009), ecosystem responses and subsequent feedbacks to

atmospheric and climate changes may depend on how soil arthropod

communitiesrespondtotheseperturbations(e.g.,reduceddecomposition rates will decrease feedbacks to global carbon (C) input, while

increased decomposition rates will increase feedbacksto the C cycle;

Bardgett and others 2008). As such, understanding the response of

these species and communities to climate change is critical if we are

to predict how such perturbations are going to alter both biodiversity

and the functioning of ecosystems.

	 While natural communities may be able to adapt to changing

climates to some extent, via shifts in the distribution of their local or

geographic ranges, paleontological data suggest that communities do

not respond to climate change as entities(Coope 1995;Jablonski and

Sepkoski 1996). Rather, species responses tend to be idiosyncratic,

resulting in the development of new assemblages and associations

(Graham and Grimm 1990; Voight and others 2003). These results

underscore the need for field-based inventories to better understand

current diversity and distribution patterns, which in turn establishes

a baseline for future comparisons and ecological studies to track

how species and communities respond to future climate change in

the Southwest.

	 We will use data acquired from our own collecting and from specimens housed inmuseumcollectionsincluding the UAICto determine

climatic boundaries for target species; species-specific boundaries
146 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

will then be integrated with climatological modelsto predict changes

in arthropod communities and distributions in the wake of potential

rapid climate change.To date, the scientific community knows almost

nothingabouthowarthropodspecies andcommunitieswillrespondto

climate change.But because changesin both species composition and

abundancesofarthropodscaninfluenceimportantecosystemprocesses,

it is vital to understand how arthropodsspecies and communities will

respond to climate change and how arthropods influence ecosystem

processes. Additionally, because climate change is predicted to lead

to a widespread reorganization of species and community patterns, it

is important to document the biogeography of Sky Island arthropod

species now, to determine patterns of endemism and identify species

at risk of extinction.

	 TheASAPprojectwillmonitordistributionsofSkyIslandarthropod

species over the next 20–30 years to see if there are detectable shifts

in species distributions as climate changes. We will use ecological

niche models and phylogeographic analysesin combination to assess

how arthropods have responded to past changes in climate by shifting distributions and/or adapting in situ, which in turn will help us

predict how species will likely respond to increasing isolation and

future climate change. Ultimately, we aim to identify species in the

Sky Island Region that might be at risk of extirpation or extinction

as the Southwest warms, and to identify locations of refugia from

past glaciation events that may be areas of high allelic diversity that

merit conservation attention.We further predict that effects of climate

change may be seen in thisregion of the country and on these isolated

mountains before most other areas, and that by studying the effects of

climate change here we can make better predictions as to what will

happen elsewhere later in time.

Purpose of This Paper

	 The purpose of this paper is to describe the study system of the

ASAP, including our concept ofthe Sky IslandRegion and its biomes,

and to present preliminary data derived from our collections in the

Santa Catalina Mountains during the first year of the project. In the

initial phase of the project we are documenting the ground-dwelling

arthropods ofthe SantaCatalinas, and the first year’ssampling (2011)

has been completed. The project will be expanded to other ranges in

southern Arizona, and eventually to selected ranges in Mexico and

to other arthropods (not just ground-dwelling taxa). Data collected

in early years of the project will provide a baseline for future/continuing surveys designed to elucidate how populations, species, and

communities change as climate changes in the Southwest.

Biogeographic Parameters of the Study 

Boundaries and Concept of the Sky Island Region—When the

term “Madrean Sky Islands” was first coined in the 1950s, it referred

to those ranges possessing strong shared elements with the Sierra

Madre Occidental (the “Madrean” flora). In fact, the term “Madrean

Sky Islands” is not a perfect descriptor of the region, because many

northern (e.g., Rocky Mountain or Petran) (Brown and others 1979;

Brown 1994) species reach their southernmost range limits here, just

as many southern (e.g., Madrean) species reach their northernmost

range limits in this region. Although we are not suggesting a name

change here, conceptually it might be more accurate to think of these

mountain ranges collectively as “Cordilleran Gap Sky Islands” and

for the ASAP project we broaden the definition of the Madrean Sky

Islands to include all ~65 of the isolated mountain ranges spanning

the Cordilleran Gap (that are high enough to have Oak Woodlands)

regardlessofthedegreeofsharedMadreanflora (fig.1).We,therefore,

include the northern Pinal and Superstition Mountains(Arizona) and

the easternBig and Little Hatchet andAlamo Hueco Mountains(New

Mexico),which are traditionally notincluded onmaps oftheMadrean

Sky Islands. We define the northern boundary of the Madrean Sky

Island Region as the Salt River Valley; north of this lie the Mazatzal

and Bradshaw Mountains, and other ranges of Arizona’s Transition

Zone that grade into the base of the escarpment of the Colorado

Plateau’s Mogollon Rim. The southernmost limit of the Sky Island

Region is somewhat arbitrarily set at 29° N latitude (about the latitude of Hermosillo, Sonora). So, the northernmost Sky Islands are

the Pinals and Superstitions; the westernmost Sky Islands are Sierra

el Humo (in Sonora) and the Baboquivari Mountains (in Arizona);

the easternmost are the Alamo Hueco and Big Hatchet Mountains of

New Mexico; and the southernmost Sky Island is the Sierra Mazatán

70 km east of Hermosillo, Sonora.

	 Nearly all of the Madrean Sky Islands exceed 1525 m in elevation,

or if falling short of that, they are connected by woodland habitat to

an adjacent higher range and thus form part of a Sky Island complex.

The Sierra el Humo reaches an elevation of 1650 m and has ~1036

ha of Oak Woodland (Flesch and Hahn 2005). Two small ranges to

the west of Sierra el Humo have oak patches, but they do not exceed

1370 m elevation and do not have sizeable Oak Woodlands (Sierra

El Cobre, 1350 m; Sierra El Duranzo, 1210 m). The Sierra Mazatán

reaches only 1545 m, but supports a large area of Oak Woodland

(~3626 ha, with five oak species) at its highest elevations, and it is

surrounded by a “sea” offoothillsthornscrub (Flesch and Hahn 2005;

Dimmitt and others 2011).

	 The northernmost Sky Island ranges (the Pinal and Superstition

Mountains)largelylackcommonMadreanspeciessuchasthesilverleaf

oak (Quercus hypoleucoides) that is so common in more southern

ranges ofArizona. Instead, they harbor woodland of northern species

such as Gambel oak (Quercus gambelii) the only winter-deciduous

oak in the Sky Island Region, and Palmer oak (Quercus palmeri).

The Pinals, which reach over 2380 m in elevation, support a large

Petran Montane Pine Forest, surrounded by well-developed Oak

Woodlands that connect to the woodlands of the Superstitions, the

latter having only a few patches of ponderosa pine (the highest peak

in the Superstitions is 1900 m).

	 In addition to these 65 ranges (fig. 1), several other small ranges

in Mexico are apparently high enough to support Oak Woodlands,

notably a few east of the Sierra Mazatán. However, plant data for

these ranges are too sparse to draw firm conclusions at this time.

We do not consider ranges with only scattered oak patches but

lacking true Oak Woodland— such as Arizona’s Tucson Mountains

(Wasson Peak, 1429 m)— to be Sky Islands. Similarly, many of the

small mountains in Sonora, west of the Sierra Madre Occidental,

are high enough to support scattered patches of oaks, especially on

their north-facing slopes (e.g., Sierra Cucurpe), but in general these

are not large or contiguous encinals (Oak Woodlands). To the east,

we include all of the ranges of appropriate elevation and isolation

west of the Continental Divide, plus three east of the divide (Big and

Little Hatchets and Alamo Hueco in New Mexico). Some authors

have included the far eastern Cedar Mountains of New Mexico and

others have included the Sierra San Javier, 130 km east-southeast of

Hermosillo. Although included on some Sky Island maps, we do not

include Sonora’s huge Sierra el Tigre mountain complex (i.e., Sierras

El Tigre, San Diego, el Oso, and los Pilares de Teras) because the

woodlands of those ranges connect to the Sierra Madre Occidental

south of Huachinera (via the Mesa Los Tabachines, Sierra El Gato,

and Sierra Los Tules), which connect to the Sierras at elevations over

1525m, effectivelymaking the SierraElTigre complex a “woodlands
USDA Forest Service Proceedings RMRS-P-67. 2013 147

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Figure 1—Elevation map of the Sky Island Region with 65 Sky Island mountains labeled. Green shading indicates approximate area above 1600 m

(5250 ft), roughly the elevation at which oak woodlands first begin, although there is considerable variation in this due to latitude, slope and aspect,

rainfall patterns, etc. (Map based on cartographic GIS research by Joel Viers/Lirica.)
148 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

peninsula” of the Sierra Madre Occidental. Thissaid, it is known that

thesewoodlandconnectionshavebeengreatlydiminishedbydrought,

fires, and tree cutting and in the near future this “peninsula” may

become an “island” as it is surrounded on the other three sides by the

RíoBavispe. Nevertheless,for historical biogeographic purposes, the

El Tigre mountain complex is part of the Sierra Madre. South of the

huge Sierra el Tigre complex in Sonora, thornscrub surrounds most

of the ranges south to the Sierra de Mazatán.

	 The “Apachian Floristic District” (of McLaughlin 1995) and the

“Apachian Subprovince” (of the larger Madrean Floristic Province)

largely coincide with our definitions of the Madrean Sky Island Region. The 12.146-million ha “Apache Highlands Ecoregion” (of The

Nature Conservancy) somewhat exceeds our view of the Sky Island

Region in that it extends northwest through the Verde River and Big

Chino Valleys of central Arizona to the Mogollon Rim, and south to

include the northernmost region of the Sierra Madre Occidental.

Mountain Islands vs. Habitat Islands—A mountain range in this

region istraditionally defined as a “Sky Island” if it is high enough to

includeOakWoodlandhabitat andisnot connectedbyOakWoodlands

to the Cordilleran ranges of the Rocky Mountains/Colorado Plateau

or the Sierra Madre Occidental (Heald 1951; Dimmitt and others

2011). Using this definition, we recognize ~65 mountain ranges that

should be designated as Sky Islands, 32 of which are in the United

States (table 1).

	 In reality, 18 of the 32 named mountain ranges north of the border

are connected to at least one other range by contiguous Oak Woodland and thus these 32 ranges can be classified into 21 distinct Sky

Island complexes (table 2). Many of the Sonoran Sky Islands are

connected to one another by Oak Woodland also, and, in the future,

with improved information these could be combined into mountain

complexes as we’ve done in table 2.The 21 U.S.mountain complexes

that are connected by Oak Woodland can be thought of, ecologically

and biogeographically as 21 “Oak Woodland islands” or “habitat

islands.” Many of these mountain ranges are high enough to also

harbor higher elevation biomes (e.g., Pine-Oak Woodlands, Chaparral, Pine Forest). However, to date, we have not determined how

many contiguous vs. isolated landscapes of these higher elevation

biomes exist. These concepts have considerable relevance for the

biogeography of arthropods that inhabit these montane biomes.

	 Inaddition,someSkyIslandmountainranges actuallycomprise two

ormore subranges, eventhoughmaps andcommonusage typicallyuse

the name of only the largest of these. For example, the Baboquivari

Mountains actuallycomprise fourranges—theCoyote,Quinlan,Pozo

Verde, and Baboquivari Mountains—separated from one another by

distinct valleys. But, popular usage refers to them collectively as the

Baboquivari Mountains. This situation is especially troublesome in

Mexico where we have tried to determine the most commonly used

name for such ranges. Many Mexican ranges also have more than

one name due to local usage, etc., and we have relied on the INEGI

(Instituto Nacional de Estadística y Geografía) 1:250,000 maps of

Sonora and Chihuahua for these names. We used USGS and INEGI

information, and a Garmin 62S GPS device, for estimating elevation

data.
USDA Forest Service Proceedings RMRS-P-67. 2013 149

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Biomes of Arizona’s Sky Islands

	 Ecologists generally agree that there are about a dozen major plant

community types that occur worldwide, e.g., tundra/alpine, temperate broadleaf forests, Mediterranean woodlands/shrublands, tropical

rain forests, coniferous forests, temperate and tropical Grasslands,

temperate and tropical savannahs, tropical dry forests, Chaparral,

savannahs, and deserts. Examples of all but the first four are found

in the Madrean Sky Island Region. We refer to these as “biomes” in

this study.

	 Here, we follow theWhittaker and Niering (1968b),Whittaker and

others(1968), and Niering and Lowe (1985)studies of the vegetation

of the Santa Catalina Mountainsin recognizing the following biomes

for the Sky Islands of Arizona: Desertscrub, Desert Grassland, grazing disturbed Grassland, Oak-Grassland, Oak Woodland, Pine-Oak

Woodland, Chaparral, Pine Forest, and Mixed Conifer Forest. Whittaker and Niering (1965, 1968) listed the signature species for each

of these biomes in their now classic diagrams depicting the biomes

of the Santa Catalina Mountains. Not all of the Madrean Sky Islands

are high enough to have all of these biomes, and many lack Pine

Forest and/or Mixed Conifer Forest, especially in Mexico. However,

all eight of these biomes occur in the Santa Catalina Mountains.

	 The elevational ordering of these biomes, and the strong Madrean

component of most of the Sky Islands, were first formally described

by Forrest Shreve early in the twentieth century (Shreve 1915, 1919,

1922, 1936, 1951).Joseph Marshall’s 1957 study ofthe birds of PineOak Woodlands of the border region also described the stacking of

biotic communities on each island mountain from the Mogollon Rim

to the Sierra Madre, and it was Marshall who defined the “Madrean

Archipelago” asthosemountainswithMexican Pine-OakWoodlands

in the Cordilleran Gap. In 1951, Weldon Heald, studying and living

in the Chiricahua Mountains, coined the evocative and descriptive phrase “Sky Islands” for these ranges. Also in the 1960s, the

pioneering ecological studies of Robert Whittaker, William Niering,

and Charles Lowe described the botany of these ranges in growing

detail. Comprehensive descriptions of Arizona’s Sky Island biomes

can also be found in the forthcoming book A Natural History of the 

Santa Catalina Mountains, Arizona, with an Introduction to the 

Madrean Sky Islands (Moore and Brusca, in press, Arizona-Sonora

Desert Museum Press).

ASAP Surveys in the Santa Catalina 

Mountains

Catalinas Elevational Gradients

	 Situated 140 km north of the United States-Mexico border, the

Santa Catalina Mountains are one of the best-known Sky Islands. To

assess the diversity and distribution of ground-dwelling arthropods

along elevation and environmental gradients, 66 sampling sites were

identified in recognizable biomes along the elevation gradients of the

southern and northern sides of the Santa Catalina Mountains, along

theMt.LemmonHighway (south side),theControlRoad (north side),

and on Mt. Lemmon and Mt. Bigelow (fig. 2). These same elevation

gradients along the same roads were studied botanically by Lowe

(1961), Whittaker and Niering (1964, 1965, 1968a,b, 1975), Niering

and Lowe (1985), and Whittaker and others (1968). Whittaker and

others (1968) established 30 0.1-ha quadrats (20 x 50 m) for their

plant censuses. Our study used 66 belt transects, each 0.02 ha in size

(2x100m)forspecies-X-abundance plant censuses.Neitherthe study

byWhittaker and others(1968) nor ourinitialstudy examined riparian

or “wet canyon” sites; i.e., all transectsites established during the first

year of the project are upland sites. In total we established 28 sites

each on the southern and northern slopes of the Catalina Mountains

and 10 mixed conifer sites on Mt. Lemmon and Mt. Bigelow.

	 While past workers have been in agreement about the over-arching

sense of plant species turnover and plant community change with elevation,they have agreed only partly inwhere to drawlinesseparating

these biomes, and what to call them. Forrest Shreve, in his benchmark

1915 paper,separated the slopes of the Catalina Mountainsinto three

broad zones based on the biogeographic roots of the predominant

plants: Desertscrub, “encinal” (dominated by Madrean species), and

“forest” (dominated by Rocky Mountain/Petran species). Shreve’s

idea was compelling, but it does not work well for all the other Sky

Islands, especially those on the fringes of the Sky Island region

whose plant community affinities are not so clear-cut. It also largely

ignored the grasses and the importance of Grassland habitat on the

Sky Islands. In a detailed series of papers by Whittaker, Niering, and

Lowe, Shreve’s ideas were refined into something very close to what

we use in this paper. Our scheme is a slight alteration of the scheme

presented in Niering and Lowe’s 1985 publication, and it is based

on vegetation analyses of 66 transect sites in the Catalina Mountains

established by the ASAP Project (table 3, fig. 2).

	 In each biome, 100-m long transects were placed farther than 0.25

kmfromthe road, tominimize possible road effects. On the south side

ofthe rangewe establishedfive transectsinDesertscrub (at elevations

of 1045–1172 m), six in Oak-Grassland (at elevations of 1384–1433

m),seven in Pine-OakWoodland (at elevations of 1803–2422m), two

in Chaparral (at elevations of 1923–2052 m), and eight in Pine Forest

(at elevations of 2224–2463 m). On the north side of the range we

established seven transectsindisturbedDesertGrassland (historically

grazed areas at elevations of 1323–1451 m), six in relatively undisturbed Desert Grassland (at elevations of 1330–1645 m), two in Oak

Woodland(at elevationsof1939–2000m),five inPine-OakWoodland

(2032–2149m),five inChaparral(at elevations of 1845–1971m), and

four in Pine Forest (at elevations of 2218–2305 m). Ten transects in

mixed conifer habitat were established in the Mt. Lemmon and Mt.

Bigelow areas (at elevations of 2442–2777 m).

	 The southern slope elevation gradient along the Mt. Lemmon

Highway does not have good Oak Woodland or Grassland; all grass

habitats on the south side were deemed to be Oak-Grassland, a fact

also observed byWhittaker andNiering (1965), although goodDesert

Grassland can be found by hiking west from Molino Basin a few

miles on the Arizona Trail. The northern slope elevation gradient

along the Control Road has no Desertscrub, it begins near the town of

Oracle at 1220 m in elevation, a fact observed by Forrest Shreve and

all subsequent works since the 1920s. However, the northern slope

does have highly disturbed, overgrazed Grassland that has converted

to scrubland (which we sampled) that resembles Desertscrub as

overgrazed Grassland does throughout southern Arizona. The reduction of grasses by livestock and fire suppression, and the subsequent

invasion by woody and shrubby desert plants in the Southwest, has

been known since Aldo Leopold (1924) and a large literature exists

on this subject. The north side also had no Oak-Grassland; it did,

however, have relatively undisturbed Desert Grassland and some

patches of OakWoodland. Of course, it haslong been recognized that

these biomes, or plant communities, intergrade continuously (Whittaker and others 1968; Brown and Lowe 1980), and local variations

in vegetation are due to gradients of slope, soil type, slope aspect,

etc. This was a major focus of Whittaker and his colleagues’ work

in the Sky Islands for many years. A detailed series of papers (Whittaker and Niering 1964, 1965, 1968a,b, 1975; Whittaker and others

1968) explored relationships between plant species and a variety of
150 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Figure 2—TheSantaCatalinaMountainsfromspace showinglocationofmajortopographic features aswell astheMt.LemmonHighwayandtheControlRoad.
USDA Forest Service Proceedings RMRS-P-67. 2013 151

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

(continued)
152 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

(continued)
USDA Forest Service Proceedings RMRS-P-67. 2013 153

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

environmental factors. It is not our goal to repeat those analyses but

rather to accurately describe and classify the botany of our arthropod

transect sites.

	 Transect sites for our 2011 fieldwork in the Catalina Mountains

were categorized according to six environmental variables: elevation,

slope, aspect, biome, as well as ground temperature and humidity.

A Log-Tag HAXO-8 temperature and humidity recorder was placed

2 cm above the soil/litter in the center of each of the 66 transects.

Log-Tags were placed in the field on May 6, 2011 and they recorded

data every 30 min. Average temperature data for each site along the

elevation gradients from May 7, 2011, through September 14, 2011,

is plotted according to elevation and biome (fig. 3). Although we

did not statistically analyze the temperature data, it is apparent that

north- and south-side temperatures (per elevation) were not markedly different, and this is probably partly due to the fact that this was

the hottest time of the year, although differences in slope aspect and

otherfactors probably also contributed to evening out these averages.

Nonetheless,the overall negative correlation between temperature vs.

elevation and biome is clear.

Ground Dwelling Arthropod Surveys

	 We set10pitfalltraps arranged10mapart alongeach100-mtransect

line. Pitfall trap design was adopted from Higgins (2010). Each trap

consists of a heavy Pyrex glass “test tube” (3.2 cm diameter, 25 cm

deep) inserted in a PVC sleeve (3.8 cm in diameter, 28 cm long) that

had been buried in the ground. The opening of the trap is flush with

the soilsurface, and the glasssampling tubes can be exchanged while

leaving the PVC sleeve in place. Traps were charged with propylene

glycol (50% full). A PVC rain shield covers the opening of the trap,

3–4 cm above the ground. When the glass tubes are not in place, the

PVC sleeves are capped to prevent them from filling with dirt or

inadvertently capturing any animals.

	 In 2011 we sampled for 2 weeks in the spring (pre-monsoon,

May1–15)and 2weeksin the late summer(post-monsoon, September

1–15). This project representsthe first effort to extensively and quantitatively sample the arthropod fauna of any of the Sky Islands of the

Madrean Archipelago and, as such, will provide baseline data on the

abundance, diversity, and community structure of this exceptionally

diverse group for future studies.
154 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Figure 3—Average summer temperatures collected 7-May-2011 to 14-Sept-2011 along the transect elevation gradients in the 

Santa Catalina Mountains. Top: A comparison of average temperatures at sites on the South and North sides of the mountain, 

as well as those found at the highest elevations on Mt. Lemmon and Mt. Bigelow. Bottom: A comparison of average temperatures at sites in different plant biomes.

	 Here, we present preliminary species lists of the ants and carabid

beetles based on our first round of pitfall-trap sampling. These data

were supplemented from specimen-level data in the University of

Arizona Insect Collection to develop working lists of the known

species from the Santa Catalina Mountains. We expect these species

lists to grow with future collecting and monitoring efforts.

Ants—The ants, family Formicidae, are possibly the numerically

dominantfamilyofinsects.Antsrepresent10–15%ofthe entire animal

biomassin terrestrial ecosystems(Hölldobler andWilson 1990).Ants

have played a significant role in the evolution of modern terrestrial

biotic communities for at least the last 40–50 million years. More

species of ants occur inArizona than in any other U.S. State (Johnson

1996).While there have been no extensive surveys of the ant fauna of

the Santa Catalina Mountains, we expect their diversity to be similar

to that of other Sky Islands of comparable area and elevation range.

The most well-known Sky Island ant fauna is that of the Chiricahua

Mountains where 187 species have been documented, representing

59% ofthe total known antfauna ofArizona (StephanCover, personal
USDA Forest Service Proceedings RMRS-P-67. 2013 155

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

communication).To date, in the SantaCatalinas, 88 species have been

identifiedthroughourpitfalltraps andtheUniversityofArizona Insect

Collection (table 4).

Ground Beetles—The ground beetles, family Carabidae, comprise

one ofthe largest beetle familieswith atleast 40,000 described species

worldwide.Most ground beetlesrepresent apex predators ofmostsoil

arthropod communities and, thus, play an important ecological role

in almost every terrestrial habitat. Because of this, they have been

important subjects in ecological and climate change studies such as

the long-term climate change study presently being conducted by the

National Ecological Observatory Network (NEON). There are over

2500 described species of Carabidae known from North America.

Based on the holdings of the UAIC, over 300 species occur in the

Sky Island Region of Arizona. Transect samples and UAIC collection records document 69 species of Carabidae occurring in the Santa

Catalina Mountains (table 5).

	 Initialanalysesofpatternsofspeciesdistributionofcarabidsandants

find patterns for the fauna of the Santa Catalina Mountains similar to
156 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

those Halffter(1987)recognized in themountains ofMexico. Species

with temperate ancestral distributions are generally found at higher

elevations(Pine Forest and MixedConifer Forest), while Neotropical

species are generally found at lower elevations (Desertscrub, Desert

Grassland, and Oak Woodland). This pattern is also similar to those

reported in the studies of Ball (1968), Liebherr (1994), and Marshall

and Liebherr (2000) for montane carabids in Mexico. For example,

in the Catalinas, the temperate genus Scaphinotus is represented

by one species, Scaphinotus petersi catalinae Van Dyke, which is

restricted to Pine Forest and Mixed Conifer Forest and the tropical

genus Goniotropis, represented by Goniotropis kuntzeni Bänninger,

which is restricted to Oak-Grassland. The Madrean Archipelago and

the neighboring northern Sierra Madre mountains are the only areas

in the world where members of these two genera, with such disparate

ancestral distributions, can be found within the same mountain range.
USDA Forest Service Proceedings RMRS-P-67. 2013 157

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Plant Surveys

	 Plant surveys were conducted August 5-14, 2011. Surveys were

made for each transect site in two ways. First, the transect line was

walked and every plant recorded within one meter on either side of

the line. This produced a 200-m2 belt transect record of plant species

and abundance. Then, the transect was walked again, this time scanning the broader area outside the 200-m2 area, noting the presence

but not abundances of any plants that might not have been in the belt

transect itself. For practicalreasons,Rocky Mountain ponderosa pine

and Arizona pine were counted together, and no attempt was made

to distinguish between these two species. In the Catalina Mountains,

the Coronado National Forest manages these as a “single species,”

and past workers have considered Arizona pine to be a variety of

ponderosa (P. ponderosa var. arizonica), although this is not current

opinion.

	 Surveys of plantsin the 66 transectsitesin the Catalina Mountains

recorded a total of 316 species: 24 trees, 30woody shrubs, 23 stemand

rosette succulents (including one hybrid agave), 38 grasses, and 201

small shrubs, herbs and annuals. Data are summarized in table 6, and

a complete list of plants-by-biome is given in table 7. For each biome,

there are two species lists, one of “common species” that occurred in

two or more transects and the other of “uncommon species” that occurred in only one transect. The complete plant database is available

from W. Moore. Whittaker and Niering’s (1964) benchmark study of

the southern slopes oftheCatalina Mountainslisted 700 plantspecies

(those above 2743 m being from the southern slopes of the Pinaleño

Mountains), but they did not report these plant occurrences relative

to specific biomes as we do in this preliminary study. Instead, they

reported them relative to a large matrix of environmental variables

and growth forms.Although we do not use that approach here (we are

interested in documenting plantspecies associatedwith our arthropod

transectsites), theASAP project is collecting detailed environmental

data and may undertake such an analysis in the future as relevant to

arthropod species and communities.Also, none ofthe publications by

Whittaker, Niering, or Lowe analyzed plant occurrences statistically

in the Catalina Mountains relative to elevation or biome.

	 After classifying sites according to biome using the categorical

systempreviously described, we used a series of analysis ofsimilarity

(ANOSIM)tests asimplementedinPRIMER5.2.9(Clarke andGorley

2001) to ask if the plant communities in these biomes significantly

differ from one another. We calculated the Bray-Curtis distance

among all possible pairs of sites based on species composition and

abundance ofthe 77 species oftrees, perennialwoody shrubs, and succulentsfound in oursites.We then usedANOSIM to establish overall

differences among plant biomes, followed by a series of pairwise

biome comparisons. To visualize the relationships among sites we

provide the multidimensional scaling (MDS) ordination plot (fig. 4).

ANOSIMrevealed significant overall differencesin plant community

composition among biomes (R = 0.858, P < 0.001). Furthermore,

pairwise comparisonsindicate that allplantbiomessignificantlydiffer

from one another, which suggests that our classification system does

identify sites that are unique with regard to their plant assemblages.

Oak-Woodland sites were excluded from these analyses because of

the low sample size (two sites). To the best of our knowledge, the

ANOSIM analysis of our 66 sites based on plant abundance and species composition is the first statistical analysis of plant distributions

in this range.

Table 6—Biomes sampled in the Santa Catalinas, with numbers of plant species recorded from transects in each.

Biome a

Common

species b

Uncommon

species c Total species

Elevation range 

sampled

Desertscrub (S)

5 transect sites

44 30 74 1045-1172 m

(3428-3845 ft)

Desert Grassland (N)

5 transect sites

61 70 131 1330-1645 m

(4364-5397 ft)

Grazing-Disturbed Grassland (N)

7 transect sites

51 48 99 1323-1451 m

(4340-4760 ft)

Oak-Grassland (S)

6 transect sites

52 36 88 1384-1433 m

(4541-5030 ft)

Oak Woodland (N)

2 transect sites

9 21 30 1939-2000 m

(6362-6562 ft)

Chaparral (N)

5 transect sites

32 22 54 1845-1971 m

(6053-6467 ft)

Chaparral (S)

2 transect sites

10 17 27 1923-2052 m

(6309-6732 ft)

Pine-Oak Woodland (N)

5 transect sites

29 17 46 2032-2149 m

(6667-7051 ft)

Pine-Oak Woodland (S)

7 transect sites

26 24 50 1803-2422 m

(5915-7946 ft)

Pine Forest (N)

4 transect sites

10 13 23 2218-2305 m

(7277-7562 ft)

Pine Forest (S)

8 transect sites

25 20 45 2224-2463 m

(7297-8081 ft)

Mixed Conifer Forest (MTN) 10 

transect sites

24 19 43 2442-2777 m

(8012-9111 ft)

a

 N = northern slopes of range (along Control Road). S = southern slopes of range (along Mt. Lemmon 

 Highway). MTN = Mt. Lemmon and Mt. Bigelow sites. 

b

 “Common Species” are species found in two or more transects in the given biome. 

c

 “Uncommon Species” are species recorded from only a single transect in the given biome. 
158 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Cont.
USDA Forest Service Proceedings RMRS-P-67. 2013 159

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Table 7—Continued

Cont.
160 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Table 7—Continued

Cont.
USDA Forest Service Proceedings RMRS-P-67. 2013 161

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Table 7—Continued

Cont.
162 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Table 7—Continued

Cont.
USDA Forest Service Proceedings RMRS-P-67. 2013 163

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Table 7—Continued

	 Plant diversity data are summarized in tables 6 and 7. The highest

plant diversity occurred in our Desert Grassland sites on the north

side of the Catalina Mountains, where the relatively undisturbed

transectsites had a total of 131 species and the grazing-disturbed sites

had a total of 99 species. This was followed by the southern slope

Oak-Grassland sites with 88 species, and the Desertscrub sites with

74 species. Pine-Oak Woodland sites had 46 plant species (northern

slopes) and 50 species (southern slopes). The lowest diversity sites

were Pine Forest of the northern slopes (23 species) and Chaparral

on the southern slopes (27 species). This is the same biome diversity

trend noted by Whittaker and Niering (1965) and other workers over

the years. A number of our plant records are new species to the Catalina Mountains, or significant elevation extensions, and these will be

discussed in future papers.

Desertscrub—Because the base of the Catalina Mountains on its

northern side is ~1220 m, there is no Desertscrub. Instead, the biome

at the base (the start of the Control Road, in the town of Oracle) is

Grassland. However, a well-developed Arizona Upland Desertscrub

is at the base of the mountain on the south side, and it contains all

of the signature plants of this Sonoran Desert subprovince (table 7).

Whittaker and Niering (1965) called this the “Sonoran Desert of

Mountain Slopes” biome, even though Forrest Shreve had coined

the well-accepted name Arizona Upland in 1951 for this subprovince of the Sonoran Desert. Three of our Desertscrub sites are near

Babad Do’ag Viewpoint, along the Babad Do’ag Trail. Because of

its elevation (1133–1160 m) and proximity to the opening of Molino

Canyon, the Babad Do’ag sites are wetter than the other Desertscrub

site (1045m), and thus velvetmesquite (Prosopis velutina) and desert
164 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

hackberry (Celtis pallida) are present(and palo verde/Parkinsonia sp.

and saguaro/Carnegiea gigantea are fewer in number); also shindagger agave (Agave schottii), turpentinebush (Ericameria laricifolia),

limberbush (Jatropha cardiophylla),sprawling prickly pear(Opuntia 

phaeacantha var. major), andmany wildflowers. For example, purple

ground cherry (Physalis crassifolia), desert windmills (Allionia incarnata) and spike moss (Selaginella arizonica) occur at this wetter

site,indicative ofits proximity to the transition intoGrassland habitat.

Although none of our transects had invasive buffelgrass(Pennisetum 

ciliare), large patches of it occur acrossthe Desertscrub landscape on

the southern slopes oftheCatalina Mountains and in theBabad Do’ag

Trail area large patches of invasive Lehmann lovegrass (Eragrostis 

lehmanniana) also are found. Desertscrub habitat extends to ~1177

m on the southern slopes of the Catalina Mountains along the Mt.

Lemmon Highway.

Undisturbed Savanna-like Habitats: Desert Grassland and OakGrassland—About halfthe plantspeciesfound in these two savannalike biomes are shared, so we discuss them together. There are no

large swaths of Desert Grassland along the Mt. Lemmon Highway on

the south side of range, and there Desertscrub transitions quickly into

Oak-Grassland dominated byEmory oak,Arizonawhite oak, and four

grasses - side-oats grama (Bouteloua curtipendula), cane beardgrass

(Bothriochloa barbinodis),Arizonapanicgrass(Urochloa arizonica),

andtheelevationallywide-rangingbullgrass(Muhlenbergia emersleyi)

(a total of 14 grasses were recorded in our Oak-Grassland transects).

Other abundant plants of our Oak-Grassland sites are pointleaf manzanita (Arctostaphylos pungens),shindagger agave,sotol(Dasylirion 

wheeleri), beargrass (Nolina microcarpa), mountain yucca (Yucca 

madrensis), scarlet creeper (Ipomoea cristulata), Amaranth (Amaranthus nr. palmeri), and three species of Portulaca. Although our

Grassland and Oak-Grassland sitesshare many species, each also has

many unique components(table 7).The transition between Grassland

and Oak-Grassland biomesismost easily seen in the turnover of grass

species along elevational gradients and the increasing abundance of

oaks with higher elevations. Although Arizona white oak (Quercus 

arizonica) and manzanita make their first appearance in these grassy

habitats,theseareelevationallybroad-rangingplantsthatarealsofound

in Chaparral, Oak Woodland, and Pine-Oak Woodland. In contrast

to the southern slopes, the north side of the Catalina Mountains has

accessible, expansive, rolling hills of Desert Grassland but no welldevelopedOak-Grassland (along theControlRoad).These north-side

Desert Grassland sites harbored 131 plant species, making this the

most botanically diverse biome in our study, including 22 grass species, although 4 species dominated in abundance: side-oats grama,

Arizona panic grass, Mexican panic grass (Panicum hirticaule), and

the invasive Lehmann lovegrass. In addition to these grasses, velvet

mesquite, whitethorn acacia (Acacia constricta), wait-a-minute bush

(Mimosa aculeaticarpa), fairy duster (Calliandra eriophylla) were

all present in high numbers, indicative of some history of grazing

disturbance. On Grassland sites with strong limestone presence, the

most abundant plants included a half-dozen typical Desertscrub species, with ocotillo (Fouquieria splendens) dominating. Ground cover

Figure 4—MDS ordination of sites based on Bray-Curtis similarity values. Sites were classified according to the composition and abundance of 77 species of trees, perennial woody shrubs, and succulents found in the transect sites. Plant biome 

designations for each site were determined a-priori using the criteria outlined in this paper. Note that the MDS ordination 

shown in this figure is a 2-dimensional compression of a multi-dimensional clustering of the sites, so graphical representation is not perfect. However, the stress value of 0.07 indicates that information in the 2-dimensional figure does a good job 

of describing patterns in the multi-dimensional space.
USDA Forest Service Proceedings RMRS-P-67. 2013 165

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

by grasses was high in all Desert Grassland transects, often 90%, and

sometimes dominated by the invasive Lehmann’s lovegrass.

Grazing-Disturbed Grassland—On the north side of the Catalina

Mountains, most Grassland habitats have been converted to “scrubland” by over a century of grazing, giving the landscape a desert-like

appearance. Seven transects were established in this highly disturbed

DesertGrasslandnearthebaseofthemountainalongtheControlRoad,

from 1337 to 1451 m. All of these sites showed evidence of decades

of heavy livestock traffic, with a great deal of disturbed, bare dirt and

disrupted soil and, in some areas, erosion down to bedrock. These

transects had 99 plant species that were a mix of Desert Grassland

andDesertscrub species characteristic oflong-overgrazedGrasslands

of southeastern Arizona. Typical invasive native Desertscrub plants

included velvet mesquite, whitethorn acacia, catclaw acacia (Acacia 

greggii), fishhook barrel cactus(Ferocactus wislizeni),slim ragweed

(Ambrosia confertiflora), and fairy duster, and the abundance and

biomass of these species generally exceeded that of the Grassland

species. The large number of grass species (19) in these disturbed

sites might be due, in part, to the history of livestock grazing and

imported forage. This was the only site where we found red brome

(Bromus rubens), an exotic invasive grass.

Oak Woodland—OakWoodlandisgenerallycharacterizedbystands

of oaks, with junipers and pinyons. This biome reaches its maximum

development in the Sierra Madre of Mexico, and in general, more

northern Sky Islands have less Oak Woodland than more southern

ranges. In the Catalina Mountains, well-developed Oak Woodland

does not occur along the Mt. Lemmon Highway, on the south side

of the mountains. However, on the north side of the Catalina Mountains, patches of Oak Woodland do occur along the Control Road,

and in our two transect sites in this habitat there was a total of 30

plant species (tables 6 and 7). Seven of the nine signature species

of this biome listed by Whittaker and Niering (1968b) occurred in

our site transects. These woodlands are dense with foliage, the large

dominants being Arizona white oak, silverleaf oak, Chihuahua pine

(Pinus chihuahuana), alligator juniper (Juniperus deppeana), and

Fendler buckbrush (Ceonothus fendleri). Bullgrass covers much of

ground between the trees. In the Catalinas, the elevationally broadly

occurring alligator juniper, Fendler buckbrush, and adder’s tongue

make their first appearances in Oak Woodlands and are then found

all the way into Pine Forest habitat and in the case of adder’s tongue

into the Mixed Conifer Forest.

Chaparral—Interior Chaparral occurs on both the north and south

sides of the Catalina Mountains, but it is more extensively developed

along theControlRoad on the northern slopes. OurChaparralsites on

the south side of the range had 27 plantspecies, whereasthe northern

sites had 54. Even though the plant diversity on the north-side sites

was greater, neither border pinyon norsotol were found in those transect sites. The north-side Chaparral transects had abundant alligator

juniper, Arizona white oak, Emory oak (Quercus emoryi), Fendler

buckbrush, mountain mahogany (Cercocarpus montanus), silktassle

bush (Garrya wrightii), and mountain yucca—all missing from the

more depauperate south-side sites (tables 6 and 7). This is not to say

these typical Chaparral plants do not occur on the southern slopes

of the Catalina Mountains; they do, but just not in our transect sites.

Virtually all of the Chaparral along the Mt. Lemmon Highway was

burned in the 2003 Aspen fire, including our two south-side transect

sites(nearManzanitaVista andWindy PointVista), and it islikely this

was at least partly the reason these sites were depauperate relative to

those on the north side of the mountains (none of our five north-side

Chaparralsites burned in theAspenFire).These south-side plant communities are only now beginning to show strong recovery. Chaparral

transects on both sides of the range were dominated by pointleaf

manzanita, silverleaf oak, golden-flower agave (Agave chrysantha),

beargrass, and spidergrass (Aristida ternipes var. ternipes). Higher

plant diversity in the north-slope sites is likely also due to the greater

variety of soil types there, including limestone soils. Whittaker and

Niering (1965) and others have shown that limestone soilsshift plant

biomesupwardonSkyIslandmountains,suchthat at elevationswhere

Grassland would normally occur one finds Desertscrub plants (with

a strong presence of limestone-loving species such as ocotillo). This

shift presumably occurs because limestone is porous and does not

retain moisture, creating a more xeric soil condition that Desertscrub

plants are adapted to. Several of ourChaparralsiteswere on limestone

soils and these showed the expected up-elevation shift in Desertscrub

plants.

Pine-Oak Woodlands—Our Pine-Oak Woodland transects had a

total of 50 species(south slopes) and 46 species(north slopes).Abundant on both slopes were alligator juniper, Chihuahua pine, Rocky

Mountain ponderosa pine/Arizona pine,RockyMountainDouglas-fir

(Pseudotsuga menziesii var. glauca),silverleafoak,Fendlerbuckbrush,

mountain yucca, cudweed (Pseudognaphalium sp.), New Mexico

groundsel (Packera neomexicana), goldenrod (Solidago wrightii),

and bullgrass. It is here that Rocky Mountain Douglas-fir and Rocky

Mountain ponderosa pine/Arizona pinemake their first appearance in

the Catalina Mountains, to continue all the way into Mixed Conifer

Forest. Silverleaf oak, with its broad elevational range, occurs from

Pine-Oak Woodland and Chaparral well into Pine Forest habitat. All

six of Whittaker and Niering’s (1968b) signature plant species of

Pine-Oak Woodland occurred in our transect sites.

Pine Forest—OurPineForesttransectsites had a total of 45 species

(south slopes) and 23 species(north slopes).Abundant on both slopes

were Rocky Mountain ponderosa pine/Arizona pine, southwestern

white pine (Pinus strobiformis), Rocky Mountain Douglas-fir,silverleafoak,Fendlerbuckbrush,brickellbush(Brickelliasp.),butterweed,

goldenrod, pineland dwarf mistletoe (Arceuthobium vaginatum),

cudweed,brackenfern(Pteridum aquilinum), andbullgrass.Although

white fir occurred in some of the south-side Pine Forest transects,

this tree is more typical of the Mixed Conifer Forest and did not

occur below 2255 m in our transects. Rocky Mountain ponderosa

pine/Arizona pine was dominant conifers in both the Pine Forest

and Mixed Conifer biomes. Whittaker and Niering (1968b) noted

five signature plant species of Pine Forest, although two were lowerelevation transitionals to Pine-Oak Forest/Woodland, and all but one

of these (their transitional netleaf oak/Quercus rugosa) also occurred

in our Pine Forest transects. It isin the upper Pine-OakWoodland and

Pine Forest biomes that ponderosa Pine Forests dominate the visual

landscape oftheCatalina Mountains andmost ofArizona’s other high

Sky Islands.

Mixed Conifer Forest—Transectsitesin our Mixed Conifer Forest

sites, on Mt. Lemmon and Mt. Bigelow, at 2442–2777 m had 43 species (tables 6 and 7). Dominants, in terms of numbers of individuals,

were Rocky Mountain ponderosa pine/Arizona pine, white fir (Abies 

concolor), southwestern white pine, Rocky Mountain Douglas-fir,

pineywoods geranium (Geranium caespitosum), Oxalis sp. (an

unidentified Oxalidaceae), mountain parsley (Pseudocymopterus 

montanus), braken fern, meadow rue (Thalictrum fendleri), and

Viola spp (an unidentified violet). Corkbark fir, which is rare south

of the Mogollon Rim, occurred in only 2 of our 10 Mixed Conifer

Forest transects. There are no naturally occurring spruces (Picea)

in the Santa Catalina Mountains, and probably also no limber pine

(Pinus flexilis), a more northern species, despite some old (dubious)

records from the Catalina Mountains. Whittaker and Niering (1964)
166 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

andWhittaker and others(1968)subdivided theMixedConiferForest

into two elevational zones, “montane fir forest” and “subalPine Forest” (above 2440 m/8000 ft). They distinguished their “montane fir

forest” biome by six species that occurred on our sampling transects.

Summary

	 TheArizonaSky IslandArthropodProject(ASAP)waslaunched in

2011.Parametersandover-archinggoalsoftheproject,andpreliminary

results of year one, are presented here. We define the boundaries of

the Madrean Sky Island Region somewhat more broadly than they

have been in the past, with less of an emphasis on the Madrean flora

components and more of an emphasis on their location as isolated

Cordilleran Gap ranges. The Mexican Sky Islands are still not well

known and adjustments regarding the range and extent of the Sky

Island Region will no doubt be fine tuned in the future. We point out

the important difference between “mountain islands” vs. “habitat

islands.” We present the first statistical analysis of plant species and

biomes in the Santa Catalina Mountains, and use this to preliminarily establish the biome names and boundaries that we will use in our

arthropod analyses. Year one ant and ground beetle (Carabidae) data

for the Catalinas are presented; both groups are high in diversity (88

ant species, 69 ground beetle species) and show a trend of species

affiliation to plant biomes.

Acknowledgments

	 This paper would not have been possible without the assistance of

manypeople,whogenerouslyhelpedinthefieldandwiththe research,

and who put up with our endless badgering with questions. Grateful thanks go to David Bertelsen, Mark Dimmitt, George Ferguson,

George Montgomery, and Julie Wiens for their botanical tutorials.

Thanks to Aaron Flesch for information on Sonoran Sky Islands and

for his review of the maps. Our thanks are also extended to Steven

Cover for his help identifying the ants in this study. This work would

not have been possible without the assistance of many undergraduate students including Rebecca Compoy, Kayla Jones, Hee Ju, Chris

McGinnis, Gabe Oropeza, Chelsea Powers, Shawna Rodgers, Carol

Tepper, and Eryn Wouri. Special thanks go to Emmanuel Bernal,

Reilly McManus, Jeff Henkel, Ryan McInroy, Garrett Hughes, Paul

Marek,Jason Schaller, and JamesRobertson fortheir help setting and

harvesting pitfall traps and sorting pitfall trap samples. Our work in

the Catalinasis facilitated by U.S. Forest Service Special Use Permit

# SAN0287.ThankstoLindaBrewer andGoggyDavidowitz fortheir

careful reviews of this manuscript that greatly improved the quality

ofthis paper.The University ofArizona’s Department of Entomology

and Center for Insect Science, as well as National Geographic Society (8964-11) and the National Science Foundation (EF - 1206382)

provided funding for this project.

Authors Note

This paper represents the combination of four presentations made by

the authors in a Session on Biodiversity of Arthropods at the conference. The titles of these presentations were “TheArizona Sky Island

ArthropodProject(ASAP):Systematics,Biogeography,Ecology, and

Population Genetics of Ground-Dwelling Arthropods,” “Do Plant

BiomesAlongtheElevationGradientintheSantaCatalinaMountains

Harbor Unique Arthropod Communities?” “Diversity and Species

Composition of Ant Assemblages in the Santa Catalina Mountains,”

and “Hidden Biodiversity of Sky Island Arthropods.”

References

Ball, G.E. 1968. Barriers and southward dispersal of the Holarctic boreomontane element of the family Carabidae in the mountains of Mexico.

Anales de la Escuela Nacional de Ciencias Biologicas 17:91-112.

Barber, P.H. 1999a. Patterns of gene flow and population genetic structure in

the canyontreefrog,Hyla arenicolor(Cope).MolecularEcology.8:563-576.

Barber, P.H. 1999b. Phylogeography of the canyon treefrog, Hyla arenicolor

(Cope) based on mitochondrial DNA sequence data. Molecular Ecology.

8:547-562.

Bardgett, R.D., C. Freeman, and N.J. Ostle. 2008. Microbial contributions to

climate change through carbon cycle feedbacks. ISME Journal. 2:805-814.

Behan-Pelletier, V., and G. Newton. 1999. Linking soil biodiversity and

ecosystem function—the taxonomic dilemma. BioScience. 49:149-153.

Betancourt, J., T.R. Van Devender, and P.S. Martin (eds.). 1990. Packrat

Middens: The Last 40,000 Years of Biotic Change. University Arizona

Press, Tucson, AZ.

Boyd, A.E. 2002. Morphological analysis of Sky Island populations of

Macromeria viridiflora (Boraginaceae). Systematic Botany. 27:116-126.

Bowers, J.E., and S.P. McLaughlin. 1996. Flora of the Huachuca Mountains,

a botanically rich and historically significant Sky Island inCochiseCounty,

Arizona.Journal of theArizona-NevadaAcademy of Sciences. 29:66-107.

Brown, J.H. 1971. Mammals on mountaintops: non-equilibrium insular biogeography. The American Naturalist. 105:467-478.

Brown,D.E.(ed.). 1994.BioticCommunities.SouthwesternUnitedStates and

Northwestern Mexico. University of Utah Press, Salt Lake City.

Brown, D.E. and C.H. Lowe. 1980. Biotic communities of the Southwest.

GTR-RM-78. U.S. Department ofAgriculture, U.S. Forest Service, Rocky

Mountain Forest and Range Experiment Station, Fort Collins, CO.

Brown,D.E.,C.H.Lowe, andC.P.Pase. 1979.Adigitized classification system

for the biotic communities of NorthAmerica, with community (series) and

association examples for the Southwest. Journal of the Arizona-Nevada

Academy of Science. 14:1-16.

Clarke R. and Gorley R. N. 2001. PRIMER v5: User manual/tutorial. Plymouth: PRIMER-E.

CLIMAS (ClimateAssessment for the Southwest) Project, University of Arizona. http://www.climas.arizona.edu/projects/southwest-climate-outlook

(accessed March 2012).

Coope, G.R. 1995. The effects of Quaternary climatic changes in insect

populations: Lessons from the past. Pp. 30-49, in R. Harrington and N. E.

Stork (eds.),Insectsin aChanging Environment. Academic Press, London.

Coviella, C.E., and J.T. Trumble. 1999. Effects of elevated carbon dioxide on

insect plant interactions. Conservation Biology. 13:700-712.

Davis, R., and D.E. Brown. 1988. Documentation of the transplanting of

Abert’s squirrels. Southwestern Naturalist. 33:490-492.

Dimmitt, M.A.,R.C.Brusca, K. Krebbs,C.L. McIntyre,J.Viers,C. Filippone,

and A. Berry. 2011. Natural Resource Condition Assessment: Chiricahua

National Monument, Coronado National Memorial, and Fort Bowie National Historic Site.Arizona-Sonora Desert Museumand Sonoran Institute,

Tucson, AZ.

Dominguez, F.,J. Canon, and J. Valdes. 2010. IPCC-AR4 climate simulations

for the Southwestern US: the importance of future ENSO projections.

Climatic Change. 99:499-514.

Downie, D.A. 2004. Phylogeography in a galling insect, grape phylloxera,

Daktulosphaira vitifoliae (Phylloxeridae) in the fragmented habitat of the

Southwest USA. Journal of Biogeography. 31:1759-1768.

Flesch, A.D., and L.A. Hahn. 2005. Distribution of birds and plants at the

western and southern edges ofthe Madrean Sky Islandsin Sonora, Mexico.

Pp. 80-87 in Gottfried, G.J., B.S. Gebow, L.G. Eskew, and C.B Edminster.

Connectingmountain islands and desertseas:biodiversityandmanagement

of the Madrean Archipelago II. Proceedings RMRS-P-36. Fort Collins,

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station. 631 p.
USDA Forest Service Proceedings RMRS-P-67. 2013 167

Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . . Moore and others

Graham, R.W., and E.C. Grimm. 1990. Effects of global climate change on

the patterns of terrestrial biological communities. Trends in Ecology and

Evolution. 5:289-292.

Gutzler,D.S., andT.O.Robbins.2010.Climatevariabilityandprojectedchange

in the western United States: regional downscaling and drought statistics.

Climate Dynamics. 34(7-8):10.1007/s00382-010-0838-7.

Halffter, G. 1987. Biogeography of the montane entomofauna of Mexico and

Central America. Annual Reviews of Entomology. 32:95-114.

Heald, W.F. 1951. Sky Islands of Arizona. Natural History. 60:56-63, 95-96.

Higgins, J.W. 2010. Ground dwelling arthropod responses to successional

endpoints: burned versus old growth pinyon-juniper woodlands(Master’s

thesis). Northern Arizona University, Flagstaff, AZ.

Hölldobler, B., and E.O. Wilson. 1990. The Ants. Harvard University Press,

Cambridge, MA.

Hughes, L. 2003. Climate change and Australia: trends, projections and

impacts. Australian Ecology. 28:423-443.

IPPC (Intergovernmental Panel on Climate Change). 2001. Climate change

2001 synthesis report: Summary for policy makers. Intergovernmental

Panel on Climate Change. Geneva, Switzerland.

IPCC (Intergovernmental Panel on Climate Change). 2007. Climate change

2007: The Physical Science Basis. Cambridge University Press, Cambridge, UK.

Jablonski, D., and J.J. Sepkoski. 1996. Paleobiology, community ecology, and

scales of ecological pattern. Ecology. 77:1367-1378.

Johnson, R.A. 1996. Arizona ants. Arizona Wildlife Views. June, pp. 2-5.

Kupfer,J.A.,J.Balmat, andJ.L.Smith.2005.Shiftsinthepotentialdistribution

of Sky Island plant communities in response to climate change. Proceedings RMRS-P-36. U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. Fort Collins, CO.

Leopold,A.S. 1924. Grass, brush, timber, and fire in southernArizona.Journal

of Forestry. 22:1-10.

Liebherr, J.K. 1994. Biogeographic patterns of montane Mexican and

Central American Carabidae (Coleoptera). The Canadian Entomologist.

126:841-860.

Linhart, Y.B., and A.C. Permoli. 1994. Genetic variation in central and disjunct populations of Lilium parryi. Canadian Journal of Botany. 72:79-85.

Lowe,C.H.1961.Biotic communitiesinthe sub-Mogollon regionofthe inland

Southwest. Journal Arizona Academy Science. 42(2):40-49.

Maddison, W.P., and M.M. McMahon. 2000. Divergence and reticulation

among montane populations of the jumping spider Habronattus pugillis

Griswold. Systematic Biology. 49:400-421.

Marshall,C.J., andJ.K.Liebherr. 2000.Cladistic biogeography oftheMexican

transition zone. Journal of Biogeography. 27:203-216.

Marshall, J.T., Jr. 1957. Birds of the Pine-Oak Woodland in southern Arizona

and adjacent Mexico. Pacific Coast Avifauna. 32:1-125.

Martin, P.S. 1963. The Last 10,000 Years: a Fossil Pollen Record of the

American Southwest. University of Arizona Press, Tucson.

Masta, S. 2000. Phylogeography of the jumping spider Habronatus pugillis

(Araneae: Salticidae): recent vicariance of Sky Island populations? Evolution. 54:1699-1711.

McCord,R.D. 1995. Phylogeny and biogeography ofthe land snail, Sonorella,

in theMadreanArchipelago. Pp. 317-323 in,L. F. DeBano, P. F. Ffolliott,A.

Ortega-Rubio, G. Gottfried, R.H. Hamre, C.B. Edminster (Tech. Coord.).

Biodiversity andmanagement ofthe MadreanArchipelago: the Sky Islands

of Southwestern United States and Northwestern Mexico; 1994, Tucson,

AZ. Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture,

U.S. Forest Service, Rocky Mountain Research Station. Fort Collins, CO.

McCormack,J.E.,B.S.Bowen, andT.B.Smith.2008.Integratingpaleoecology

and genetics of bird populations in two Sky Island archipelagos. BioMed

Central Biology. 6:28.

McLaughlin, S.P. 1986.Afloristic analysis of the southwestern United States.

Great Basin Naturalist. 46:46-65.

McLaughlin, S.P. 1995.An overview ofthe flora ofthe Sky Islands, Southeastern Arizona: diversity, affinities and insularity. Pp. 60-70 in DeBano, L.F.,

P.F. Ffolliott, A. Ortega-Rubio, G. Gottfried, R.H. Hamre, C.B. Edminster

(Tech.Coord.).Biodiversity andmanagement oftheMadreanArchipelago:

the Sky Islands of Southwestern United States and Northwestern Mexico;

1994, Tucson, AZ. Gen. Tech. Rep. RM-GTR-264. U.S. Department of

Agriculture, U.S. Forest Service, Rocky Mountain Research Station. Fort

Collins, CO.

Moore, W., and R.C. Brusca. [In press]. A Natural History of the Santa Catalina Mountains,Arizona, with an Introduction to the Madrean Sky Islands. 

Arizona-Sonora Desert Museum Press, Tucson, AZ.

Niering, W.A., and C.H. Lowe. 1985. Vegetation of the Santa Catalina Mountains: community types and dynamics. Pp. 159-184 in, R. K. Peet (ed.),

Plant Community Ecology: papers in Honor of Robert H. Whittaker. W.

Junk Publishers/Kluwer-Academic Publishing, Boston, MA.

Ober, K.A., B. Matthews, A. Ferrieri, and S. Kuhn. 2011. The evolution and

age of populations of Scaphinotus petersiRoeschke onArizona Sky Islands

(Coleoptera, Carabidae, Cychrini). Zookeys. 147:183-197.

Overpeck,J.T., andB. Udall. 2010. Dry times ahead. Science. 328:1642-1643.

Powers, J.S., R.A. Montgomery, E.C. Adair, [and others]. 2009. Decomposition in tropical forests: a pan-tropical study of the effects of litter type,

litter placement and mesofaunal exclusion across a precipitation gradient.

Journal of Ecology. 97:801-811.

Shreve, F. 1915. The vegetation of a desert mountain range as conditioned by

climatic factors.Carnegie Institute ofWashington, Publication 217, 112 pp.

Shreve, F. 1919. A comparison of the vegetational features of two desert

mountain ranges. Plant World. 22:291-307.

Shreve, F. 1922. Conditionsindirectly affecting vertical distribution on desert

mountains. Ecology. 3: 269-274.

Shreve, F. 1936.The transition fromdesert toChaparral.Madroño. 3:257-264.

Shreve, F. 1951. Vegetation of the Sonoran Desert. Carnegie Institute of

Washington, Pub. 591: 1-129.

Slentz, S., A.E. Boyd, and L.A. McDade. 1999. Patterns of morphological

differentiation among Madrean Sky Island populations of Castilleja austromontana (Scrophulariaceae). Madroño. 46:100-111.

Smith, C.I., and B.D. Farrell. 2005a. Phylogeography of the longhorn cactus

beetle Moneilema appressum LeConte (Coleoptera: Cerambycidae): was

the differentiation oftheMadreanSky Islands driven byPleistocene climate

changes? Molecular Ecology. 14:3049-3065.

Smith,C.I., andB.D.Farrell.2005b.Rangeexpansionsintheflightlesslonghorn

cactus beetles, Moneilema gigas and Moneilema armatum, in response to

Pleistocene climate changes. Molecular Ecology. 14:1025-1044.

Sullivan, R.M. 1994. Micro-evolutionary differentiation and biogeographic

structure among coniferous forest populations of the Mexican woodrat

(Neotoma mexicana) in the American Southwest: a test of the vicariance

hypothesis. Journal of Biogeography. 21:369-389.

Tennessen, J.A., and K.R. Zamudio. 2008. Genetic differentiation among

mountain island populations of the striped plateau lizard, Sceloporus 

virgatus (Squamata: Phrynosomatidae). Copeia. 3:558-564.

VanDevender,T.R.2002.EnvironmentalhistoryoftheSonoranDesert.In,T.H.

FlemingandA.Valiente-Banuet(eds.),Columnar cacti andtheirmutualists:

evolution, ecology, andconservation. UniversityArizonaPress,Tucson,AZ.

Van Devender, T.R., and W.G. Spaulding. 1990. Development of vegetation

and climate in southwestern United States. Science. 204:701-710.

Voight, W., J. Perner, and A.J. Davis. 2003. Trophic levels are differentially

sensitive to climate. Ecology. 84:2444-2454.

Warshall,P.1995.TheMadreanSkyIslandArchipelago:Aplanetaryoverview.

Pp.6–18in,DeBano,L.F.,P.F.Ffolliott,A.Ortega-Rubio,G.Gottfried,R.H.

Hamre,C.B.Edminster(Tech.Coord.).Biodiversityandmanagementofthe

Madrean Archipelago: the Sky Islands of Southwestern United States and

Northwestern Mexico; 1994, Tucson, AZ. Gen. Tech. Rep. RM-GTR-264.

U.S. Department of Agriculture, U.S. Forest Service, Rocky Mountain

Research Station. Fort Collins, CO.

Watling, J.I., and M.A. Donnelly. 2006. Fragments as islands: a synthesis of

faunalresponsestohabitatpatchiness.ConservationBiology.20:1016-1025.

Whittaker, R.H., S.W. Buol, W.A. Niering, and Y.H. Havens. 1968. A soil

and vegetation pattern in the Santa Catalina Mountains, Arizona. Soil

Science. 105:440-450.

Whittaker, R.H., and W.A. Niering. 1964. Vegetation of the Santa Catalina

Mountains,Arizona.I.Ecological classification and distribution ofspecies.

Journal of the Arizona Academy of Science. 3:9-34.

Whittaker, R.H., and W.A. Niering. 1965. Vegetation of the Santa Catalina

Mountains, Arizona. A gradient analysis of the south slope. Ecology.

46:429-452.
168 USDA Forest Service Proceedings RMRS-P-67. 2013

Moore and others Introduction to the Arizona Sky Island Arthropod Project (ASAP): . . .

Whittaker, R.H., and W.A. Niering. 1968a. Vegetation of the Santa Catalina

Mountains, Arizona. III. Species distribution and floristic relations on the

north slope. Journal of the Arizona Academy of Science. 5:3-21.

Whittaker, R.H., and W.A. Niering. 1968b. Vegetation of the Santa Catalina

Mountains, Arizona. IV. Limestone and acid soils. Journal of Ecology.

56:523-544.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Whittaker, R.H., and W.A. Niering. 1975. Vegetation of the Santa Catalina

Mountains,Arizona.V.Biomass, production, and diversity along the elevation gradient. Ecology. 56:771-790.

Yang, X., and J. Chen. 2009. Plant litter quality influences the contribution of

soil fauna to litter decomposition in humid tropical forests, southwestern

China. Soil Biology and Biochemistry. 41:910-918.
USDA Forest Service Proceedings RMRS-P-67. 2013 169

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Preliminary Assessment of the Moth (Lepidoptera: 

Heterocera) Fauna of Rincon de Guadalupe, Sierra de 

Bacadéhuachi, Sonora, Mexico

John D. Palting

Department of Entomology, University of Arizona, Tucson, Arizona

Abstract—The Sierra de Bacadéhuachi is a poorly sampled extension of the Sierra Madre Occidental (SMO) 

located in east-central Sonora near the town of Bacadéhuachi. Sampling of moths using mercury vapor and 

ultraviolet lights occurred in summer and fall 2011, and spring 2012 at Rincón de Guadalupe, located in 

pine-oak forest at 1680 m elevation. Approximately 400 taxa of moths were identified from this locality.

Species occurrences demonstrate connections to the famously diverse Yécora area, to the higher, more mesic SMO in Chihuahua, as well as the Rocky Mountains to the north. Several unidentified and presumably 

new moth species from Rincón de Guadalupe await further study and description. Two species previously 

known only from the type specimens (destroyed in WWII) were rediscovered at Rincón de Guadalupe. A 

complete list of the moths identified from Rincon de Guadalupe is available as a Research Species List in 

the Madrean Archipelago Biodiversity Assessment (MABA)/Southwest Environmental Information Network 

(SEINet) online database (Madrean.org).

Introduction

	 The Sierra de Bacadéhuachi is considered the westernmost massif of the Sierra Madre Occidental (SMO), and is located east of the

Pueblo deBacadéhuachi,Municipio deBacadéhuachi,Sonora.While

technically not a Sky Island, the range represents the northernmost

extension of the SMO and serves as a conduit for SMO species to

move north into the Sky Island region.Thus,speciesrecordsfromthis

area, of which there are historically few, are of interest to biologists

working in the Sky Island region. The range is located only 34 km

west of the border with Chihuahua, and 165 km south of the Arizona

border. The Yécora plateau of the SMO is approximately 120 km to

the SE. This area has historic significance as the location of Rincón

deGuadalupe (RG), a rustic enclave of buildings constructed between

1920 -1940 in the pine-oak forest on property owned by the Catholic

Diocese of Sonora. The main facility located in Arroyo Campo Los

Padres (29°50’40”N, 108°58’37”W, 1680 m elevation) was used as

a base of operations for the field activities reported herein.

	 TheSierradeBacadéhuachivegetationrangesfromlowlandfoothills

thornscrub up through desert grassland to oak woodland and finally

pine-oak forest. The area immediately around RG is dominated by

pine and oak, and bisected by a perennial stream. Eleven species of

Quercus and five species of Pinus are recorded from the area along

with Cupressus arizonica, Fraxinus velutina, and Platanus wrightii.

The most speciose families in the preliminary flora of the Sierra de

Bacadéhuachi are Asteraceae (54 taxa), Fabaceae (47 taxa), and

Poaceae (36 taxa) (Van Devender and others, Preliminary Flora of 

the Sierra Bacadéhuachi, Sonora, Mexico, this volume).

Methods

	 A combination mercury vapor/ultraviolet light or ultraviolet light

alone was used to attract the majority of moth species collected. A

few exceptions, such as Coloradia prchali, were netted during the

day. The light stations were run from dusk until dawn in the summer

and fall when the nights remained warm, and until approximately

1AM in the spring after which it became too chilly for moths to

fly. Sampling was conducted throughout the night and specimens

were spread on site and dried in an oven to ensure as many different

“cryptic” taxa as possible were recorded and to get representative

examples of sexually dimorphic species for identification. In addition, several photographers documented diversity at the lights with

photo vouchers of living moths, all of which are available records in

the Madrean Archipelago Biodiversity Assessment (MABA) database

(Madrean.org).

Results

	 About 400 species of moths were identified from all three sampling expeditions to Rincón de Guadalupe. Given that this collection

represents only three periods of activity and a total of nine collecting

nightsin a single locality of the Sierra de Bacadéhuachi, the diversity

of this area appears to be exceptionally rich. The total moth diversity

for all areas and all months known from the Huachuca Mountains 170

km to the north is 1500 species— 900 “macrolepidoptera” (Powell

and Opler 2009) and 600 “microlepidoptera.” This relatively well

surveyedarea (316km2)isbiologicallydiversewithvegetationranging
170 USDA Forest Service Proceedings RMRS-P-67. 2013

Palting Preliminary Assessment of the Moth (Lepidoptera: Heterocera) Fauna of Rincon de Guadalupe, . . .

from desert grassland up to pine-oak forest in an elevational range

of 1361 m (Bowers and McLaughlin 1966) similar to the Sierra de

Bacadéhuachi. But given the strong influence and connections with

the SMO, it is expected that further sampling of the Sierra de Bacadéhuachi in foothillsthornscrub, desert grassland, and oak woodland

along the 700 to 2370 m elevational gradient (range of 1670 m) at

other times of the year will ultimately bring the number of moth taxa

to well over that recorded for the Huachucas.

Summer Sampling (July 31-August 3, 2011)

	 Thisinitialsample during the monsoon period documented around

205 species. The number of tropical taxa more typically associated

with tropical deciduous forest (TDF) was surprising. These included

the extremely large tailed saturniid moth Dysdaemonia boreas, a

common monsoon species in TDF where the larval foodplants in

the family Bombacaceae occur, and another saturniid moth, Copaxa 

multifenestrata, which was previously thought to be restricted to

extreme southeastern Sonora where the caterpillars are thought to

feed on plants in the family Lauraceae. Also recorded were tropical

sphingidsXylophanes ceratomoides andX. tersa, andoneofthe largest

tropical sphinx moths, the Forestiera-feeding Sphinx leucophaeta.

A single specimen of the metallic, tropical wasp-mimic tiger moth,

Phoenicoprocta lydia,was also an unexpected record.Northern range

extensionsforseveralspeciesmore typical oftheYécora plateau (120

km to the south) were also significant new records: Coloradia prchali and Copaxa muellerana, (both Saturniidae), Epicrisias eschara,

Ammalo nr. paranomon, and Amastus (two species; all Erebidae;

Arctiinae) These records also reflect the association of this area with

higher and more mesic regions in the SMO in Chihuahua to the east.

A noteworthy southern record of the large silkmoth Hyalophora 

gloveri was also obtained for a Rocky Mountains species with only

a handful of Sonoran records.

	 While noctuids (cutworm moths) were not particularly speciose

in August, some of those that were collected were important records

and several are thought to represent new and undescribed species.

Unusual northern records include Cropia europs, Dypterigia dolens

(both Noctuidae:Amphypirinae), Coenipeta bibatrix (Erebidae: Erebinae), and Metria sinaloa (Erebidae: Catocolinae). Schinia simplex,

a lovely green noctuid (Heliothinae) found in the Rocky Mountain

region nearly to Canada, was an unexpected southern record, asthere

are no SE Arizona records of the species. A new species of Richia nr.

cofrensis (Noctuidae: Noctuinae) was collected along with possibly

newLicnoptera nr. gulo, a newPanthea nr. gigantea (bothNoctuidae:

Pantheinae), a new Bryolymnia nr biformata and a new Elaphria sp.

(both Noctuidae: Amphipyrinae). The possibility that they had not

been recorded as new species previously, Zale nr. obsita and Z. nr.

sabina (Erebidae: Catocolinae) were also collected. However, these

may be found to belong to the nominal taxa after DNA barcoding.

Zale sabina has a single Arizona record, while Z. obsita has been

recorded in southern Texas. Voucher specimens were forwarded to

Don Lafontaine, an Erebid/Noctuid expert at the Canadian National

Collection (CNC) in Ottawa, Canada, for DNA barcoding and further analysis. A specimen of Paraceliptera guerreronis (Erebidae:

Erebinae) and a specimen of the shiny white Chasmina mexicana

(Noctuidae: Stiriinae) collected on the August trip also proved to

be very significant records. Both of these moths were described

earlier in the 1900s from much farther south (Guerrero state) by the

German lepidopterist M. Draudt and figured in the famous German

lithograph series on Lepidoptera by Seitz. The original type and

paratype specimens of these moths were destroyed in 1945 during

WWIIin the bombing ofDresden,Germany.The specimens collected

at RG represent the firstspecimens collected since the type series and

are currently with Don Lafontaine for DNA barcoding and study.

These specimens will likely be designated neotypes of the two species. In addition to noctuids, two undescribed species of slug moths

(Limacodidae) were collected at RG and specimenssent to specialist

Marc Epstein in Sacramento, California. These specimens have been

tentatively assigned to the genera Natada and Miresa.

Fall Sampling (September 3-5, 2011) 

	 Conducted only a month after the initial sample, this trip yielded

214 identified taxa with almost half of these representing species not

recorded in August. Virtually all of the “giant moths” seen a month

earlier were absent as were most of the species with tropical affinities. Instead, there was an explosion of smaller moths, particularly

those that feed on flowers. The noctuid family of Acontiinae moths,

better known as “bird dropping moths,” was particularly plentiful

with 37 species in several closely related genera. Other noteworthy

diversity was noted in the noctuid family Stiriinae, with 14 species

records.Especiallyabundantwasthebrightorange “Goldenrodmoth”

Cirrhophanus dyari, with more than 100 showing up at the lights.

A noctuid collected on this second trip was sent to Chris Schmidt in

Canada, who recently revised the genus Charadra north of Mexico;

He confirmed the moth as new—C. nr. moneta. An additional specimen of the rare Chasmina mexicana was recorded.

	 Larval activity was noteworthy in the fall sampling, and served as

evidence that a flurry of moth activity had occurred with the onset

of the monsoon rains. When known, these larvae were photographed

and catalogued as records for MABA. Two particularly noteworthy

species were Crinotes beidermani (Notodontidae), which had virtually denuded the Ceonothus buxifolius bushes around the compound,

and Eupackardia caletta (Saturniidae) that was collected by a local

cowboy on Rhus aromatica, a previously unrecorded foodplant for

this moth.

Spring Sampling (April 1-3, 2011)

	 Despite cool evening temperatures and dry conditions, a spring

sample at RG yielded an additional 122 taxa, 104 of these not being

previously recorded.The Geometridae were especially diverse in this

sample with 50 taxa collected including two possible new species.

Several geometrids that are rarely encountered in the United States,

and onlyinSE Arizona,were found tobe common.Stamnodes apollo,

Hydriomena mediodentata and Pityeja picta were all abundant here.

While the silkmoths (Saturniidae) are typically expected during the

summer monsoon, a freshly emerged male Copaxa lavendera was an

unusual and new record for RG. Like C. multifenestrata, the larvae

of this moth are usually associated with plants in the Lauraceae. But,

this widespread tropical species has also been reared in captivity on

Quercus, amore likelyhostplantinthislocality.Theoccurrenceofthis

tropicalspeciessoearlyintheseasonindicatesthatwintertemperatures

must be comparatively mild at RG even though the resident vaqueros

reported snow on the ground only a month before. Other interesting

recordsinclude anunidentifiedwhiteArtace spp(Lasiocampidae) and

surprising spring broods (based on the numbers of individuals seen)

of Caripeta pulcherimma and Pityeja picta (Geometridae), both of

which occur as univoltine summer species in Arizona, Pityeja in the 

Chiricahuas only.
USDA Forest Service Proceedings RMRS-P-67. 2013 171

Preliminary Assessment of the Moth (Lepidoptera: Heterocera) Fauna of Rincon de Guadalupe, . . . Palting

Discussion and Conclusions

	 The mostsurprising aspect of the RG locality has been the number

oftropicaltaxaencounteredmuchfurthernorththanpreviouslyknown,

especially given the lack of contiguous records between here and

previously documentedpopulations.Particularly strikingexamplesof

this areChasmina mexicana,Copaxa multifenestrata,Cropia europs,

Metria sinaloa,Paraceliptera guerreronis and Phoenicoprocta lydia.

This likely is the result of sampling bias in the northern SMO, with

more sampling historically occurring in the Alamos area. The tropical Sphingidae encountered are less surprising given their ability to

feed and disperse across long distances as adults. Copaxa lavendera

encountered in the spring sampling was also less surprising as it is

ratherwellknownintheYécora area.Twosaturniidspreviouslyknown

from Sonora only from the vicinity of Yécora, Coloradia prchai and

Copaxa muellerana, representsignificantrange extensionsforunusual

SMO species.Coloradia prchali is an especially unique species, with

males being the only members of the genus that are diurnal while the

females only fly after dark. This species was only described in 1992

and it remains a near-endemic species for Sonora (with records from

nearby Chihuahua) with a very limited range.

Ceiba acuminata is occasional in foothills thornscrub at lower

elevations in the Sierra de Bacadéhuachi and must be the host plant

of the Dysdaemonia boreas taken in early August. While seemingly

out of place among the pines, Peter Hubbell reported that tropical

moths often stray into the upland pine-oak forest in Guatemala (pers.

comm. through T. Van Devender).Asthe adults of the Saturniidae do

not feed, they have rather limited dispersal abilities. However, they

do have significant fat energy reserves and they tend to fly up hill as

do many Lepidoptera and other insects, so the specimen must have

started life further down the slopes ofthe Sierra deBacadéhuachi.The

occurrence of this large tropical moth so far north is further evidence

of the warmer temperaturesin SMO versus northern pine-oak forests

that allow‘tropical’plantstosurvive incloseproximitytotheoak/pine

elevation.The C. multifenestrata record may also indicate a “pocket”

of unsurveyed tropical vegetation near RG as this species is known

to feed only of Lauraceae, a rather rare and restricted plant family

in Sonora with no records of members being in the Sierra deBacadéhuachi.As the related (but more widespread) C. lavendera feed on

both Lauraceae and Fagaceae, it is possible that C. multifnestrata is

utilizing a plant in another family, but the complete lack of records

of the species between RG and Alamos remains confounding.

	 TheconnectionbetweenRGandthemainSierraMadreofChihuahua

was evident through several species more commonly encountered in

Chihuahua than in Sonora. These include the two species of Amastus

and the Ammalo nr. paranomon as well as Copaxa lavendera. The

beetle Chrysina adelaida (Coleoptera; Scarabeidae) recorded during

both the summer and fallsamplings atRG is a SMO speciesrestricted

in the Yécora area in Sonora and more commonly encountered in

mountains of Chihuahua.

	 Unusually high numbers (hundreds of individuals) of the bright

orange and black tiger moth Epicrisias eschara were encountered at

RG in August. This moth does not occur in the United States and its

life history (larva and foodplant) remains unknown. The moth was

originally described from south of Mexico City (Dyar 1912) and

was previously known to occur along the SMO axis as far north as

Yécora. In 2010, two specimens, indicative of a small population,

were documented in the Sky Island of Sierra La Madera and sampling

at RG indicates a robust population in the Sierra de Bacadéhuachi.

It would be interesting to correlate the distribution of this moth with

a particular host plant, and to see if the plant species also decreased

in abundance between RG, Sierra La Madera, and Arizona.

	 The rediscovery of two taxa “lost” to science was exciting, especially given that the original types were described fromGuerrero state

nearly a century ago. This and the number of new taxa identified at

RG are indicative of how incomplete our knowledge is of biodiversity

in northern Mexico. New species recorded from RG include at least

two members of the Erebidae and six members of the Noctuidae. All

specimens of the Prominants(Notodontidae) were sent to Jim Miller,

AMNH, New York, for review, and he tentatively believes at least

four of these are undescribed, including a new Disphragus sp and

Psilacron sp. Geometridae from the spring sampling are also likely

to include two undescribed species in the genus Hydriomena (Geometridae: Laurentiinae), and the Limacodidae sample also included

two undescribed taxa.

	 The intent of this initial survey of moths is to provide a starting

point in the MABA database for future bioinventories of the Sierra

de Bacadéhuachi and a comparison to other locations, particularly

the Sky Island area to the north. SE Arizona’s Sky Islands, while

fairly well- inventoried for the last 100 years, still yield new species

of moths every few years. The gap in our knowledge of species really lies to the south in the Sky Islands of Sonora and in the northern

SMO proper. It is not at all surprising that more than a dozen species

new to science have been identified from RG, given the paucity of

sampling that has occurred in this area. It is a good reminder of the

exciting biology and endemicity that has developed in this region,

where the new world tropics and the temperate zones meld together.

As land use change in the northern sierra escalates to meet human

needs, it becomes critically important that we know what is there

so we can preserve some of it for the future. The relatively pristine

state of RG compared with much of the heavily logged SMO makes

it a standout for preservation from both natural history and human

cultural history perspective.

Acknowledgments

	 Special thanks to Don Lafontaine and Chris Schmidt, Canadian

National Collection of Arthropods, Ottawa, Canada for their expert

identification of many of the Erebid/Noctuid species found at RG.

Thanks also to Jim Miller, American Museum of Natural History,

New York and to Mark Epstein, California Department of Agriculture, Sacramento, California, for identification of Notodontidae and

Limacodidae, respectively.

References

Bowers, Janice E.; and Steven P. McLaughlin. 1996. Flora of the Huachuca

Mountains, a botanically rich and historically significant Sky Island in

Cochise County, Arizona. Journal of the Arizona-Nevada Academy of

Science 29: 66-107.

Dyar, Harrison G. 1912. Descriptions of new species and genera of Lepidoptera, chiefly from Mexico. In: Proceedings of the United States National

Museum. Washington, DC: Smithsonian Institute. Volume 42:39-106.

Powell, J.A. and Paul A. Opler. 2009. Moths of western North America,

University of California Press.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
172 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Sand Dune of Ruby, Arizona, an Anthropogenically 

Created Biodiversity Hotspot for Wasps and Their Velvet 

Ant Parasitoids

Justin O. Schmidt

Southwestern Biological Institute, Tucson, Arizona 

Abstract—A large artificial sand dune composed of finely crushed mine tailings was produced by deep 

mining operations at Ruby, Arizona. Today, the ghost town of Ruby is an important historical location and 

biodiversity refuge, with the newly formed dune forming the core of the refuge. The dune provides ideal 

nesting habitat for at least 13 species of sand-loving wasps, including cicada killers (Hymenoptera: Sphecius) 

that are sometimes present in record breaking populations. Following the colonizing wasps were at least 14 

species of velvet ants (Hymenoptera: Mutillidae) that parasitize the wasps. Host wasp species range in size 

from about 9 to 1000 mg; various opportunistic velvet ant species exploit the entire host range of available 

wasps, resulting in velvet ants ranging in size from 6 to about 250 mg. These findings indicate that colonization of the man-made Ruby dune by both wasps and their parasitoids occurred rapidly after dune formation.

Introduction

	 Ruby was a prominent mining town in the Atascosa region of

south-central Arizona near the border with Sonora. The town was

named Ruby in 1912, not for the gemstone, but for Julius Andrews’s

wife, Ruby, who he certainly must have felt was a true gem (Weiser

2010). The town formed around vast, rich ore deposits that were

mined on a small scale during the late 19th and early 20th Centuries.

Starting in 1926 with the purchase of the mine by the Eagle-Picher

LeadCompany,the settlementbecame aboomtownthroughthe1930s,

before closing in 1941. At its peak production in 1934-37, the mine

was the largest producer in Arizona of lead and zinc and third largest

in silver. Culturally, Ruby was a well operated company town reaching a population in its heyday of over 1200 inhabitants, and became

best known for a pair of gruesome double murders in 1920-21 (Ring

et. al. 2005; Weiser 2010).

	 A byproduct of the mine was a huge sand dune generated from the

pulverized tailings ofthemined ore.The sand tailingswere dumped in

the canyon next to the mine, eventually filling it to a height over 15 m

and creating two lakes upstream. Although the exact date of the sand

dune’s creation is not clear, it undoubtedly began reaching its present

size during the period of 1926-1940. Consequently, this man-made

island of sand among the Sky Islands of the Madrean Archipelago

is extremely recent in geologic terms. Currently the dune is home to

a myriad of sand-loving wasps, mainly sphecid wasps (Crabronidae

and Sphecidae) plus various paparazzi-like parasitic insects that follow their activities and exploit their nest and larval stores. The most

abundant of these parasites are the velvet ants (family Mutillidae).

Sphecid wasps, sometimes called hunting wasps, prey on various

insects and spiders, which they usually subdue via a paralyzing sting.

Inmost casesthe haplessinactivated prey issubsequently transported

to a nest constructed by the wasp. Nests consist of several cells, each

of which serves as the incubator for a wasp larva that feeds on the

provisioned prey. When cells are completed with adequate food supplies, they are sealed and abandoned, leaving the larvae to develop

on their own to form the next generation (O’Neill 2001). 

	 Sphecid wasp nests are exploited by a variety of other wasps, flies,

and others having a parasitic lifestyle.Aggregations ofsphecidwasps,

as in a limited sand dune area, present an abundance of resource opportunities,thereby often attracting large populations ofmany species

of parasites.At Ruby, the most conspicuous of these parasites are the

velvet ants. Not true ants,rather wasps with wingless ant-like females

and winged males, velvet ants are among the hardest, most colorful,

and best-defended insects known (Schmidt and Blum 1977). They

are technically parasitoids because the larvae consume and kill the

prey rather than just feeding, usually non-lethally, on a host as does

a typical parasite. The hosts of many velvet antspecies are unknown,

in large part as a product of their lifestyles of entering the hidden

cells of their hosts where they lay an egg on the quiescent host pupa

or prepupa. Velvet ants frequently are considered opportunists that

accept a wide variety of different species and sizes of hosts. Hosts

are mainly bees and wasps (Brothers 1972).

	 The purpose of this investigation is to document the colonization

of the Ruby sand dune by sand-loving wasps and their associated

parasitic velvet ants.

Materials and Methods

	 All surveys were conducted exclusively on the Ruby dune proper

fromaboutmid-Augustthroughmid-September2009-2011.Thedune

is located at 31o 27’ 30.90” N, 111o 14’ 00.04” W, elevation 1267 m.

It measures approximately 334 x 143 m in length and width and has

a total surface area of 4.00 x 104 m2 (4.00 hectares) (fig. 1). Within

the dune are various areas including broad flat bare areas, small

sparsely grassed stable hummocks elevated about a meter above the

plain, larger eroding buttes 1-3 m in height, and deep eroding cliffs
USDA Forest Service Proceedings RMRS-P-67. 2013 173

Sand Dune of Ruby, Arizona, an Anthropogenically Created Biodiversity Hotspot for Wasps . . . Schmidt

where the dune meets the lower valley floor of the drainage basin.

The sand throughout is very fine, uniform, and densely packed. 

	 Wasps were collected manually and by insect net during daylight

hours, placed onice, andtransported tothe labwhere theirliveweights

and identities were determined. Voucher specimens are deposited in

the University of Arizona Entomology Collection.

Results

	 Thirteen species of sand wasps were recorded on the dunes. Six of

these were very common to abundant.The largest and most conspicuous sand wasp in Ruby, and perhaps of Arizona, is Sphecius convallis, the Pacific cicada killer, so named because it paralyzes cicadas

to provision its larvae. These huge bright yellow and light brown

wasps attain lengths of 4 cm and weigh more than 1 g, with males

usually much smaller than females (Coelho 1997). Ruby appears to

have the largest population of these wasps known with well over a

thousand individuals present during good years.Anybody visiting the

dunes during their active period cannot avoid noticing these wasps,

especially the males actively patrolling territories and searching for

females.

	 The second largest wasp on the dune is the “great black wasp”,

Sphex pennsylvanicus, which weighs on average an estimated 300

mg and hunts katydids. This conspicuous species is common around

the cliffs and buttes regions of the dune, though its numbers are considerably fewer than the cicada killers. Ammophila sp., a caterpillar

hunter, weighs about 100 mg and can be observed resting on the sand

or hovering just above itinmany locations around the dune.Itsslender

shape, but not color pattern, makes this common wasp conspicuous.

The large bembicine wasp, Bicyrtes viduata, actively flies and chases

othersin large aggregationsin the flat areas near the dune center. This

abundant and striking black and yellow wasp weighs on average 83

mg and paralyzes Hemiptera as prey. A smaller bembicine wasp,

Microbembix sp.,is evenmore abundant,perhapsbeingthemost common species on the dune.This 26mgwasp,resembling a smallBicyrtes,

nests in the flat central dune area as well as other areas. It provisions its

young with dead arthropods and fragments. A final abundant member

of the big six common wasps is Tachysphex near terminatus. Unlike its

larger wasp co-inhabitants, thistiny wasp, weighing only about 9 mg, is

neither flashy, nor conspicuous. It is most prominently found around the

hummock area of the dune where it provisionsits nest with immature

katydids and other orthopteran relatives. 

	 In addition to the common dune wasps, are an assortment of infrequentlyencounteredspecies.These species,usuallyobservedsinglyor

in low numbers, range in size from a large second species of Bicyrtes

weighing about 150 mg to the 18 mg Stenodynerus apache. Five others species: Cerceris sp., Eucerceris sp., Dolichodynerus vandykei,

Podalonia sericea, and an unidentified sphecid are present. Others

undoubtedly also make the dune home.

	 A large, diverse guild of velvet ants shares the dune habitat with

the sand wasps. These species range widely in size from the 6 mg

Dasymutilla monticola to the nearly 250 mg D. klugii (D. magnifica 

is excluded, as only one specimen was found and the species is generally equal in size to D. klugii)(table 1). Twelve of the 14 species

form a gradual size gradient from about 6 mg through 80 mg. The

conspicuous outliers are the two ecological siblings, D. klugii and

D. magnifica, which are about 3 times as large as the nearest sized

species. In addition to filling all available size niches at the species

level, the individual velvet ant species span enormous ranges in the

size of individuals. For the most abundant species, those with three

or four stars of relative abundance in table 1, the average size range

Figure 1–Google Earth image of Ruby dune and surrounding area.
174 USDA Forest Service Proceedings RMRS-P-67. 2013

Schmidt Sand Dune of Ruby, Arizona, an Anthropogenically Created Biodiversity Hotspot for Wasps . . .

from smallest individual to largest individual is 5.26 (fig. 2). In

comparison, the average size range within the most abundant sand

wasps at Ruby is under 2.0 (data notshown). These numbersindicate

that the sand wasps are more specialized, using a limited number of

prey species, and that they additionally might have the opportunity to

optimize the prey provisioned for each of their larvae. The wide size

range for many velvet antspeciesreflectsthe fact that each velvet ant

larva can only attain the size allowed by the available food in its cell;

that is, the size of the host parasitized by its mother. If the mother’s

opportunities bring her to a host species with small individuals, the

offspring will be small; if she finds host species have large individuals, the consequent velvet ant offspring will be large.

Discussion

	 The recent formation of the Ruby sand dune about 80 years ago

provides an opportunity to investigate the speed of colonization of

this new habitat. No other large man-made or natural sand dunes are

present near Ruby and sand areasin the vicinity are scarce,small, and

likely ephemeral areas along washes and stream courses. All wasps

are partially limited by food resources (O’Neill 2001), but perhaps

a more important factor for many species is availability of suitable

nesting sites. If sand for nesting is scarce in the environment, that

factor might be the crucial limitation in the life history of the species.

The Ruby sand dune is unique in two ways: first, it is a huge sand

resource for sand wasps; second, it is extremely recent in geologic

time. The importance of the dune as a nesting resource is evident by

the extreme population densities of wasps nesting in the dune and the

large number of species present. The enormous numbers of cicada

killers, a species rarely seen in other areas, is evidence that the environmental constraint on that species is not prey availability or prey

carrying capacity, but, rather it is constraints on nesting locations.

This flagship species provides a contrast to numerous other sand

wasp species on the dune, many of which occur as in small numbers.

The question arises, are these less common species limited by prey

availability, some other factor, or were they constrained by ability to

disperse to the dune from other areas? Are they recent arrivals that

have not had adequate time to expand their populations? Long-term

studies might well shed light on these questions.

	 Unlike the sand wasps, all of which can disperse by flight to arrive

at Ruby, the velvet ant females are all flightless. That meansto follow

their wasp hosts and to colonize the dune, they needed to crawl there.

Given the potential distancesinvolved and the sizes ofthe velvet ants,

this seems a remarkable feat; especially given that at least 14 species

have colonized the dune is less than 80 years. Collections of both

sand wasps and velvet ants in the areas of Arizona and Sonora that

surroundRuby have been sparse.We, therefore, have little knowledge

concerning the distance the velvet ants might have been required to

travel to get to Ruby, or the densities of individuals in the pools from

whichthe immigrants came.Inotherlocations aroundArizona,velvets

ants are infrequently encountered, an exception being in the stabilized

dune area near Willcox, Arizona, that has as many or more species

than Ruby (Justin Schmidt, personal observations). The Willcox

area contrasts Ruby in that it is a much older sand formation and is

surrounded by mainly flat surface terrain.

	 Specifichostsforvelvet ants are infrequentlyknown.One exception

is the relationship between cicada killers and the two largest velvet

ants at Ruby, D. klugii and D. magnifica. Both of these enormous

velvet ant species require equally enormous hosts, and cicada killers

fit the bill. Indeed, western cicada killers are recorded as the host for

D. klugii(Bradley 1920).Thisrestriction to one hostspecies(perhaps,

in exceptional cases, Sphex pensylvanicus can be an alternate host

to produce small individuals) for D. klugii is reflected in the narrow

size range from smallest to largest individuals of only 3.25.Asimilar

prey limitation might also occur for D. nogalensis, the second largest

species at Ruby, which also has the small size range in the order of

2.39 among individuals. In contrast to these large species, the small

and medium sized velvet antspecies exhibit huge size ranges up to an

over 8-fold difference issize between smallest and largest individuals

in a species. Thissuggeststhat these smallerspecies are not nearly as

limited in their range of available hosts, and likely parasitize almost

any available host on the dune within a wide size range.

Acknowledgment

	 I thank Howard and Pat Frederick who generously encouraged this

research, made the study site available, and provided background

information and advice and Tom Van Devender and John Palting for

reviews and comments on the manuscript.

References

Bradley, J. Chester. 1920. Note. Entomological. News 31: 112-13.

Brothers, Denis J. 1972. Biology and immature stages of Pseudomethoca f. 

frigida, with notes on otherspecies(Hymenoptera: Mutillidae). University

of Kansas Science Bulletin. 50: 1-38.	

Coelho,Joseph R. 1997. Sexualsize dimorphism and flight behavior in cicada

killers, Sphecius speciosus. Oikos. 79: 371-75.

O’Neill, Kevin M. 2001. Solitary wasps behavior and natural history. Ithaca,

NY: Cornell University Press. 405 p.

Ring, Bob; Ring, Al; and Cahoon, Tallia P. 2005. Ruby, Arizona–mining,

mayhem, and murder. Tucson: U.S. Press & Graphics. 280 p.

Schmidt, Justin O.; and Blum, Murray S. 1977. Adaptations and responses

of Dasymutilla occidentalis (Hymenoptera: Mutillidae) to predators. Entomologia Experimentalis et Applicata. 21: 99-111.

Weiser, Kathy. 2010. Mining and murder in Ruby, Arizona. Legends of

America [Online]. Available: http://www.legendsofamerica.com/az-ruby.

html [May 9, 2012].

Figure 2–Size extremes in two velvet ant species, Dasymutilla chiron

and D. asteria.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 175

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Bird List of San Bernardino Ranch in Agua Prieta, 

Sonora, Mexico

Melinda Cárdenas-García and Mónica C. Olguín-Villa

Universidad de Sonora, Hermosillo, Sonora, Mexico

Abstract—Interest and investigation of birds has been increasing over the last decades due to the loss of their 

habitats, and declination and fragmentation of their populations. San Bernardino Ranch is located in the 

desert grassland region of northeastern Sonora, México. Over the last decade, restoration efforts have tried to 

address the effects of long deteriorating economic activities, like agriculture and livestock, that used to take 

place there. The generation of annual lists of the wildlife (flora and fauna) will be important information as 

we monitor the progress of restoration of this area. As part of our professional training, during the summer 

and winter (2011-2012) a taxonomic list of bird species of the ranch was made. During this season, a total 

of 85 species and 65 genera, distributed over 30 families were found. We found that five species are on a 

risk category in NOM-059-ECOL-2010 and 76 species are included in the Red List of the International Union 

for Conservation of Nature (IUCN). It will be important to continue this type of study in places that are attempting restoration and conservation techniques. We have observed a huge change, because of restoration 

activities, in the lands in the San Bernardino Ranch. 

Introduction

	 Birds represent one of the most remarkable elements of our environment, because they’re easy to observe and it’s possible to find

them practically anywhere. Approximately 12% of all bird species

in the planet are found in Mexico, this corresponds to 1060 species

of a total of 8500 (INECOL 1996). This enormous diversity is due,

among other reasons, to a bird species with Nearctic (NorthAmerica)

and Neotropical (Middle and South America) affinities; Mexico is

the contact zone between these two biogeographic regions(INECOL,

1996).

	 Sonora varies seasonally from moist and cool to dry and hot.

Here the northern fringe of the tropics meets the southern end of the

temperature region, bringing plants and animals from both sources

into a single state (Rusell and Monson 1998). Due to this, Sonora has

varied habitats like the deserts, grassy plains, and rugged mountains

that may be arid and barren, or forest clad and split by great canyons.

River valleys are bordered by gallery woodlands or by irrigated fields

(Russell and Monson 1998). These characteristics produce a rich,

diversified, and interesting biota, which still awaits full study.

	 According to the book Diversidad Biológica de Sonora (MolinaFreaner and Van Devender 2009). the birds of Sonora include 556

species in 73 families and 20 orders, with 227 resident species, 46

summer residents, 223 long distance migratory and 50 partially

migratory (Villaseñor-Gómez et al., 2010). Twenty-eight of those

species are considered at risk on a global scale, and are included in

the Red List of the International Union for Conservation of Nature

(IUCN). Two-hundred-ninety are in the Neotropical Migratory Bird

Conservation Act (NMBCA) of the United States and 66 species in

the Mexican Official Norm NOM-059-ECOL-2001 (SEMARNAT

2010).

	 The length of the migration period may vary considerably among

different species. Most of the Sonoran birds (41%) are year-round

residents. Only 8.3% are summer residents that nest after returning

from a winter stay to the South. Another group is composed of birds

that breed in the United States and Canada, and then migrate south

for varying distancesto spend the colder monthsin Sonora or passing

through to continue as far as southern South America (Russell and

Monson 1998).

	 The studyarea is consideredasthe continuation ofArizona’sSoutheast birds and the extreme Southeast New Mexico’s birds, bringing

a mix of species with tropical and neartic distributions, becoming an

ecotone between the Chihuahuan and Sonoran Desert, and the Sierra

Madre Occidental that divides both deserts (Molina-Freaner and

VanDevender, 2009). Thisregion is part of the MadreanArchipelago

that constitutes mountain formations isolated by “oceans” of grass

and deserted shrubs, acting like barrels or bridges that limit or make

a possible genetic inter, 2011).

	 The areaofSanBernardinohas ahistoryof agriculture andlivestock

exploitation beyond the recovery capacity of the land. In the last 20

years,restorationprocesseshavebeenimplementedwithgoodresults,

making it necessary to intensify politics and economical resourcesto

maintain and continue the ecological improvement (CEDES 2011).

	 Birds are very sensitive to environmental changes and, therefore,

they can be used as indicators of environmental transformations,

through changesin their distribution and temporal differencesin their

abundance (Hutto 1998). Research has increased in the last decades,
176 USDA Forest Service Proceedings RMRS-P-67. 2013

Cárdenas-García and Olguín-Villa Bird List of San Bernardino Ranch in Agua Prieta, Sonora, Mexico

Table 1—SBR total species of birds.

Order Family Common name Summer Winter NOM-059 IUCNa

Scientific name -ECOL-2010a

Accipitriformes Accipritidae

Accipiter cooperi Cooper’s Hawk x Protección especial (PR)

Buteo albonotatus Zone-tailed Hawk x Protección especial (PR)

Buteo jamaicensis Red-tailed Hawk x x Least Concern

Buteo regalis Ferruginous Hawk x Protección especial (PR) Least Concern

Circus cyaneus Northern Harrier x Least Concern

Parabuteo unicinctus Harris’s Hawk x Protección especial (PR) Least Concern

Cathartidae

Cathartes aura Turkey Vulture x x Least Concern

Anseriformes Anatidae

Anas strepera Gadwall x Least Concern

Aythya collaris Ring-necked Duck x Least Concern

Oxyura jamaicensis Ruddy Duck x Least Concern

Apodiformes Apodidae

Aeronautes saxatalis White-throated Swift x Least Concern

Trochilidae

Archilochus alexandri Black-chinned Hummingbird x Least Concern

Cynanthus latirostris Broad-billed Hummingbird x x Least Concern

Selasphorus platycerus Broad-tailed Hummingbird x

Caprimulgiformes Caprimulgidae

Chordeiles acutipennis Lesser Nighthawk x Least Concern

Chordeiles minor Common Nighthawk x x Least Concern

Charadriiformes Charadriidae

Charadrius vociferus Killdeer x x Least Concern

Scolopacidae

Actitis macularius Spotted Sandpiper x Least Concern

Columbiformes Columbidae

Columbina inca Inca Dove x x Least Concern

Streptopelia decaocto Eurasian Collared-Dove x Least Concern

Zenaida asiatica White-winged Dove x x Least Concern

Zenaida macroura Mourning Dove x x Least Concern

Cuculiformes Cuculidae

Geococcyx californianus Greater Roadrunner x x Least Concern

Falconiformes Falconidae

Falco columbarius Merlin x Least Concern

due to the decline of many populations and fragmentation or loss of

their habitats (Vickery et al., 1999). In recent years, vast areas have

been cleared of all existing plant life and planted with African buffel

grass (Poaceae: Pennisetm ciliare), creating a monoculture of little

value to wildlife. Even more of Sonora is destined to be altered this

way (Bowden 1993).

	 The objective or this work is to know the composition of bird

species present in the San Bernardino Ranch, and contribute to the

listing update of the wildlife of the area.

Area Description and Methods

	 The San Bernardino Ranch is located 31 kilometers from Agua

Prieta-Janos road 31° 19’ 0.20” N & 109°15’ 59.91” inside the State

of Sonora. The observations occur on five differentsites, locality one

(L1) main house, locality two (L2) Silver Creek River, locality three

(L3) San Bernardino River, locality four (L4) dam, locality five (L5)

Cienega. The total area of observation was 3.9 km2.

	 Threeperiodsofobservationswereperformedtwotimesindaylight

around 5 am to 11 am and from 5 pm to 7 pm. The first period was

on July 4 to 10, 2011, called summer observation, and the other two

periods were December 13 to 18, 2011, and March 14 to 18, 2012,

being the winter observations. These dates were decided due to being part of our professional practices in our career as an obligatory

process for our graduation.

	 Bird species were registered using direct observation by binoculars

(10x42ALPENand12x42BUSHNELL) andthe identificationbyfield

guides of birds such as Kaufman (2005), and Sibley (2000). There

was no singing identification. A list of recorded species is presented

as a main result of the study.

Results and Discussions

	 The total of species for all periods was of 85 species distributed

in 30 families and 65 genus (table 1). On the summer observation

(June) we observed a total of 46 species in 23 families and 42 genus.

The winter observation’s (December and March) results were of 63
USDA Forest Service Proceedings RMRS-P-67. 2013 177

Bird List of San Bernardino Ranch in Agua Prieta, Sonora, Mexico Cárdenas-García and Olguín-Villa

Table 1—Continued.

Order Family Common name Summer Winter NOM-059 IUCNa

Scientific name -ECOL-2010a

Galliformes Odontophoridae

Callipepla gambelii Gambel’s Quail x x Least Concern

Gruiformes Rallidae

Fulica americana American Coot x x Least Concern

Passeriformes Bombycillidae

Bombycilla cedrorum Cedar waxwing x Least Concern

Cardinalidae

Cardinalis cardinalis Northern Cardinal x x Least Concern

Cardinalis sinuatus Pyrrhuloxia x x Least Concern

Passerina caerulea Blue Grosbeak x Least Concern

Piranga rubra Summer Tanager x Least Concern

Corvidae

Corvus corax Common Raven x Least Concern

Corvus cryptoleucus Chihuahuan Raven x x Least Concern

Emberizidae

Aimophila carpalis Rufous-winged Sparrow x

Aimophila ruficeps Rufous-crowned Sparrow x Least Concern

Amphispiza belli Sage Sparrow x Least Concern

Amphispiza bilineata Black-throated Sparrow x x

Amphispiza quinquestriata Five-striped Sparrow x Least Concern

Chondestes grammacus Lark Sparrow x Least Concern

Melospiza melodia Song Sparrow x Least Concern

Passerculus sandwichensis Savannah Sparrow x Amenazada (A) Least Concern

Pipilo chlorurus Green-tailed Towhee x Least Concern

Pipilo fuscus Canyon Towhee x x Least Concern

Pooecetes gramineus Vesper Sparrow x x Least Concern

Spizella atrogularis Black-chinned Sparrow x x Least Concern

Spizella passerina Chipping Sparrow x Least Concern

Zonotrichia leucophrys White-crowned Sparrow x Least Concern

Fringillidae

Carduelis psaltria Lesser Goldfinch x Least Concern

Carpodacus mexicanus House Finch x x Least Concern

Spinus tristis American Goldfinch x

Hirundinidae

Hirundo rustica Barn Swallow x Least Concern

Tachycineta bicolor Tree Swallow x Least Concern

Icteridae

Agelaius phoeniceus Red-winged Blackbird x Least Concern

Euphagus cyanocephalus Brewer’s Blackbird x Least Concern

Molothrus ater Brown-headed Cowbird x

Molothrus aeneus Bronzed Cowbird x Least Concern

Quiscalus mexicanus Great-tailed Grackle x Least Concern

Mimidae

Oreoscoptes montanus Sage Trasher x Least Concern

Parulidae

Dendroica petechia Yellow Warbler x Least Concern

Geothlypis trichas Common Yellowthroat x Least Concern

Vermivora luciae Lucy’s Warbler x

Vermivora ruficapilla Nashville Warbler x Least Concern

Troglodytidae

Cistothorus palustris Marsh Wren x Least Concern

Thryomanes bewickii Bewick’s Wren x Least Concern

Turdidae

Turdus migratorius American Robin x Least Concern

Tyrannidae

Contopus cooperi Olive-sided Flycatcher x Near Table 
178 USDA Forest Service Proceedings RMRS-P-67. 2013

Cárdenas-García and Olguín-Villa Bird List of San Bernardino Ranch in Agua Prieta, Sonora, Mexico

Table 1—Continued.

Order Family Common name Summer Winter NOM-059 IUCNa

Scientific name -ECOL-2010a

Threatened

Empidonax hammondii Hammond’s Flycatcher x Least Concern

Empidonax wrightii Gray Flycatcher x Least Concern

Pyrocephalus rubinus Vermilion Flycatcher x x Least Concern

Sayornis nigricans Black Phoebe x x Least Concern

Sayornis phoebe Eastern Phoebe x Least Concern

Sayornis saya Say’s Phoebe x x Least Concern

Myiarchus cinerascens Ash-throated Flycatcher x Least Concern

Tyrannus melancholicus Tropical Kingbird x Least Concern

Vireonidae

Vireo gilvus Warbling Vireo x Least Concern

Vireo huttoni Hutton’s Vireo x Least Concern

Pelecaniformes Ardeidae

Ardea heroidas Great Blue Heron x

Egretta caerulea Little Blue Heron x x Least Concern

Piciformes Picidae

Melanerpes formicivorus Acorn Woodpecker x Least Concern

Melanerpes uropygialis Gila Woodpecker x x Least Concern

Picoides arizonae Arizona Woodpecker x Least Concern

Picoides villosus Hairy Woodpecker x Least Concern

Sphyrapicus nuchalis Red-naped Sapsucker x Least Concern

Strigiformes Strigidae

Bubo virginianus Great Horned Owl x x Least Concern

Tytonidae

Tyto alba Barn Owl x Least Concern

Total 30 46 63 5 76

85

a

Species found in the NOM-059-ECOL-2010 and the International Union for Conservation of Nature are marked with the risk category under where they are established.

species in 28 families and 51 genus. In December we identified 35

species and in March 43 species were identified.

	 The 24 species that were in all observations were Black-throated

sparrow(Amphispiza bilineata),GreatHornedOwl(Bubo virginanus),

Red-tailed Hawk (Buteo jamaicensis), Gambel’s Quail (Callipepla 

gambelii), House Finch (Capodarcus mexicanus), Northern Cardinal

(Cardinalis cardinalis), Pyrrhuloxia (Cardinalis sinuatus), Turkey

Vulture (Cathartes aura), Killdeer (Charadrius vociferus), Common

Nighthawk (Chordeiles minor), Inca Dove (Columbina inca), Chihuahuan Raven (Corvus cryptoleucus), Broad-billed Hummingbird

(Cynanthus latirostris),LittleBlueHeron(Egretta caerulea),American

Coot(Fulica americana),GreaterRoadrunner(Geococcyx californianus),Acorn Woodpecker (Melanerpes uropigyalis), Canyon Towhee

(Pipilo fuscus), Vesper Sparrow (Pooecetes gramineus), Vermilion

Flycatcher (Pyrocephalus rubinus), Black Phoebe (Sayornis nigricans), Black-chinned Sparrow (Spizella atrogularis), White-winged

Dove (Zenaida asiatica), Mourning Dove (Zenaida macroura).

	 We found that five species are on a risk category in NOM-059-

ECOL-2010, four are in Special Protection (Pr): Cooper’s Hawk

(Accipiter cooperi),Zone-tailedHawk(Buteo albonotatus),Red-tailed

Hawk (Buteo jamaicensis), Ferruginous Hawk (Buteo regalis), Harris’s

Hawk (Parabuteo unicinctus); one isthreatened (A): Savannah Sparrow (Passerculus sandwichhensis); and none is endemic of Mexico;

76 species are included in the Red List of the International Union for

ConservationofNature (IUCN) asleast concern“LC” andone species

is near threatened “NT” (Olive-sided Flycatcher). This is important

because the risk species help to declare protected areas, ensuring the

preservation of the rich flora, fauna, and environmental services.

	 In August and October of 2005, Rob Hunt made a report of bird

observations in the San Bernardino Ranch, and found 61 species of

birds. We found 31 of those observed; we didn’t know exactly the

area where the observationstook place,so we can’t compare the total

or both results (Cuenca Los Ojos 2012).

	 The San Bernardino National Wildlife Refuge (SBNWR) located

in the border east of Douglas Arizona, next to the San Bernardino

Ranch, has 30 years of protecting water resources and habitats. They

have registered over 270 species of birds; if we consider those data

as 100% of the bird species present in SBNWR, we can say that we

have observed 31.48% of species. Also, the SBNWR has an area of

9.34 km2 and our area of observations was made on 3.9 km2 (U.S.

Fish & Wildlife Service 2011).

Conclusions

	 It is important to keep having this type of study in places that are

actively implementing restoration and conservation techniques. We

have seen a huge change in the lands in the San Bernardino Ranch;

there is more retained soil and water, which attracts many diverse

species. Also, we didn’t find considerable threatened species in the

Red List of IUCN and NOM-059-ECOL-2010, but we are aware

that actions of conservation and restoration in the future will bring a

healthy habitat that would help in the improvement and development

of the ecological interactions.
USDA Forest Service Proceedings RMRS-P-67. 2013 179

Bird List of San Bernardino Ranch in Agua Prieta, Sonora, Mexico Cárdenas-García and Olguín-Villa

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Acknowledgments 

	 We would like to thank to Valer Austin for all the support and time

he shared with us by getting to know the Ranch and working in it.

Also, to Cindy Tolle who helped in the making of this work, and to all

the people oftheRanch, Don Pancho, Lázaro, Lili,Juana,Bernardino

and Samuel. Special thanks to our professor M.Sc. Gilberto Solís

who introduced us to the Ranch; to Dra. Reyna A. Castillo Gámez

for letting us use her field material and helpful comments; and to Dr.

Alejandro Varela Romero for his help in the professional practices,

and some comments on the manuscript.Thanksto our friends Erandi,

Valeria, Cali, Carlos, Gert, David and JR for their support.

References

Bowden C. 1993. The secret forest. New Mexico: University of New Mexico.

141 p.

Comisión de Ecología y Desarrollo Sustentable del Estado de Sonora. 2011.

Estudio previo justificativo para proponer la región de San Bernadino, 

municipio de Agua Prieta, Sonora como Reserva Estatal. Gobierno del

Estado de Sonora, Cuenca Los Ojos, A. C. [Online]. Available:www.

cedes.gob.mx/LinkClick.aspx?fileticket=sZq5s6%2FMWdg%3D&tab

id=830 [April 23,2012]

Cuenca Los Ojos. 2012. Wildlife reintroduction. [Online] Available: http://

cuencalosojos.org/

Hutto, R. L. 1998. Using landbirds as an indicator species group. En: J.M.

Marzluff yR. Sallabanks, eds.Avian Conservation: Research and Management. Token of Diversidad Biológica de Sonora 385 p. Molina Freaner F.

E.; Van Devender T. 2009. 1ra edición. México: UNAM.

Instituto Nacional de Ecología (INECOL). 1996. Guía de aves canoras y de

ornato. 1era edición. México: CONABIO. 180 p.

Kaufman, K. 2005. Field guide to birds of North America. Houghton Mifflin

Company, New York.

Molina-Freaner,F. andT.VanDevender,2009.DiversidadbiológicadeSonora.

México: UNAM. 500 p.

Russell, S. and G. Monson. 1998. Birds of Sonora. University of Arizona

Press. Tucson, United States of America.

Secretaría de Medio Ambiente y Recursos Naturales. 2009. Plan de

Manejo tipo de aves canoras y de ornato. [Online].Available: www.semarnat.mx/temas/gestionambiental/vidasilvestre/Documents/Aves_Canoras_2011-2012.pdf. [April 12, 2012]

SEMARNAT. 2010. Norma Oficial Mexicana NOM-059-ECOL-2010.

Secretaría del Medio Ambiente y Recursos Naturales. Diario Oficial de

la Federación

Sibley, D. A. 2000. The Sibley guide to birds. National Audubon Society.

Allen A. Knopf, Inc. 544 pp.

U.S. Fish and Wildlife Service. 2011. San Bernardino and Leslie Canyon

National Wildlife Refuges. [Online] Available: www.fws.gov/southwest/

refuges/arizona/sanbernardino.html

Van Rossem, A.J. 1945. A distributional survey of the birds of Sonora,

Mexico. Louisiana State University. Occasional Papers of the Museum

of Zoology 21: 1-379.

Villaseñor-Gómez, J. F., O. Hinojosa-Huerta, E. Gómez-Limón, D. Kruepes,

andA. Flesh. 2009.Avifauna. In: Molina-Freaner, F.; and T. Van Devender,

(eds). Diversidad biológica de Sonora. México: UNAM.
180 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Preliminary Assessment of Species Richness and Avian 

Community Dynamics in the Madrean Sky Islands, Arizona

Jamie S. Sanderlin, William M. Block, Joseph L. Ganey, and Jose M. Iniguez

U.S. Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Abstract—The Sky Island mountain ranges of southeastern Arizona contain a unique and rich avifaunal 

community, including many Neotropical migratory species whose northern breeding range extends to these 

mountains along with many species typical of similar habitats throughout western North America. Understanding ecological factors that influence species richness and biological diversity of both resident and migratory 

species is important for conservation of this unique bird assemblage. We used a 5-year data set to evaluate 

avian species distribution across montane habitat types within the Santa Rita, Santa Catalina, Huachuca, 

Chiricahua, and Pinaleño Mountains. Using point-count data from spring-summer breeding seasons, we describe avian diversity and community dynamics. We use a Bayesian hierarchical model to describe occupancy 

as a function of vegetative cover type and mountain range latitude, and detection probability as a function 

of species heterogeneity and sampling effort. By identifying important habitat correlates for avian species, 

these results can help guide management decisions to minimize loss of key habitats and guide restoration 

efforts in response to disturbance events in the Madrean Archipelago.

Introduction

	 The Sky Island mountain ranges of southeastern Arizona, USA,

contain a unique and rich avifaunal community. Habitat diversity

frommixing ofMadrean andCordilleran flora supportsmany species,

including Neotropical migratory species whose northern breeding

rangesextendtothesemountains, andspeciestypicalofsimilarhabitats

throughout western North America. Many species of concern from

theNationalAudubonSociety/AmericanBirdConservancyWatchlist

(2007)andU.S.FishandWildlifeService(2008)BirdsofConservation

Concern(http://www.fws.gov/birdhabitat/Grants/NMBCA/BirdList.

shtm) occur within this area. The Mexican Highlands Partners In

Flight (PIF) Physiographic Area Plan (http://www.blm.gov/wildlife/

pl_81sum.htm, Sonoran Joint Venture Technical Committee 2006)

and Arizona PIF plan (Latta and others 1999) indicate conservation

issues of water use, urban development, overgrazing, and recreation.

Understanding ecological factors that influence species richness of

both resident and migratory species is important for conservation of

this unique bird assemblage, yet few studies describe bird habitat

requirements in the Sky Islands (e.g., Balda 1967; Block and others

1992; Block and Severson 1992; Hall and Mannan 1999; Marshall

1957). Past studies indicate riparian areas support the most bird species (Balda 1967; Strong and Bock 1990), but upland habitats also

contribute to species diversity (Marshall 1957).

	 We studied occupancy and cover type associations of forest birds

across montane vegetative cover types in the Santa Rita, Santa

Catalina, Huachuca, Chiricahua, and Pinaleño Mountains of southeastern Arizona from 1991 to 1995 (Block and others 1992; Block

and Severson 1992). We used occupancy models (MacKenzie and

others 2006) to estimate species richness and community dynamics

(local species extinction, local species colonization), while accounting for detection (e.g., Dorazio and others 2006). Imperfect detection

of species is important to include in analyses, especially with rare

or elusive species, or when trying to assess change over time. Estimates of species richness and community dynamics could be biased

if species occupy an area but were never detected during a survey or

multiple surveys. This could lead to biased study conclusions used

to guide management actions. Our objectives were to (1) estimate

species richness across mountain range and cover type; (2) relate

species occurrence to forest cover types within these ranges; and (3)

estimate probability of local species extinction and colonization.

Methods

Study Area

	 Ourstudy area (elevation: 1,470–3,000m) consisted ofwoodlands,

pine-oak forests(Pinusspp. – Quercusspp.), pine forests, and mixedconifer forests within the Santa Rita, Santa Catalina, Huachuca,

Chiricahua, and Pinaleño Mountains, of the Coronado National

Forest and Fort Huachuca (Department of Defense) in southeastern

Arizona, USA (fig. 1). Major tree species included Arizona white

(Q. arizonica), silverleaf (Q. hypoleucoides), Emory (Q. emoryi),

Gambel (Q. gambeli), and netleaf (Q. reticulata) oaks; ponderosa

(P. ponderosa), Apache (P. engelmanni), Chihuahua (P. leiophylla),

Mexican white (P. strobiformis), and border pinyon (P. discolor)
USDA Forest Service Proceedings RMRS-P-67. 2013 181

Preliminary Assessment of Species Richness and Avian Community Dynamics . . . Sanderlin and others

pines; alligator-bark juniper (Juniperus deppeana); white fir (Abies 

concolor); and Douglas-fir (Pseudotsuga menziesii) (Iniguez and

others 2005).

Field Sampling

	 Count points (n = 344) were located along transects consisting

of 12 points spaced at 300 m intervals, with the exception of one

transect that had only 8 points. Transects (n = 29) were established

using a systematic-random sampling design (Cochran 1977), and

occurred in proportion to occurrence of vegetation types in the

landscape. Using the variable-radius point count method (Reynolds

and others 1980), we sampled birds at each point three times during

each breeding season (Apr-July) from 1991 to 1995. Counts (5 min/

point) began within 30 min of sunrise and completed no later than 4

hr after sunrise. Observers remained still for 1-2 min after reaching

a point, then recorded species, age, and sex of birds detected.

	 We sampled diameter at breast height (DBH) of all trees and snags

in four 0.1-ha circular plots located within 100 m of each point count

station. We used these data to estimate a weighted percent basal area

average (WBA) of live and dead trees for each point count plot. We

used cluster analysis with WBA(Iniguez and others 2005) to classify

plots into 7 broad vegetative types for covariates in our occupancy

models: deciduous forest, mixed-conifer forest, pine-oak forest,

broadleaf evergreen woodland, conifer woodland, conifer riparian,

and ponderosa pine.

Analytical Analyses

	 Oursampling design was a robust design (Pollock 1982), in which

sampling occurred on up to three secondary periods (visits) during

each primary period (years). Occupancy states of species can change

with local extinction and colonization between primary periods,

but not during secondary periods. We condensed data across points

within each mountain × cover type combination to increase species

detectionsper combinationandreducemodel complexity.Usingthese

occupancy states with latent variable zihmt for occupancy of species i,

cover type h, mountain m at time t, we calculated derived parameters,

including species richness (N), species colonization (γ) and species

extinction (ε). We calculated the average number of species in each

mountain range and cover type over all 5 years:

	 We also calculated the average over all 5 years for local species

colonization (γ), probability that a species selected at random from

the community was not present in the community at time t-1, and

local species extinction (ε), or probability that a species selected at

random from the community was present at time t-1 but not at time

t (Williams and others 2002):

Figure 1—Study area from the Sky Island mountain ranges of Santa Catalina, Piñaleno, Santa Rita, Huachuca, and Chiricahua mountains of southeastern Arizona, USA from 1991 to 1995. Labeled dots within the mountain ranges indicate 

transect locations for avian point counts.
182 USDA Forest Service Proceedings RMRS-P-67. 2013

Sanderlin and others Preliminary Assessment of Species Richness and Avian Community Dynamics . . .

	 We used a data matrix y, where element yimht was a binary indicator

ofspecies detection.When yimht = 1,speciesi(i = 1,...,N) was detected

in mountain range m (m = 1,...,5)in covertype h (h = 1,..., 7) of year

t (t = 1,..., 5). We used the sum of binary species detection indicators

for each species × mountain range × cover type × year combination

over all three secondary periods during that combination (yimht = {0,

1, 2, 3}). We used species and effort (eff) to model species detection

and expected detection probability to increase with effort and to vary

among species. We used latitude (lat) and cover type (hab) to model

probability of occupancy of species. Since we condensed points by

cover type and mountain range, we were unable to use elevation as an

explanatory variable; however, there is a weak relationship between

covertype andelevation(table1).We expectedthatspeciesoccupancy

patterns would vary among cover types, and species richness would

decrease with increasing latitude due to loss ofspecies with Mexican

and Central American distributions. For numerical reasons, we used

normalized covariates for effort and latitude. We defined effort as

number of points in each mountain range × cover type combination

(table 1). Each sampling unit (n = 35) consists of points classified by

cover type within each mountain range. We used the mean latitude

from UTM coordinates of each mountain range as a covariate (Chiricahuas = 3533798, Huachucas = 3474145, Pinaleños = 3618746,

Santa Catalinas = 3589560, Santa Ritas = 3509745).

	 Sincewewere interested in thewhole avian community, quantification of both observed species and species we did not detect, but may

havebeenpresent,wereofinterest.Bayesianhierarchicalmodelswith

unknown species richness (Royle and Dorazio 2008: 384-387) that

rely on data augmentation (e.g., Dorazio and others 2006) allow usto

make inference on the entire community. We used a fixed supercommunity size M,so the posterior distribution ofΩ, probability a species

from the supercommunity was available forsampling during primary

sampling periods, was centered well below its upper limit (e.g., Ω ≤

0.5). We augmented the species observation matrix y with (M - nobs)

rows of zeros for all mountain range × cover type × primary period

sampling sessions. We modeled probability of latent variable wi of

species i being available for sampling during all primary sampling

periods given Ω as a Bernoulli trial:

To model community change, we assumed a Markovian process

described the current occupancy state of each species, whereby

probability of occupancy at time t was partially dependent on probability of occupancy at time t-1.We modeled probability of Bernoulli

latent variable zimht for occupancy given probability of occupancy

ψimht and wi as:

where occupancy probability ψihmt is a function of covariates and the

previous occupancy state:
USDA Forest Service Proceedings RMRS-P-67. 2013 183

Preliminary Assessment of Species Richness and Avian Community Dynamics . . . Sanderlin and others

Values ai1(0) and ai1 describe local probability of survival for species

i. These terms describe species-specific normal random effects: ai0

(intercept), ai3h (cover type), and ai2 (latitude).

	 We modeled probability of observation of species i, yihmt, given

the number of secondary periods J, probability of detection pihm and

latent variable zihmt, using a binomial distribution with J trials and

probability of success pihm×zihmt:

	 We used species and sampling effort (eff) as covariates to model

probability of detection p:

We modeled heterogeneity among species using a covariance term

between species intercepts of occurrence (ai0) and detection probability (bi0) (Royle and Dorazio 2008: p. 391), which allows us to

make inference on occupancy and detection probability of species

we never detected.

	 We used Bayesian hierarchical models (Gelman and others 2004)

in R2WinBUGS (Sturtz and others 2005), an R (R Development

Core Team 2011) package that interfaces with WinBUGS (Lunn and

others 2000) to obtain parameter estimates. We used independent

non-informative priors for parameters (uniform: Ω, ai1(0); Bernoulli:

zihm0; Normal: ai0, ai1, ai2, ai3h, bi0, b1; e.g., ). We ran four

parallel chains(length 35,000 it, burn-in period 20,000 it, thinning 10

it) to obtain median parameter estimates and 95% Bayesian Credible

Intervals (BCI). Convergence was reached (Ȓ = 1.0-1.1[Brooks and

Gelman 1998]).We assessed goodness-of-fit(GOF) using the squared

loss and deviance test statistics for Bayesian p-values (Gelman and

others 2004: p. 162).

Results

	 Over all mountain ranges, cover types, and years, we detected

158 bird species (table 2). Number of species observed (nobs) was

comparable (nobs,1991 = 119, nobs,1992 = 117, nobs,1993 = 107, nobs,1994 = 120,

nobs,1995 = 110) across years. There was evidence of overdispersion

(Bayesian p-value squared loss = 1.0, Bayesian p-value deviance =

0.66). The probability of a species being a supercommunity member

(median Ω = 0.506, 95% BCI: 0.408-0.68) indicates that our data

augmentation step was appropriate. Detection probabilities varied by

species (logit scale hyperprior medians: = 0.078, = 0.797)

and increased with increased sampling effort (median 1

ˆ

b = 0.671,

95% BCI: 0.627-0.714).

	 Average speciesrichnesswas differentfor eachmountainrange and

cover type, although 95% BCIs overlapped (fig. 2). For all mountain

ranges,speciesrichnesswasgreatestinbroadleaf evergreenwoodland

andconiferwoodlandcovertypes andleastindeciduousforest.Species

richness decreased as latitude increased in mixed-conifer, pine-oak

woodland, broadleaf evergreen woodland and conifer riparian cover

types. Deciduousforest, conifer woodland, and ponderosa pine cover

types had no association with species richness and latitude. Average

local extinction and colonization were similar for all cover types and

mountain ranges (fig. 3), except for Pinaleño Mountain ponderosa

pine ( : 0.205 [0.156, 0.256]; : 0.383 [0.282, 0.549]). Species

turnover, orthe process oflocal extinction and replacement ofspecies

through local colonization, was greatest within deciduous forest for

most mountain ranges and smallest within pine-oak forest, broadleaf

evergreen woodland, and conifer woodland cover types.

	 Fourteenspecieshadpositive slopeswithlatitude and18specieshad

negative slopes associated with latitude (95% BCIs for these species

did not cross zero; fig. 4). Some species had positive median cover

type slopes, although the 95% BCIs overlapped zero, for deciduous

forest (n = 3), mixed-conifer forest (n = 35), pine-oak forest (n = 25),

broadleaf evergreen woodland (n = 57), conifer woodland (n = 64),

conifer riparian (n = 6) and ponderosa pine (n = 17) (table 2).

Discussion

	 We identified associations between forest covertype and avian species:broadleaf evergreenwoodlandandconiferwoodlandcontributed

the most species. Deciduous forest cover type, which occurred at

higher elevations in these mountain ranges, had the greatest species

turnover, but fewest species.As expected, species richness increased

withdecreasedlatitudesinfourofthe sevencovertypes.Manyspecies

that occur in southerly latitudes have Mexican and CentralAmerican

distributions (e.g., elegant trogon, eared trogon, thick-billed parrot

[scientific names given in table 2]). Southernmost Arizona mountain

ranges encompassthe northern part of these speciesranges. These relationshipsinmixed-conifer,pine-oakwoodland,broadleaf evergreen

woodland and conifer riparian cover types may have implicationsfor

avian range shifts with climate change. Most climate models predict

warmer and drier future conditionsin the southwestern United States

(Seager andothers2007),thusspecieswhose ranges aremore southern

may move northward with warming climates, particularly if changes

create expanded habitat availabilities. Lower elevation cover types

may shift upwards and higher elevation cover types may shrink or

disappear, leading to decline or loss of species with strong habitat

associations.

	 Our results suggest not all species present were detected during

sampling anddetection probability increasedwithincreased sampling

effort. This highlights the importance of sample design and accounting for detection probabilities within occupancy models(MacKenzie

and others 2006). Detection probabilities are critical in studies that

assess changes over time or responsesto disturbance events, because

detection probability may change over time or with habitat changes.

Several species in our study had low detectability, which is known

to cause bias in occupancy estimates (MacKenzie and others 2006:

p. 107) and can lead to inflated estimates of species richness.

	 Our preliminary analyses suggest that cover type may not be the

best explanatoryvariable fortheprobabilityofoccupancydue tosome

evidence of overdispersion.Otherfactorslike elevation,microhabitat

structure, slope and aspect, mountain range area, and distance to

other mountains may have more explanatory power, which warrants

future exploration. Future analyses will focus on estimating relative

abundances(sensuRoyle andNichols2003),detailedhabitatrelationships of individual avian species, especially species of conservation

concern, and evaluation of resident and migratory species for their

relative contributions to species richness and community dynamics.

Thisinformation will be valuable to managers charged with conserving

this unique avifauna, and aid in assessing potential impacts of changing climates.

	 Our study also provides a baseline for future studies in this Global

Important Bird Area (Audubon Society). The Sky Islands have been

impacted by several large-scale, habitat altering disturbance events

in recent years (e.g., the 2011 Horseshoe Two fire burned 90,307 ha
184 USDA Forest Service Proceedings RMRS-P-67. 2013

Sanderlin and others Preliminary Assessment of Species Richness and Avian Community Dynamics . . .

Table 2—Avian species with positive posterior medians associated with cover types within the five Sky Island mountain 

ranges in southeastern Arizona, USA from 1991 to 1995. The number ‘1’ indicates a positive posterior median for 

the specific habitat type. Some species did not have any positive associations and some species were associated 

with more than one cover type.

Common species name Scientific name

Dec. 

for.

Mix.-

conf. 

for.

Pineoak 

for.

Brd. 

Evgr. 

wd.

Conf. 

wd.

Conf. 

ripa.

Pond. 

pine

Mallard Anas platyrhynchos 0 0 0 0 1 0 0

Gambel’s quail Callipepla gambelii 0 0 0 1 1 0 0

Scaled quail Callipepla squamata 0 0 0 0 0 0 0

Montezuma quail Cyrtonyx montezumae 0 0 0 1 1 0 0

Wild turkey Meleagris gallopavo 0 0 0 0 0 0 0

Turkey vulture Cathartes aura 0 0 0 1 1 0 0

Cooper’s hawk Accipiter cooperii 0 0 0 0 0 0 0

Northern goshawk Accipiter gentilis 0 1 0 0 0 0 1

Sharp-shinned hawk Accipiter striatus 0 0 0 0 1 0 0

Gray hawk Buteo nitidus 0 0 0 0 0 0 0

Zone-tailed hawk Buteo albonotatus 0 0 0 0 0 0 1

Red-tailed hawk Buteo jamaicensis 0 0 0 1 0 0 0

Common black-hawk Buteogallus anthracinus 0 0 0 0 0 0 1

Golden eagle Aquila chrysaetos 0 0 0 0 0 0 0

Prairie falcon Falco mexicanus 0 0 0 0 1 0 0

American kestrel Falco sparverius 0 0 0 0 0 0 0

Band-tailed pigeon Patagioenas fasciata 0 0 1 0 0 0 0

Rock dove Columba livia 0 0 0 0 0 0 1

White-winged dove Zenaida asiatica 0 0 0 1 1 0 0

Mourning dove Zenaida macroura 0 0 0 1 1 0 0

Thick-billed parrot Rhynchopsitta 

pachyrhyncha 0 1 0 0 0 0 0

Greater roadrunner Geococcyx californianus 0 0 0 1 1 0 0

Great horned owl Bubo virginianus 0 0 0 0 0 0 0

Northern pygmy-owl Glaucidium gnoma 0 0 1 0 0 0 0

Flammulated owl Otus flammeolus 0 1 0 0 0 0 0

Whiskered screech-owl Megascops trichopsis 0 0 1 0 0 0 0

Spotted owl Strix occidentalis 0 1 0 0 0 0 0

Lesser nighthawk Chordeiles acutipennis 0 0 0 0 1 0 0

Eastern whip-poor-will Caprimulgus vociferus 0 0 0 1 0 0 1

White-throated swift Aeronautes saxatalis 0 1 0 0 0 1 0

Black-chinned hummingbird Archilochus alexandri 0 0 1 1 1 0 0

Anna’s hummingbird Calypte anna 0 0 0 1 0 0 0

Costa’s hummingbird Calypte costae 0 0 0 0 0 0 0

Broad-billed hummingbird Cynanthus latirostris 0 0 0 0 1 1 0

Magnificent hummingbird Eugenes fulgens 0 1 0 0 0 0 0

(continued)

within the Chiricahua Mountains) and these large-scale disturbance

events may become more frequent as climate in the southwestern

United States becomes warmer and drier (McKenzie and others

2004). Consequent changesin habitatstructure could influence avian

community dynamics in these ranges. We know little about effects

of fire and other disturbances on Neotropical migratory birds in this

region (Ganey and others 1996; Kirkpatrick and others 2006), thus,

futuremonitoringaimedat evaluatingeffectsoffire events andclimate

shifts on avian species in the region are paramount.
USDA Forest Service Proceedings RMRS-P-67. 2013 185

Preliminary Assessment of Species Richness and Avian Community Dynamics . . . Sanderlin and others

Table 2—Continued

Common species name Scientific name

Dec. 

for.

Mix.-

conf. 

for.

Pineoak 

for.

Brd. 

Evgr. 

wd.

Conf. 

wd.

Conf. 

ripa.

Pond. 

pine

Blue-throated hummingbird Lampornis clemenciae 0 0 0 0 0 1 0

Broad-tailed hummingbird Selasphorus platycercus 0 1 0 0 0 0 0

Rufous hummingbird Selasphorus rufus 0 0 0 0 0 0 0

Eared quetzal Euptilotis neoxenus 0 0 0 0 0 0 0

Elegant trogon Trogon elegans 0 0 1 0 0 1 0

Northern flicker Colaptes auratus 0 0 0 0 0 0 0

Acorn woodpecker Melanerpes formicivorus 0 0 1 1 1 0 0

Gila woodpecker Melanerpes uropygialis 0 0 0 1 0 0 0

Ladder-backed woodpecker Picoides scalaris 0 0 0 1 1 0 0

Strickland’s woodpecker Picoides stricklandii 0 0 0 1 1 0 0

Hairy woodpecker Picoides villosus 0 1 1 0 0 0 1

Red-naped sapsucker Sphyrapicus nuchalis 0 1 0 0 0 0 0

Northern beardless-tyrannulet Camptostoma imberbe 0 0 0 1 1 0 0

Black phoebe Sayornis nigricans 0 0 0 0 1 0 0

Say’s phoebe Sayornis saya 0 0 0 1 0 0 0

Olive-sided flycatcher Contopus cooperi 0 0 0 0 0 0 0

Greater pewee Contopus pertinax 0 1 1 0 0 0 1

Western wood-pewee Contopus sordidulus 0 0 1 0 0 0 0

Pacific-slope flycatcher Empidonax difficilis 0 1 0 0 0 0 0

Buff-breasted flycatcher Empidonax fulvifrons 0 1 0 0 0 0 0

Hammond’s flycatcher Empidonax hammondii 0 1 0 0 0 0 0

Dusky flycatcher Empidonax oberholseri 0 0 0 0 1 0 0

Cordilleran flycatcher Empidonax occidentalis 0 1 0 0 0 0 0

Gray flycatcher Empidonax wrightii 0 0 0 1 0 0 0

Dusky-capped flycatcher Myiarchus tuberculifer 0 0 0 0 1 0 0

Ash-throated flycatcher Myiarchus cinerascens 0 0 0 1 1 0 0

Brown-crested flycatcher Myiarchus tyrannulus 0 0 0 0 1 0 0

Sulphur-bellied flycatcher Myiodynastes luteiventris 0 0 1 0 0 0 0

Thick-billed kingbird Tyrannus crassirostris 0 0 0 0 1 0 0

Western kingbird Tyrannus verticalis 0 0 0 0 0 0 0

Cassin’s kingbird Tyrannus vociferans 0 0 0 1 1 0 0

Rose-throated becard Pachyramphus aglaiae 0 0 0 1 0 0 0

Loggerhead shrike Lanius ludovicianus 0 0 0 1 0 0 0

Bell’s vireo Vireo bellii 0 0 0 0 1 0 0

Warbling vireo Vireo gilvus 0 1 1 0 0 0 0

Hutton’s vireo Vireo huttoni 0 0 1 1 0 0 0

Plumbeous vireo Vireo plumbeus 0 0 1 0 0 0 0

Gray vireo Vireo vicinior 0 0 0 0 1 0 0

Western scrub-jay Aphelocoma californica 0 0 0 1 1 0 0

Stellar’s jay Cyanocitta stelleri 0 0 1 0 0 0 0

Gray jay Perisoreus canadensis 0 0 1 1 1 0 0

Common raven Corvus corax 0 0 0 1 0 0 0

Chihuahan raven Corvus cryptoleucus 0 0 0 0 0 0 0

Tree swallow Tachycineta bicolor 0 1 0 0 0 0 0

(continued)
186 USDA Forest Service Proceedings RMRS-P-67. 2013

Sanderlin and others Preliminary Assessment of Species Richness and Avian Community Dynamics . . .

Table 2—Continued

Common species name Scientific name

Dec. 

for.

Mix.-

conf. 

for.

Pineoak 

for.

Brd. 

Evgr. 

wd.

Conf. 

wd.

Conf. 

ripa.

Pond. 

pine

Violet-green swallow Tachycineta thalassina 0 1 0 0 0 0 0

Mountain chickadee Poecile gambeli 0 0 0 0 0 0 0

Mexican chickadee Poecile sclateri 0 0 0 0 0 0 0

Juniper titmouse Baeoluphus ridgwayi 0 0 0 1 1 0 0

Bridled titmouse Baeoluphus wollweberi 0 0 0 1 1 0 0

Verdin Auriparus flaviceps 0 0 0 0 1 0 0

Bushtit Psaltriparus minimus 0 0 0 1 1 0 0

Red-breasted nuthatch Sitta canadensis 0 1 0 0 0 0 0

White-breasted nuthatch Sitta carolinensis 0 0 0 0 0 0 0

Pygmy nuthatch Sitta pygmaea 0 1 0 0 0 0 1

Brown creeper Certhia americana 0 1 1 0 0 0 1

Canyon wren Catherpes mexicanus 0 0 0 1 1 1 0

Rock wren Salpinctes obsoletus 0 0 0 1 1 0 0

Bewick’s wren Thryomanes bewickii 0 0 0 1 1 0 0

House wren Troglodytes aedon 0 1 0 0 0 0 0

Winter wren Troglodytes hiemalis 0 0 0 0 0 0 1

Blue-gray gnatcatcher Polioptila caerulea 0 0 0 1 1 0 0

Black-tailed gnatcatcher Polioptila melanura 0 0 0 1 0 0 0

Ruby-crowned kinglet Regulus calendula 0 0 0 1 1 0 0

Golden-crowned kinglet Regulus satrapa 0 1 0 0 0 0 0

Mountain bluebird Sialia currucoides 0 0 0 0 0 0 1

Western bluebird Sialia mexicana 0 0 0 0 0 0 0

Eastern bluebird Sialia sialis 0 1 0 0 0 0 0

Townsend’s solitaire Myadestes townsendi 0 0 0 0 0 0 1

Hermit thrush Catharus guttatus 0 1 1 0 0 0 0

Swainson’s thrush Catharus ustulatus 0 1 0 0 0 0 0

American robin Turdus migratorius 0 0 1 0 0 0 0

Northern mockingbird Mimus polyglottos 0 0 0 1 1 0 0

Bendire’s thrasher Toxostoma bendirei 0 0 0 0 0 0 0

Crissal thrasher Toxostoma crissale 0 0 0 1 1 0 0

Curve-billed thrasher Toxostoma curvirostre 0 0 0 1 1 0 0

Cedar waxwing Bombycilla cedrorum 0 0 0 1 1 0 0

Phainopepla Phainopepla nitens 0 0 0 1 0 1 0

Olive warbler Peucedramus taeniatus 0 1 0 0 0 0 1

Red-faced warbler Cardellina rubrifrons 0 1 1 0 0 0 0

Yellow-rumped warbler Setophaga coronata 0 1 1 0 0 0 0

Grace’s warbler Setophaga graciae 0 1 1 0 0 0 0

Black-throated gray warbler Setophaga nigrescens 0 0 1 1 1 0 0

Hermit warbler Setophaga occidentalis 0 1 0 0 0 0 0

Townsend’s warbler Setophaga townsendi 0 0 0 1 0 0 1

Painted redstart Myioborus picta 0 0 1 0 0 0 0

MacGillivray’s warbler Geothlypsis tolmiei 1 0 0 0 0 0 0

Orange-crowned warbler Oreothlypsis celata 1 1 0 0 0 0 0

Lucy’s warbler Oreothlypsis luciae 0 0 0 0 1 0 0

(continued)
USDA Forest Service Proceedings RMRS-P-67. 2013 187

Preliminary Assessment of Species Richness and Avian Community Dynamics . . . Sanderlin and others

Table 2—Continued

Nashville warbler Oreothlypsis ruficapilla 0 0 0 0 1 0 0

Virginia’s warbler Oreothlypsis virginiae 0 0 1 0 0 0 0

Wilson’s warbler Cardellina pusilla 0 0 0 1 0 0 0

Abert’s towhee Melozone aberti 0 0 0 0 1 0 0

Green-tailed towhee Pipilo chlorurus 0 0 0 1 1 0 0

Canyon towhee Melozone fusca 0 0 0 1 1 0 0

Spotted towhee Pipilo maculatus 0 0 0 0 0 0 0

Rufous-crowned sparrow Aimophila ruficeps 0 0 0 1 1 0 0

Black-throated sparrow Amphispiza bilineata 0 0 0 1 1 0 0

Lark sparrow Chondestes grammacus 0 0 0 0 0 0 0

Black-chinned sparrow Spizella atrogularis 0 0 0 1 1 0 0

Chipping sparrow Spizella passerina 0 0 0 1 1 0 0

White-crowned sparrow Zonotrichia leucophrys 0 0 0 0 1 0 0

Dark-eyed junco Junco hyemalis 0 0 0 1 1 0 0

Yellow-eyed junco Junco phaeonotus 0 1 1 0 0 0 0

Hepatic tanager Piranga flava 0 0 0 1 1 0 0

Western tanager Piranga ludoviciana 0 0 0 0 0 0 0

Summer tanager Piranga rubra 0 0 0 0 1 0 0

Northern cardinal Cardinalis cardinalis 0 0 0 1 1 0 0

Pyrrhuloxia Cardinalis sinuatus 0 0 0 0 1 0 0

Black-headed grosbeak Pheucticus melanocephalus 0 0 0 0 0 0 0

Blue grosbeak Passerina caerulea 0 0 0 1 0 0 0

Lazuli bunting Passerina amoena 0 0 0 0 1 0 0

Eastern meadowlark Sturnella magna 0 0 0 0 0 0 0

Western meadowlark Sturnella neglecta 0 0 0 0 1 0 0

Brown-headed cowbird Molothrus ater 0 0 0 1 1 0 0

Hooded oriole Icterus cucullatus 0 0 0 0 1 0 0

Bullock’s oriole Icterus bullockii 0 0 0 1 1 0 0

Scott’s oriole Icterus parisorum 0 0 0 1 1 0 0

Pine siskin Spinus pinus 0 1 0 0 0 0 1

Lesser goldfinch Spinus psaltria 0 0 0 1 1 0 0

Cassin’s finch Carpodacus cassinii 0 1 0 0 0 0 0

House finch Carpodacus mexicanus 0 0 0 1 1 0 0

Evening grosbeak Coccothraustes vespertinus 1 1 0 0 0 0 0

Red crossbill Loxia curvirostra 0 0 0 0 0 0 1

Common species name Scientific name

Dec. 

for.

Mix.-

conf. 

for.

Pineoak 

for.

Brd. 

Evgr. 

wd.

Conf. 

wd.

Conf. 

ripa.

Pond. 

pine
188 USDA Forest Service Proceedings RMRS-P-67. 2013

Sanderlin and others Preliminary Assessment of Species Richness and Avian Community Dynamics . . .

Figure 2—Median posterior estimates of average 

species richness (SR) and 95% Bayesian credible 

intervals over all years from 1991 to 1995 for the 

five Sky Island mountain ranges of southeastern 

Arizona, USA. Estimates are separated by cover 

type (1 = deciduous forest, 2 = mixed-conifer forest, 3 = pine-oak forest, 4 = broadleaf evergreen 

woodland, 5 = conifer woodland, 6 = conifer 

riparian, 7 = ponderosa pine).

Figure 3—Median posterior estimates of local extinction and local colonization and 95% Bayesian credible 

intervals over all years from 1991 to 1995 for the five 

Sky Island mountain ranges of southeastern Arizona, 

USA. Estimates are separated by cover type (1 = deciduous forest, 2 = mixed-conifer forest, 3 = pine-oak 

forest, 4 = broadleaf evergreen woodland, 5 = conifer 

woodland, 6 = conifer riparian, 7= ponderosa pine).
USDA Forest Service Proceedings RMRS-P-67. 2013 189

Preliminary Assessment of Species Richness and Avian Community Dynamics . . . Sanderlin and others

Figure 4—Significant median posterior estimates for the latitude slope parameter (âlatitude) and 95% Bayesian credible intervals of individual avian 

species from a study conducted from 1991 to 1995 in five Sky Island mountain ranges of southeastern Arizona, USA. Positive associations 

indicate that the probability of occupancy for those species increases with increasing latitude, while negative associations indicate probability 

of occupancy decreases with increasing latitude. Scientific names for these species are given within Table 2. Species are ordered according to 

taxon or family group (letters indicate separate groups).

Acknowledgments

	 We thank field crews for collecting avian and vegetation data; The

Coronado National Forest, The Nature Conservancy, Fort Huachuca,

Santa Rita Experimental Station, and Audubon Research Ranch for

logistics/technical assistance; National Fire Plan for funding (JSS);

L. Hall, Q. Latif, and D. Turner for helpful comments during review.

References

Balda,R.P.1967.EcologicalrelationshipsofthebreedingbirdsoftheChiricahua

Mountains, Arizona. Ph.D. Dissertation, University of Illinois, Urbana.

Block, W.M.; Ganey, J.L.; Severson, K.E.; and Morrison, M.L. 1992. Use of

oaksbyNeotropicalmigratorybirdsinthe southwest.Pp.65-70in: Ffolliott,

P.F.; Gottfried, G.J., Bennett, D.A.; Hernandez, C.; Ortega-Rubio, A.; and

Hamre, R.H., tech cord. Ecology and management of oak and associated

woodlands: perspectives in the southwestern United States and northern

Mexico. Gen. Tech. Rep. RM-218. U.S. Department of Agriculture, For-
190 USDA Forest Service Proceedings RMRS-P-67. 2013

Sanderlin and others Preliminary Assessment of Species Richness and Avian Community Dynamics . . .

est Service, Rocky Mountain Forest and Range Experiment Station. Fort

Collins, CO.

Block, W.M.; and Ganey, J.L. 1992. Study Plan Number RM-4251-3-12:

Habitat associations of Neotropical migrant birds in the insular mountains

ofsoutheasternArizona. Rocky MountainResearch Station. Flagstaff,AZ.

Block, W.M.; and Severson, K.E. 1992. Habitat associations of birds and

herpetofauna in southeasternArizona. Pp. 55-57 in: Chiricahua Mountains

Research Symposium Proceedings; 1992 16-17 March; Chiricahua Mountains, AZ; Southwestern Parks and Monuments Association.

Brooks, S.B.; and Gelman, A. 1998. General methods for monitoring convergence of iterative simulations. Journal of Computational and Graphical

Statistics 7:434-455.

Cochran, W.G. 1977. Sampling techniques, Third edition. Wiley, New York.

Dorazio, R.M.; Royle,J.A.; Söderström, B.; and Glimskär,A. 2006. Estimating speciesrichness and accumulation by modeling species occurrence and

detectability. Ecology 87:842-854.

Ganey, J.L.; Block, W.M.; and Boucher, P.F. 1996. Effects of fire on birds in

Madrean forests and woodlands. Pp. 146-154 in: Ffolliott, P.F.; DeBano,

L.F.;Baker,M.B.;[andothers],techcoords.Gen.Tech.Rep.RM-GTR-289.

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest

and Range Experiment Station. Fort Collins, CO.

Gelman, A.; Carlin, J.B.; Stern, H.S.; and Rubin, D.B. 2004. Bayesian data

analysis, 2nd edition. Chapman and Hall/CRC, New York.

Hall,L.S.;andMannan,R.W. 1999.Multiscaledhabitatselectionbyeleganttrogonsin southeasternArizona.Journal ofWildlifeManagement 63:451-461.

Iniguez,J.M.;Ganey,J.L.;Dougherty,P.J.; andBailey,J.D.2005.Usingcluster

analysis and a classification and regression tree model to identify cover

types in the sky islands of southeastern Arizona. Pp.195-200 in: Gottfried,

G.J.; Gebow, B.S.; Eskew, L.G.; and Edminster, C.B., comps. Connecting

mountain islands and desert seas: biodiversity and management of the

Madrean Archipelago II. Proc. RMRS-P-36. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. Fort Collins, CO.

Kirkpatrick, C.; Conway, C.J.; and Jones, P.B. 2006. Distribution and relative abundance of forest birds in relation to burn severity in southeastern

Arizona. Journal of Wildlife Management 70:1005-1012.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Latta, M.J.; Beardmore, C.J.; and Corman, T.E. 1999. Arizona Partners in

Flight Bird Conservation Plan. Version 1.0. Nongame and Endangered

Wildlife Program Technical Report 142. Arizona Game and Fish Department, Phoenix, AZ.

Lunn, D.J.; Thomas, A.; Best, N.; and Spiegelhalter, D. 2000. WinBUGS – a

Bayesian modeling framework: concepts, structure, and extensibility.

Statistics and Computing 10:325-337.

MacKenzie, D.I.; Nichols, J.D.; Royle, J.A.; Pollock, K.H.; Bailey, L.L.;

and Hines, J.E. 2006. Occupancy estimation and modeling. Academic

Press, New York.

Marshall, J.T.Jr. 1957. Birds of the pine-oak woodland in southern Arizona

and adjacent Mexico. Pacific Coast Avifauna 32.

McKenzie,D.;Gedalof,Z.;Peterson,D.L.; andMote,P.2004.Climate change,

wildfire, and conservation. Conservation Biology 18:890-902.

Pollock, K.H. 1982. A capture-recapture design robust to unequal probability

of capture. Journal of Wildlife Management 46:757-760.

R Development Core Team. 2011. R: a language and environment forstatistical computing. R Foundation for Statistical Computing, Vienna, Austria.

ISBN 3-900051-07-0, URL http://www.R-project.org/.

Reynolds, R.T.; Scott, J.M.; and Nussbaum, R.A. 1980. A variable circular

plot method for estimating bird numbers. Condor 82:309-313.

Royle, J.A.; and Nichols, J.D. 2003. Estimating abundance from repeated

presence absence data or point counts. Ecology 84:777-790.

Royle, J.A.; and Dorazio, R.M. 2008. Hierarchical modeling and inference

in ecology. Academic Press, New York.

Seager, R.; and others 2007. Model projections of an imminent transition to a

more arid climate in southwestern NorthAmerica. Science 316:1181-1184.

Strong,T.R.; andBock,C.E.1990.Birdspeciesdistribution patternsinriparian

habitats in southeastern Arizona. Condor 866-885.

Sturtz, S.; Ligges, U.; and Gelman A. 2005. R2WinBUGS: A package for

running WinBUGS from R. Journal of Statistical Software 12:1-16.

Williams, B.K.; Nichols, J.D.; and Conroy, M.J. 2002. Estimation of community parameters. Pp. 555-573 in Analysis and management of animal

populations. Academic Press, San Diego.
USDA Forest Service Proceedings RMRS-P-67. 2013 191

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Biodiversity Effects on Ecosystem Function Due to Land 

Use: The Case of Buffel Savannas in the Sky Islands Seas in 

the Central Region of Sonora

A. E. Castellanos, H. Celaya, C. Hinojo, and A. Ibarra 

DICTUS, Universidad de Sonora, Hermosillo, Sonora

J. R. Romo 

School of Natural Resources and Environment, University of Arizona, Tucson, Arizona

Abstract—Buffel savannas have been an important landscape on cattle grazing ranches in Sonora over the past 

50 years or more. Changes in land use result in biodiversity changes that may produce ecosystem functional 

changes; however, these are less well documented. Although fire driven processes have been proposed for 

Buffel savannas, this is not generally the case, and other processes seem to be driving ecosystem function. 

Several years of studying above- and below ground processes allow us to propose how microclimate, water 

and nutrient dynamics change in established Buffel savannas, as well as how biodiversity changes may affect 

functional processes in arid and semiarid ecosystems. Water and nutrient biogeochemical cycles changed 

in Buffel dominated savannas in comparison with those in natural ecosystems, following land use changes. 

Our findings may be extrapolated to other highly invaded areas were Buffel grass is becoming a dominant 

exotic species.

Keywords: biodiversity effects on ecosystem function, Buffel savanna, desert scrub, Sonoran Desert, soil 

microclimate, soil moisture.

Land Use in Arid Sonoran Ecosystems

	 The Sonoran Desert in the States of Sonora,BajaCalifornia (North

and South), Arizona and California covers close to 300,000 Km-2. It

hasimportantgradientsofincreasingplantspeciesdiversityfromnorth

(Larrea dominated ecosystems) to south in México and from west

to east (with increasing tropical floristic influence), which results in

a large plant functional diversity and biological forms (Peinado and

others1990;Shreve1942).Those changesintaxonomic andfunctional

biodiversity, following phytogeographic and aridity gradients, make

the Central Sonora Region (CSR) (broadly limited to 28 - 30°N and

102 - 106°W), the ecotone for those two major gradients, and thus, a

particularlyimportant area for ecological andbiogeographicalstudies.

The biological importance of the region is because it is the northern

most limit to a large number of neotropical species, as well as the

southernmost limit to many neartic species (Rzedowski 1978). It is

important as well because it is an arid ecotone where large speciation

events (Stebbins 1952), high endemism (Shreve and others 1964;

Turner and others 1995) and ecological legacies (Castellanos 1992;

Castellanos and others 2010) are found.

	 Land use and cover change have been rapid and greatly modified

by cattle ranching in the last half century. Cattle grazing was introduced into Northwestern Mexico and the Central Sonora Region

(CSR) during the late 1600’s (Camou 1998; Castellanos and others

2010). Large herds of cattle have been present for over 400 years

in the CSR, diminishing palatable forage plant species, increasing

shrubland and degrading habitat (Aguirre-Murrieta and others 1974;

COTECOCA2002). Because of its aridity and dryness, Sonora grazing lands are known to support as few as one animal unit per 20 - 30

hectares (Aguirre-Murrieta and others 1974). However, increasing

opportunities for cattle export to the United States in the 1950’s led

to a change in the ranching technological paradigm in Sonora (BravoPeña and others 2010; Camou 1998; Castellanos and others 2010).

Socioeconomic drivers, such as increasing cattle market demands,

led to overstocking and increasing habitat degradation, which led

to increasing exotic buffel (Pennisetum ciliare L.) grasslands being

established in Sonora, following research from TexasAM University

(Johnson and Murrieta 1992). In order to increase land productivity

to support market needs, extensive tracts of land were converted to

intensive monocultures of P. ciliare.

	 Overthe last 40-50 yearsin Sonora, a common practice has been to

chain and bulldoze large tracts of terrain, clearing (in early years all,

but more recently) most shrubs and trees, seeding buffel grass and

rotatingcattle seasonally.Despite the apparentproductive advantages,

thismassive disturbance has posed a number of ecological challenges

over the years, not only because it decreases species richness and

plant diversity (Castellanos and others 2002; Saucedo-Monarque

1994; Saucedo-Monarque and others 1997), increases the potential

forsoil erosion(Perramond2000;Saucedo-Monarque1994), changes
192 USDA Forest Service Proceedings RMRS-P-67. 2013

Castellanos and others Biodiversity Effects on Ecosystem Function Due to Land Use: . . .

microclimatic soil conditions(Castellanos and others 2002); but also

because it decreases important soil nitrogen (Castellanos and others

2002; Dalal and others 2005; Ibarra-Flores and others 1999), and

changes nutrient dynamics (Castellanos and others 2010; CelayaMichel and others 2011; López 2007).

	 Shrubland conversion to buffelsavannas has changed biodiversity

composition and ecosystem structure in such a way that no detailed

knowledge exists on how those highly modified ecosystems will

function, remain, change or affect different ecosystem processes,

as well as biotic and trophic links. Buffel savannas dominated by a

highly invasive grass species are an important part of the landscape

in Sonora, but their ecological effects are unknown (Arriaga and others 2004). We believe that a better understanding of how introduced

buffel savannas functions is needed and can help to understand the

short and long-term ecological implications of this management

policy, such as to understand some of the main causes for its spreading, as well as to help us establish better management practices to

control its spreading into native adjacent ecosystems. Our approach

also provides some badly needed basic knowledge on how Sonoran

Desert and arid ecosystems function.

	 Over the last 20 years we have studied paired sitesin different arid

ecosystemsintheCentralSonoraRegion.Pairedsites(buffelsavannas

and “native” vegetation cover) have been studied in order to answer

some important questions in relation to the role of biodiversity in

ecosystem function:

•	 How are microclimate, soil moisture, and fertility dynamics

modified in native compared to buffel savannas?

•	 What are the changes in ecosystem nitrogen dynamics and how

do they relate to changes in structure and species composition?

•	 How are ecosystem degradation and resilience changed in a

nearly monospecific-dominated ecosystem of buffel savannas?

Biodiversity Change and Ecosystem 

Function

Biodiversity is an important component of ecosystems and its

qualitative and quantitative changes have been found to affect its

function, an important consideration with increasing land cover use

andchange (Hooper andothers2005;Naeemandothers2002;Schulze

and others 1994). A well- established paradigm relates biodiversity

with net primary productivity (Tilman 1988;Tilman and others 2002;

Tilman and others 1997; Wright and others 2006) and ecosystem

stability (Tilman and others 2002; Tilman and others 2006).

Even though buffelsavannas have been proposed as a management

alternative to increase land net primary productivity, some studies

question that such a goal has been achieved. Using NDVI, as a proxy

to measure vegetation productivity, hasshown that plant productivity

is much lower than proposed or expected in a range of conditions

alonga rainfallgradient(Bravo-Peña2009;Romo2006),intheCentral

Region of Sonora and a region north of it (Franklin and others 2006).

	 Questionsremainonhowchangesfromnatural ecosystemstobuffel

savannas affect biodiversity and biotic links at different levels, how

and what ecosystem processes change, and how stability of these

induced and simplified ecosystems is modified.

Structural Changes in Plant Cover

	 Plant removal for buffel grass establishment changed plant structure and cover (fig.1). While the herbaceous stratum is significantly

increased by buffel cover in the savanna, total plant cover diminishes

compared to native vegetation stands. It has been found (Romo 2006;

Lopez 2007)that cover could further diminish aftersome 30-40 years

ofsavanna establishmenttoabouthalf comparedtonaturalvegetation.

	 We have proposed that a “green rush” effect drives cattle ranchers

to adopt buffelsavanna establishment (Bravo and others 2010a). The

effect happens afterthe dry season andwith thefirstsummermonsoon

rains. Once buffel has been established, because it is a fast growing

herbaceous perennialspecies,it hasthe ability to immediately leaf-out

afterthe very firstrainfall pulses, producing an immediate “greeness”

given that the whole savanna biomassis biologically, physically, and

visually concentrated in only one vegetation stratum. Meanwhile,

most neighboring plant species within native woody vegetation will

green out later, resulting in large patches of brown and green because

the greeness is distributed multi-dimensionally over the landscape.

Soil Abiotic Controls

Soil Temperature and Moisture

	 Temperature and moisture are important drivers of ecosystem

processes in the soil. Mean annual soil temperatures at 10 cm were

not different at different sites of buffel cover, but their temperature

dynamicsweredifferent(fig.2).As expected,maximumandminimum

soil temperatures were found in bare areas (ISN and ISB), which

showed the largest soil temperature fluctuations in a given time.

	 Vegetation cover dampssoil temperature in fall and winter months

and during most of the year its coefficient of variation wasthe lowest.

Higher minimum soil temperatures were found under O. tesota trees

and under buffel grass. Mean soil temperatures were higher under

trees and lowest in the savanna because of large temperature fluctuations during the day. Buffel microsites had the lowest soil maximum

temperatures during fall and winter but not in summer when PFB had

about 15 - 20 °C lower temperatures compared to similar microsites

in natural vegetation (PFN).

	 Largest daily and seasonal variations in soil temperatures in bare

and buffel areas may have important implications to establishment

of successional native species (Morales-Romero and others 2008;

Saucedo-Monarque 1994).

	 Soil maximum (Tsmax) and average (Taav) temperatures as well

as minimum (Tamin) and average (Taav) air temperatures were

significantly related to soil respiration. Highest significances were

found between soil respiration and Tsmax. Soil respiration was also

directly related to volumetric soil moisture at all times during the

Figure 1—Changes in plant cover in Buffel savannas of different age of 

establishment. Old (≥30 yrs), Medium (10 - 30 yrs) and Recent (0 - 10 

yrs). Natural is the predominant native vegetation at the site (modified 

from Romo (2006) and López (2007)).
USDA Forest Service Proceedings RMRS-P-67. 2013 193

Biodiversity Effects on Ecosystem Function Due to Land Use: . . . Castellanos and others

Figure 2—Mean soil temperature at different microsites during the year. PFN = under ironwood (O. 

tesota) trees in natural vegetation, PFB = similar but 

within the Buffel savanna; ISN = interspace (bare areas between trees, shrubs or herbaceous perennials) 

in natural vegetation, ISB = similar but within Buffel 

savannas; BB = under the canopy of Buffel grass.

year, although this relationship decreased with depth to 50 cm and

deeper.

	 Soil moisture changed with site (table 1; fig. 3), year, and time of

the year. Soil moisture in the buffel savanna was significantly lower

because lower soil moisture in the interspaces and under O. tesota

trees, compared to similar habitats in natural ecosystems. Soil moisture under ironwood in the savanna (PFB) had the lowest mean soil

moisture (2.60 ± 2.2 g H2O g-1 soil p<0.01) but not significantly

lower than the one found for the same microsite in natural vegetation

(PFN, 4.15 ± 4.9 g H2O g-1 soil). Although not expected, because

of late summer and early fall rainfalls during the last 2 years, soil

moisture was higher at depths below 100 cm in sites with increasing

aboveground leaf cover, including fewer than 100 cm in buffel grass.

This is below depths of usable soil moisture (close to 60 - 70 cm in

buffel), and depths of 130 - 150 cm below O. tesota canopies.

Soil Biotic Interactions

Termite Activity

	 Termite diversity was affected by changesin plant cover and structure in the natural vegetation and buffelsavanna. In both ecosystems,

termite richness did not change and the same six species were found

in both ecosystemsthroughout the year (fig. 4). Of the speciesfound,

Gnathamitermes perplexus (41%) and Heterotermes aureus (33%)

were the two most frequent termite species. Other species were

Amitermes spp, Hoplotermes spp and Tenuirostritermes spp as well

as an unidentified Amitermitinae species.

	 Small numbers of plots yielded small numbers of termite species.

Expected termite richness ("Mao Tau" computed by the EstimateS®

program using pooled plot data) was almost 25% higher for natural

systems(8 spp) comparedwith buffelsavanna (Sobs infig. 4). "Chao2,"

another EstimateS richness output that assumes a different distribution of sampling error, gave similar results, with highest values for

termite species richness during September on natural sites. During

winter months (not shown on fig. 4), richness was lowest in natural

systems. Chao2 curves confirm that the sampling effort on study sites

was appropriate (fig. 4).

	 Termite presence, seasonal activity, frequencies and trophic substrates did change between natural and buffel savanna ecosystems.

Seasonal presence of termite species changed in both ecosystems.

buffel savannas termites were most active during autumn-winter

seasons,whilemajorseasonal activity in natural ecosystems occurred

at the end ofsummer-early fallseason under O. tesota trees. Changes

were alsofoundinthe interspaces—bare zonesbetweenspecies—with

most important termite activity at the end of winter-spring seasons,

while under buffel in the Savanna, activity periods were at the end of

summer rainy season and during winter months. In bare unprotected

zones,diversitywaslower andmajor activitywasfoundduringspring.

	 An importantfindingwasthatGnathamitermes perplexus,themost

common species in the buffel savanna, was found foraging mostly

Table 1—Changes in water availability at different depths in native vegetation and Buffel savanna 

sites. Volumetric water percentage obtained with TRIME® access tubes.

Natural SABANA

Mean±s.e. Mean±s.e. N F P

0 to 50 cm 3.38±0.28a 33.25±.25a 30 0.1251 0.072*

0 to 100 cm 8.22±0.52a 5.78±.47b 30 12.140 0.0007**

0 to 150 cm 11.56±0.75a 7.65±.68b 30 14.8735 0.0002**
194 USDA Forest Service Proceedings RMRS-P-67. 2013

Castellanos and others Biodiversity Effects on Ecosystem Function Due to Land Use: . . .

Figure 4—Expected termite richness in natural and Buffel savanna sites using Estimates®.

Figure 3—Changes in available soil moisture (amount of water in cm, in a profile), for different microsites in natural and Buffel savanna sites.
USDA Forest Service Proceedings RMRS-P-67. 2013 195

Biodiversity Effects on Ecosystem Function Due to Land Use: . . . Castellanos and others

on this grass species, behaving as an opportunistic species, changing

its trophic substrate to the more abundant buffel litter. In a different

study under Sonoran Desert conditions in Arizona, higher activity

of Gnathamitermes perplexus was found in the range of 9 to 49 °C,

while Heterotermes aureus, had a slightly lower, 7 to 47 °C, range

(table 2; Haverty 2001).

Discussion

	 Chambers and others(2007)mention the hypothesis of fluctuating

resources and say that invasibility is related to the availability of

resources.In particular, invasibility increasesifincreasing resources

are not utilized by natives. The availability of resources to buffel

grass will be related to (a) an increase in supply reliability or

(b) a decrease in uptake by natives. Many studies relate increasesd

availability of nitrogen with increased invasibility.

Acknowledgments

	 We thank CONACYT for its support to project (CB66,,,,) and

as scholarships (A.I., C.H., H.C., J.R.R.) during these studies. We

thank Mr. Luis Sierra for allowing us to perform our research on his

property.

References

Aguirre-Murrieta R et al. (1974) Coeficientes de agostadero para el estado de

Sonora. In. COTECOCA - SAG, México, p 133.

Arriaga L et al. (2004) Potential ecological distribution of alien invasive species and risk assessment: A case study for buffel grass in arid regions of

Mexico. Conservation Biology 18:1504-1514.

Bravo-Peña LC (2009) Ganadería y cambios de la cubierta vegetal en la

RegiónCentral de Sonora.Impactossobre el paisaje y vulnerabilidad de los

productores pecuarios en el contexto del cambio climático. In: Desarrollo

Regional. CIAD, A. C., Hermosillo.

Bravo-Peña LC et al. (2010) Políticas rurales y pérdida de cobertura vegetal.

Elementos para reformular instrumentos de fomento agropecuario relacionados con la apertura de praderas ganaderas. Región y Sociedad 22:3-35

Camou E (1998) De Rancheros, Poquiteros, Orejanos y Criollos: Los Productores Ganaderos de Sonora y el Mercado Internacional. El Colegio de

Michoacán y CIAD, A.C., Zamora, Mich.

Castellanos AE (1992) Ecología, utilización y conservación de las comunidades vegetales en el estado de Sonora. Un análisis. In: Moreno JL(ed) Los

Recursos Naturales del Estado de Sonora. SIUE - El Colegio de Sonora,

Hermosillo, pp 25-37.

Castellanos AE et al. (2010) Impactos Ecologicos por el Uso del Terreno en

el Funcionamiento de Ecosistemas Aridos Semi-Aridos de Sonora. In:

Molina-Freaner F,Van DevenderTR(eds) DiversidadBiologica delEstado

de Sonora. CONABIO - UNAM., México, D. F., pp 157 - 186.

Castellanos AE et al. (2002) Drought - tolerant exotic buffel - grass and desertification. In: Tellman B (ed) Weeds Across Borders. Arizona-Sonora

Desert Museum, Tucson, AZ.

Celaya-Michel H et al. (2011) Mineralización de nitrógeno en el suelo de

zonas áridas y semiáridas. Terra Latinoamericana 29:343-356.

Chambers. KC et al. 2007. What makes Great Basin sagebrush ecosystems

invasible by Bromus tectorum. Ecological Monographs 77: 117-145.

COTECOCA (2002) Diagnóstico de los agostaderos del Estado de Sonora.

In. COTECOCA, Hermosillo, Sonora, p 52.

Dalal RC et al. (2005) Total soil carbon and nitrogen and their pools following Mulga (Acacia aneura) clearing for pasture development and cropping

1. Total and labile carbon. Australian Journal of Soil Research 43:13-20.

Franklin KA et al. (2006) buffelgrass (Pennisetum ciliare) land conversion

and productivity in the plains of Sonora, Mexico. Biological Conservation

127:62 - 71.

Haverty MI (2001) The role of toilet paper in studies of desert subterranean

termites (Isoptera) in Arizona, USA: A substrate for nondestructive observations of foraging activity. Sociobiology 37:245-252.

Hooper DU et al. (2005) Effects of biodiversity on ecosystem functioning: a

consensus of current knowledge. Ecological Monographs 75:3 - 35.

Ibarra-FloresFetal.(1999)Soilphysicochemicalchangesfollowingbuffelgrass

establishment in Mexico. Arid soil research and rehabilitation 13:39 - 52.

López D (2007)Changesin carbon and nitrogen with buffelgrass(Pennisetum 

ciliare) introduction in Sonora, Mexico. In: Biological Sciences. Northern

Arizona University, Flagstaff, AZ, p 67.

Morales-Romero D et al. (2008) Influence of buffelgrass pasture conversion

on the regeneration and reproduction of the columnar cactus, Pachycereus

pecten-aboriginum, in northwesternMexico.Journal ofAridEnvironments

72:228-237.

NaeemSet al.(2002)Biodiversity and ecosystemfunctioning:The emergence

of a synthetic ecological framework. In: Loreau M, Naeem S, Inchausti P

(eds)Biodiversity andEcosystemFunctioning.Synthesis and perspectives.

Oxford University Press, Oxford, pp 33-11.

Peinado M et al.(1990)Introducción al conocimiento fito-topográfico deBaja

California (México). Studia Botanica 9:25-39.

Perramond E (2000) A preliminary analysis of soil erosion and buffelgrass in

Sonora, Mexico. Yearbook, Conference of Latin American Geographers

26:131-138.

Romo JR (2006) Conservation and the changing pattern of land cover and

land use in central Sonora, Mexico.In: Environmental Sciences and Policy.

Northern Arizona University, Flagstaff, AZ, p 103.

Rzedowski J (1978) Tipos de Vegetación de México. LIMUSA, México.

Saucedo-Monarque E (1994) La Introducción de Zacate buffel (Cenchrus 

ciliaris L) en el Estado de Sonora, un Factor de Modificación de la Veg-
196 USDA Forest Service Proceedings RMRS-P-67. 2013

Castellanos and others Biodiversity Effects on Ecosystem Function Due to Land Use: . . .

etación. In: Instituto de Recursos Naturales. Colegio de Postgraduados,

Montecillo, México, p 129.

Saucedo-Monarque E et al. (1997) La riqueza, una variable de respuesta de

la vegetación a la introducción del zacate buffel. AgroCiencia 31:83-90

SchulzeEDet al.(eds)(1994)BiodiversityandEcosystemFunction.SpringerVerlag, Berlin.

Shreve F (1942) The desert vegetation of North America. The Botanical

Review 8:195-246.

Shreve F et al. (1964) Vegetation and flora of the Sonoran Desert. Stanford

University Press, Stanford.

Stebbins GL (1952) Aridity as a stimulus to plant evolution. The American

Naturalist 86:33-44.

Tilman D (1988) Plant Strategies and the dynamics and structure of plant

communities. Princeton University Press, New Jersey.

Tilman D et al.(2002) Plant diversity and composition: effects on productivity

and nutrient dynamics of experimental grasslands. In: Loreau M, Naeem

S, Inchausti P (eds) Biodiversity and Ecosystem Funtioning. Synthesis and

perspectives. Oxford University Press, Oxford, pp 21 - 35.

Tilman D et al. (1997) Plant diversity and ecosystem productivity: Theoretical considerations. Proceedings of the National Academy of Siciences

94:1857 - 1861

Tilman D et al. (2006) Biodiversity and ecosystem stability in a decade-long

grassland experiment. Nature 441:629-632

Turner RM et al. (1995) Sonoran Desert Plants: An Ecological Atlas. The

University of Arizona Press, Tucson

Wright JP et al. (2006) Conventional functional classification schemes underestimate the relationship with ecosystem functioning. Ecology Letters

9:111 – 120

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 197

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Biogeography and Diversity of Pines in the Madrean 

Archipelago

George M. Ferguson

University of Arizona, Tucson, Arizona

Aaron D. Flesch

Avian Science Center, Division of Biological Sciences, University of Montana, Missoula, Montana

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Abstract—Pines are important dominants in pine-oak, pine and mixed-conifer forests across the Colorado 

Plateau, southern Rocky Mountains, Sierra Madre Occidental, and in the intervening Sky Islands of the 

United States-Mexico borderlands. All 17 native species of pines in the Sky Islands region or their adjacent 

mountain mainlands reach the northern or southern margins of their geographic ranges and most (12) 

have Madrean vs. Petran affinities. We compiled data on the occurrence and diversity of pines across 31 

Sky Islands based on extensive fieldwork that complemented data from museum collections. Lower and 

smaller mountains supported zero to four species of pines whereas higher ones with greater area and thus 

larger population sizes supported five to seven species. Diversity of pines increased by 0.34% with each 

1% increase in area (km2

) of forest in a mountain range with similar effects for maximum elevation (P < 

0.001); diversity decreased by 0.70% with each 1% increase in distance (km) to the nearest mainland (P 

< 0.015). Forest area explained >2 times more variation in pine diversity (R2 = 0.42) than distance from the 

nearest mainland (R2 = 0.19), suggesting that larger population sizes of pines on bigger mountains reduce 

rates of stochastic extinctions and that this process is more important than dispersal limitation in driving pine 

diversity. Although pine species have no legal conservation status in the Sky Islands, pine forests, especially 

those on small isolated mountaintops, are among the most threatened plant communities in this region due 

to increased fire disturbance, insect outbreaks, and climate change.

Introduction

The genus Pinus includes about 110 species and is the largest genus of conifers and among the most widespread trees in the northern 

hemisphere. Mexico and Central America support the highest diversity of pine species (47) including 30 that are endemic to Mexico, 

whereas a second center of diversity of about 25 species occurs in 

eastern Asia (Price and others 1998). As noted in 1942 by Howard 

Gentry, Mexico is a cornucopia of pine species with high diversity 

that crosses the United States-Mexico border into the Madrean Sky 

Islands region (Martin and others 1998). Of 17 native species of pines 

in the Sky Islands and their adjacent cordilleran mainlands, 12 have 

stronger biogeographic affinities to the more southerly Sierra Madre 

Occidental (SMO). Notably, in this broad biogeographic transition 

zone, all 17 species of pines reach the northern or southern margins 

of their geographic ranges. 

We studied the distribution of pines in the Madrean Sky Islands 

region and assessed how mountain area, elevation, and distance to 

the nearest mainland (SMO or Colorado Plateau) affected diversity 

of pines across 31 Sky Island mountain ranges in the United States 

and Mexico. We present these results and discuss implications for 

understanding the distribution, biogeography, and conservation of 

pines in the region. 

Methods

Study Area

We considered three geographic areas: 1) mountain ranges across 

the entire Sky Islands region in southern Arizona, southwest New 

Mexico, northeast Sonora, and northwest Chihuahua, 2) the southern Colorado Plateau and Rocky Mountains (CP-RM) in southern 

Colorado, central and northern Arizona and northern and western 

New Mexico, and 3) the continuous highlands of the northern SMO 

in east-central Sonora and west-central Chihuahua that are located 

directly south and east of the Sky Islands region.

The Sky Islands region encompasses the Apachian subprovince of 

the Madrean floristic province (McLaughlin 1995, 2007, 2008) and is 

bounded to the north by the Gila River or by a level grassland plain in 

lowest saddle on the Continental Divide (1370 m). Although the region’s southern boundary has been defined based on the biogeographic 
198 USDA Forest Service Proceedings RMRS-P-67. 2013

Ferguson and others Biogeography and Diversity of Pines in the Madrean Archipelagor

affinities of the local flora, it is less clear exactly where it should be 

placed. The east and west boundaries of the Sky Islands region are 

more well defined by vast level plains of low relief beyond the Big 

Hatchet mountains in New Mexico and the Baboquivari mountains 

in Arizona/Sierra el Humo in Sonora, respectively (Marshall 1957; 

Flesch and Hahn 2005).

Design

We considered mountain ranges that supported at least one species 

of pine in the flora and that supported woodlands of oaks and pines 

that were isolated and discrete from other such patches. Mountain 

ranges containing pines have peak elevations greater than 1885 m, 

and we considered mountain ranges as discrete (a) if they were not 

connected by elevations >1555 m (the highest saddle with a wide 

grassland ecotone); (b) if there were no continuous woodland across 

a saddle if higher than 1555 m; and (c) if continuous woodland exists without pines if lower. The role of vegetation was important in 

determining three: (1) a desert grassland saddle at 1675 m between 

the Animas and Pan Duro/Sierra San Luis mountains separates them 

as islands on the eastern edge of the Sky Island region bordered by 

drier desert grassland and Chihuahuan desertscrub; (2) a continuous 

oak woodland saddle at 1350 m between the Sierra de los Ajos/Púrica 

and the Sierra Juriquipa that has a broad area connecting the ranges 

(on a rapidly rising Pacific slope gradient); and (3) a problematic zone 

noted by McLaughlin (1995) that includes highland extensions of SMO 

woodlands in the Sierra Hachita Hueca and El Gato (Huachinera) 

with saddles >1555 whereas the 2400 m high Sierra Bacadéhuachi 

is an island because of the low 1340 m saddle of oak woodland that 

separates Sierra Bacadéhuachi from more extensive woodlands of 

the SMO. 

Data Collection and Analysis 

We compiled information on the presence and distribution of pines 

during many years (1995-2011) of fieldwork. We supplemented these 

data with historical records from museum records and databases such 

as the Southwest Environmental Information Network (SEINet) 

and Madrean Archipelago Biodiversity Assessment (MABA) floral 

database (http://www.madrean.org) dating back to 1880. Information 

on pine distribution outside the Sky Islands region for determining 

extent and geographic center of species ranges was obtained from 

Little (1999). 

Because the taxonomy of some species in the genus Pinus is equivocal, we verified species by examining representative specimens and 

used nomenclature from published sources (Farjon and Styles 1997; 

Ferguson and others 2001; García-Arevalo and González-Elizondo 

1998; Kral 1993; Price and others 1998). Recent phylogenetic analyses based on molecular data and phytochemistry have provided new 

information on some of these taxonomic problems. Definitions of 

some of the more problematic species are defined in the Appendix.

To assess the effects of large-scale landscape and biogeographic 

factors on diversity of pines in the Sky Islands region, we used four 

variables. As a response variable we calculated the number of Pinus 

species found in the flora of each mountain island. As explanatory 

variables we considered total area of woodland and forest (km2

) 

that could potentially support pines, the maximum elevation of each 

mountain range from sea level to summit (m), and the distance from 

each mountain range to the nearest mainland or potential source 

population of immigrants (km; CP-RM or SMO). We only considered 

mountains that supported pines because we were interested only in 

these relict and isolated populations of the Holocene, rather than the 

broader dispersal routes of past climates.

To estimate the area of woodland and forest vegetation in each 

Sky Island, we used data from The Nature Conservancy’s Apache 

Highlands Ecoregion study for mountains in Arizona, New Mexico, 

and northern Sonora and Chihuahua (Marshall and others 2004). For 

other mountain ranges in Mexico we used data from the Mexican 

National Forest Inventory and combined comparable vegetation 

communities (Palacio-Prieto and others 2000; Velazquez and others 

2000). Using this first data source, we calculated woodland area for 

each mountain range by summing the areas among seven vegetation communities: Madrean Encinal, Madrean Oak-Pine Woodland, 

Pinyon-Juniper Woodland, Interior Chaparral, Montane Mixed-Conifer 

Forest, Ponderosa Pine Forest and Woodland, and Sub-alpine SpruceFir Forest and Woodland. For this second data source, we computed 

comparable estimates by summing the area of Chaparral, Bosque de 

Encino, Tascate, Encino-Pino, and Pino-Encino.

We used estimates of the maximum elevation of each Sky Island 

from USGS and INEGI topographic maps rounded to nearest 5 m. 

Distance estimates between each Sky Island and the CP-RM and SMO 

were obtained from 1:500,000 scale NOAA Sectional Aeronautical 

Charts and from Google Earth. We measured distances between the 

summit of each mountain range and a fixed reference point for each 

mainland. The CP-RM reference point was located at the southern 

edge of this highland at 2740 m elevation (33°33’N,109°20’W; near 

U.S. Highway 191 south of Hannagan Meadow, Arizona) and the 

SMO reference point was at the north edge of this highland at 2690 m 

elevation (30°33’ N.,108°37’W.; Mesa las Guacamayas, Chihuahua).

To model the effects of area, elevation, and isolation on pine species 

diversity, we log transformed each variable and used simple linear 

regression to estimate the slope of each individual effect and the total 

variation in pine diversity that was explained by each explanatory 

variable. To assess the combined effects of different explanatory 

variables on pine diversity, we used multiple linear regression. Because maximum elevation was correlated with area of woodland and 

forest (r = 0.41, P = 0.024), we only used area and distance in the 

model. Because estimates from two mountain ranges were outliers, 

we conducted some analyses with and without these values. We also 

assess the effects of distance from the CP-RM or SMO on diversity 

of northern and southern pine species, respectively.

Results

Pine Species Occurrence and Diversity 

A total of 17 pine species occur in the study area (table 1). Nine 

species of pines occur in 31 Sky Islands including five species that 

are widely distributed in the Sky Islands. The northern mainland (CPRM) supports two species of pines (Pinus aristata and P. contorta) 

that do not occur at the southern edge of the Colorado Plateau or in 

the nearby Sky Islands. The southern mainland (SMO) supports five 

species (P. herrerae, P. leiophylla, P. lumholtzii, P. maximinoi, and P. 

oocarpa) in the north-central portion of the range that do not occur at 

the northern edge of the SMO or in the Sky Islands; a sixth Madrean 

species of pine (P. cembroides) does occur at the northern edge of 

the SMO but not in the Sky Islands. 

The highest diversity of pines in a single Sky Island is seven species, 

occurring in the Chiricahua Mountains, the second highest elevation 

island. Of the other high islands, three have six species (Pinaleño, 

Huachuca, and Santa Rita) and the lower wider Galiuro Mountain also 

have six. The most diverse Sky Islands have about the same number 

of species as found at the edge of the CP-RM mainland (seven), in 
USDA Forest Service Proceedings RMRS-P-67. 2013 199

Biogeography and Diversity of Pines in the Madrean Archipelago Ferguson and others

the CP-RM proper (six), or at the edge of the SMO (eight) although 

the species present in the north and south are different. With 13 pine 

species, the SMO has a much higher diversity than any individual 

Sky Island, although at any one location in the northern SMO only 

as many as 7 species co-occur. 

Three biotic provinces (Madrean, Cordilleran, Apachian) of Dice 

(1943) are represented by the floristic elements defined by McLaughlin 

(2008). Of the 17 species of pines in our study area, five (~30%) have 

affinities to the northern floristic element of the CP-RM subprovinces 

and 12 (~70%) have affinities to a southern floristic element of the 

Madrean floristic province. Three Madrean element pines (Pinus 

chihuahuana, P. discolor, and P. arizonica) extend as far north as the 

edge of the CP-RM, and a fourth (P. strobiformis) extends onto the 

Colorado Plateau. One CP-RM element pine (P. ponderosa) extends 

to the edge of the SMO and onto the SMO proper in the northern part 

only. 

Figure 1—Relationship of pine species diversity (ln) to woodland area, elevation, and distance to the nearest mainland (left to right) in all 31 Sky 

Island mountain ranges in the Madrean Archipelago. The regression line is the estimate of diversity for each effect. An outlier for the effect of area 

is Sierra San José.

Table 1—Biogeographic affinities and presence (X) of nine Pinus species in 31 Sky Islands and adjacent mainlands, with peak 

elevations for each island, from north (left) to south (right) in (A) southwestern U.S. and (B) northwestern Mexico.
200 USDA Forest Service Proceedings RMRS-P-67. 2013

Ferguson and others Biogeography and Diversity of Pines in the Madrean Archipelagor

Drivers of Diversity

Both the area of woodland habitat and the maximum elevation of 

a Sky Island have large effects on diversity. Pine species diversity 

increased by 0.34 ± 0.074% (± SE) with each 1% increase in area 

of woodland (km2) that was present in a Sky Island (P < 0.0001; 

fig. 1) and by 3.20 ± 0.074% (± SE) with each 1% increase in 

elevation (P < 0.0001; fig. 1). The effect of area was large even 

with a Sky Island outlier, Sierra San José included. Island isolation 

also affected pine species diversity but explained less variation than 

area or elevation when considered independently (table 2). Species 

diversity decreased by 0.70 ± 0.27% (± SE) with each 1% increase 

in distance to nearest mainland (P = 0.015; fig. 1). We found that one 

Sky Island was an outlier with respect to the distance effect; when 

Big Hatchet was removed the effects of distance on pine diversity 

increased (table 2). Distance to the nearest mainland was not as correlated with either woodland area or elevation (r = -0.37, P < 0.05 

and r = -0.35, P = 0.05 respectively). 

When only pines species in the northern element were considered 

for each island, distance to CP-RM had a large effect on species 

diversity (table 3); species diversity decreased by 0.93 ± 0.15% (± 

SE) with each 1% increase in distance from the CP-RM (P < 0.0001; 

fig. 2). When only pines in the southern element were considered for 

each island, distance to the Sierra Madre had much less of an effect 

on pine species diversity; species diversity decreased by 0.36 ± 0.18% 

(± SE) with each 1% increase in distance from the SMO (P = 0.065; 

fig. 2) and by 0.47 ± 0.15% (± SE) with each 1% increase in distance 

once Big Hatchet was removed (P = 0.004; table 3). 

When the combined effects of habitat area and isolation were 

considered, there was some evidence that both woodland area and 

isolation explained pine diversity (table 4). Species diversity increased 

by 0.30 ± 0.08% (± SE) with each 1% increase in the log number of 

km2 of woodland (P < 0.0006) after adjusting for the effect of distance to the nearest mainland, whereas, species diversity decreased 

by 0.38 ± 0.24% (± SE) with each 1% increase in log distance to the 

nearest mainland (P = 0.12) after adjusting for the area effect. With 

the Sierra San José and Big Hatchet outliers removed, the amount of 

variation in pine diversity that was explained by area and isolation 

increased somewhat (R2 = 0.61 vs. 0.47) and the effect of isolation 

became stronger. 

Discussion

Biologists have tried to define factors that explain species diversity 

across the mountain islands of the western United States and assess 

how habitat area and isolation affect the distribution of population 

and diversity of communities. Some studies suggest that both factors 

are important in explaining diversity of montane birds and mammals 

in the borderlands mountains while other studies indicate habitat is 

more important than isolation (Gehlbach, 1995). 

We found that these two variables, woodland area and distance 

to the nearest mainland, considered independently, were each good 

predictors of pine diversity across the mountain islands of the Madrean 

Archipelago. Maximum elevation was also considered a potential 

explanatory variable and found to be a significant predictor of pine 

species diversity (table 2). The relationship between a mountain’s peak 

elevation and its area of woodland that supports pines can be attributed to the “Merriam Effect” (i.e. increase in orographic precipitation 

and adiabatic cooling with increases in elevation), so the larger the 

mountain island, the more conifer forest it contains (Gehlbach, 1981). 

In our study, maximum elevation and woodland area were correlated 

and essentially measured the same attribute—habitat area and thus 

population size of pines. Isolation distance explained less variation 

in pine species diversity than area, but did have some explanatory 

power even when adjusting for the effect of area (table 4.) 

Table 2—Simple Linear Regression model for the effects of habitat area, elevation and distance to the 

nearest mainland on the diversity of pine species (ln) in all 31 Sky Island mountain ranges in the 

Madrean Archipelago and with one outlier removed.

Data set Explanatory variable R2 Estimate SE P

All data

Area of pine-oak woodland (ln km2

) 0.42 0.34 0.07 <0.0001

Maximum elevation (ln m) 0.43 3.20 0.69 <0.0001

Distance to nearest mainland (ln km) 0.19 -0.70 0.27 0.015

Hatchet removed

Distance to nearest mainland (ln km) 0.26 -0.78 0.25 0.004

Table 3—Simple Linear Regression models for the effects of distance to the Colorado Plateau on the 

diversity of northern pine species (ln) and distance to the Sierra Madre Occidental on the diversity of 

southern pine species (ln) in all 31 Sky Island mountain ranges in the Madrean Archipelago and with 

one outlier removed.

Data set Explanatory variable R2 Estimate SE P

All data

Distance to Colorado Plateau (ln 

km)

0.55 -0.93 0.15 <0.0001

Distance to Sierra Madre (ln km) 0.11 -0.36 0.18 0.065

Hatchet removed

Distance to Sierra Madre (ln km) 0.26 -0.47 0.15 0.004
USDA Forest Service Proceedings RMRS-P-67. 2013 201

Biogeography and Diversity of Pines in the Madrean Archipelago Ferguson and others

In examining the distribution of reptiles and amphibian species, 

Swann and others (2005) found that none of the Sky Islands contain 

a full complement of southern species and that few reptile and amphibian species in the Sky Islands have affinities to areas north of 

the Sky Islands. For pines, none of the 31 Sky Islands contain a full 

complement of six southern element species although eight islands 

have five of the six species, and half of those are in Mexico. All Sky 

Islands with pines have at least one pine species of southern element 

in its flora with the exception of the Big Hatchet Mountains. Only 

one Sky Island has a full compliment of three northern element pine 

species. Of the 11 Sky Islands that have at least one northern pine 

species, just one mountain, the Sierra San José, is in Mexico. 

An outlier in terms of the effect of distance from the southern 

mainland on the diversity of pines is the 2540 m high Big Hatchet 

Mountains. They are positioned close to the southern mainland and 

support no southern pines species. This could be associated with their 

longitude as they are at the easternmost edge of the of the Madrean 

Sky Islands and thus surrounded by the more arid adapted Chihuahuan 

desertscrub that is east of the Continental Divide, and the only high 

Sky Island composed entirely of limestone. The Florida Mountains 

and Cooke’s Range farther east are not limestone, but also have the 

same single northern element pine species in their flora, Pinus edulis. 

A dispersal corridor may be to the north along the Continental Divide rather than to the SMO, despite a stepping stone in the adjacent 

Animas Mountains, which supports five southern pines. On floristic 

grounds, Big Hatchet is outside of the Apachian floristic subprovince 

(McLaughlin 1995, 2008).

An outlier in terms of the effect of woodland area on pine species 

diversity is the 2520 m Sierra San José, which has much higher diversity than expected given its small area. This pattern may be partly 

explained by latitude. Sierra San José is the second highest in elevation 

and the farthest north of any Sky Island in Mexico, but its mountain 

mass (ca. 1 km2 above 2300 m) and corresponding woodland area 

are small. In addition, it is relatively close to two higher mountains 

(Huachuca Mountains in the United States, and Sierra de los Ajos, the 

highest island in Mexico) that may act as potential source populations 

of pines. Birds are known to disperse the seeds of certain pines species 

and two species that have been found in the San José, Pinus discolor 

and P. strobiformis belong to the strobi and cembroides groups of 

pines that have large and wingless seeds potentially dispersed by birds 

Figure 2—Relationship of pine species diversity (ln) to distance to the Colorado Plateau for northern pine species (left), 

and distance to the Sierra Madre Occidental for southern pine species (right) in all 31 Sky Island mountain ranges in the 

Madrean Archipelago. The regression line is the estimate of diversity for each effect. An outlier for the effect of distance 

on the diversity of southern pines is the Big Hatchet Mountains.

Table 4—Multiple Linear Regression model for the effects of habitat area and distance to the 

nearest mainland on the diversity of pine species (ln) in all 31 Sky Island mountain ranges in 

the Madrean Archipelago and with two outliers removed.

Data set Explanatory variable Estimate SE P

All data 

(R2 = 0.47)

Area of pine-oak woodland (ln km2

) 0.30 0.08 0.0006

Distance to nearest mainland (ln km) -0.38 0.24 0.12

Hatchet and San Jose removed 

(R2

 = 0.61)

Area of pine-oak woodland (ln km2

) 0.33 0.07 <0.0001

Distance to nearest mainland (ln km) -0.38 0.20 0.074
202 USDA Forest Service Proceedings RMRS-P-67. 2013

Ferguson and others Biogeography and Diversity of Pines in the Madrean Archipelagor

(Lanner 1998). Our recent fieldwork, however, failed to document P. 

strobiformis (or P. chihuahuana) in the San José that were originally 

found in 1893 by Mearns (1907; specimens examined by us). These 

species may now be locally extirpated due to greater susceptibility of 

small populations to stochastic extinctions as a continuing process, 

i.e. the few P. strobiformis individuals Mearns found near the summit 

removed by fire. 

Varying adaptive strategies of Sky Island pines might explain their 

present distributions. One pine of the CP-RM element continues 

across the Sky Islands into the northern SMO, Pinus ponderosa var. 

scopulorum; it belongs to a widespread and adaptable “ponderosae” 

complex with varieties (and species) that range from British Colombia 

to central Mexico. Pleistocene and early Holocene records are only in 

Arizona; its range expanded northward during the last 8000 years just 

as another northern element P. edulis did from refugia in the southwestern United States (Lanner and Van Devender 1998). Two pines 

of the Madrean element that range across 23 Sky Islands and reach 

the edge of the Colorado Plateau can survive at lower elevations—P. 

discolor as low as 1150 m, and P. chihuahuana, on the fringe of the 

Sky Islands as low as 1310 m—and exhibit better drought tolerance 

than other pines. P. engelmannii is the dominant pine in the SMO, 

but also occurs on the edge of the SMO and in lower Sky Islands 

where populations are small, yet does not occur into the northern 

Sky Islands. P. arizonica, widespread in the higher SMO, mostly 

occurs on mesic N-facing slopes, or canyon bottoms of the higher 

Sky Islands including the highest northern ones. P. strobiformis, the 

most widespread and highest elevation of any Madrean element pine 

in the Southwestern United States, is potentially bird-dispersed (by 

corvids and thick-billed parrots). The northernmost populations of P. 

yecorensis in the Sierra Bacadéhuachi, and adjacent Sierra Huachinera 

of the SMO, are small and perhaps cold limited; farther south the 

species is more widespread on the lower Pacific slope of the SMO 

and borders Tropical Deciduous Forest around latitude 28°N.

None of the Pinus in the Madrean Sky Islands of Mexico or the 

United States have any federal or state conservation status, although 

limited protected areas exist as reserves (e.g. Ajos-Bavispe, Ramsey 

Canyon, Saguaro National Park), have controlled access (e.g. Animas Mountains, private), or are managed by the U.S. Forest Service. 

Disturbance events have altered pine forests at large scales across the 

Sky Islands (increased fires associated with human activities, insect 

outbreaks and drought). In smaller ranges where pines occur in tiny 

isolated populations, these disturbance events are affecting pine 

diversity and distribution, e.g., a tiny population of P. engelmannii 

on privately managed land in the Sierra San Antonio may have less 

then 20 individuals at present, with many recent snags that have succumbed to drought in the past 10 years. Similarly, Pinus engelmannii 

in the Sierra Cíbuta, the westernmost outpost for pines in the Sky 

Islands of Sonora, are in serious decline and many patches of pines 

have been lost in the last decade. Although marked as present on our 

list, such pines can be rare, e.g. only a single P. edulis was recorded 

from the Pinaleño Mountains (Jim Malusa, personal observation 

1985). Other areas may be under sampled, or less explored, such 

as in Sierra Enmedio, Chihuahua, where no collections exist for a 

high mountain bound to support at least one pine species, and in the 

Dragoon Mountains, Arizona, with only one pine species recorded. 

The spread of some introduced pines should also be monitored, such 

as the Eurasian Pinus sylvestris that is locally introduced on private 

lands in the Sierra de los Ajos, Sonora. Given threats to small isolated 

populations on mountain tops and a rapidly warming and drying climate in the region, efforts to document and monitor the distribution 

and spatial extent of pine forests across the Sky Islands is critical for 

conservation. 

Acknowledgments

Nick Deyo and Alex Smith, Sky Island Alliance, Tucson, provided 

data from the vegetation inventories by island, on which this study 

depended. Elaine Nakash assisted in mapping pine distributions. 

Others who either helped with field reconnaissance or in obtaining 

observational or museum records are: John Anderson, Debra Bell (SI), 

Mark Fishbein, Steve Hale, Sky Jacobs, Mark Kaib, Shelley McMahon (ARIZ), Jim Malusa, Paul Martin, Humberto Mata-Mangueros, 

James Riser, Mike Rourke, and Jesús Sánchez-Escalante (USON). 

We thank two reviewers of an earlier version of the manuscript, Dr. 

Ronnie Sidner and Dr. Steve McLaughlin, who provided helpful comments. Support for ADF’s efforts in the Sky Islands was provided by 

the U.S. National Park Service.

References

Almaraz-Abarca, Norma; González-Elizondo, M. Socorro; Tena-Flores, Jorge 

A.; [and others]. 2006. Foliar flavonoids distinguish Pinus leiophylla and 

Pinus chihuahuana (Coniferales: Pinaceae). Proceedings of the Biological 

Society of Washington 119 (3): 426-436.

Cole, Kenneth L; Fisher, Jessica; Arundel, Samantha T.; [and others]. 2008. 

Geographical and climatic limits of needle types of one- and two-needled 

pinyon pines. Journal of Biogeography 35:257-269.

Dice, L.R. 1943. The biotic provinces of North America. Ann Arbor; University of Michigan. 

Epperson, Bryan K; Telewski, Frank W.; and Willyard, Ann. 2009. Chloroplast 

diversity in a putative hybrid swarm of Ponderosae (Pinaceae). American 

Journal of Botany 96(3): 707-712.

Farjon, Aljos; and Styles, Brian T. 1997. Pinus (Pinaceae). Flora Neotropica 

Monograph 75. New York; The New York Botanical Garden. 291 p.

Ferguson, George M.; Felger, Richard S.; and Johnson, Matthew B. 2001. 

Conifers pp. 35-58. in Felger, Richard S.; Johnson, Matthew B.; and Wilson, 

Michael F. The Trees of Sonora, Mexico. New York; Oxford University. 

Flesch, A. D.; and L. E. Hahn. 2005. Distribution of birds and plants at the 

western and southern edges of the Madrean Sky Islands in Sonora, Mexico. 

pp 80-87 in Gottfried, Gerald J.: Gebow, Brooke S.; Eskew, Lane G.; 

and Edminster, Carlton B., compilers. Connecting mountain islands and 

desert seas: biogeography and management of the Madrean Archipelago 

II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. Fort Collins, 

CO; U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Reseach Station. 631 p.

García-Arévalo, A.; and González-Elizondo, M. Socorro. 1998. Pináceas de 

Durango. Durango, México; CIIDIR-IPN, Instituto de Ecología A.C.,179 p.

Gehlbach, Frederick R. 1981. Mountain islands and desert seas: a natural history 

of the U.S.-Mexican borderlands. College Station; Texas A&M University. 

Gehlbach, Frederick R. 1995. Biogeographic controls of avifaunal richness in 

isolated coniferous forests of the U.S.-Mexican borderlands. pp 135-150 

in Istock, Conrad A.; and Hoffman, Robert S. eds. Storm over a mountain 

island. Conservation biology and the Mt. Graham affair. Tucson; University 

of Arizona.

Kral, Robert 1993. Pinus. pp 373-398 in Flora of North America Vol 2. New York; 

Oxford University. Published online http://www.fna.org/FNA/index.html

Lanner, Ronald M. 1998. Seed dispersal in Pinus. pp 281-295 in Richardson, 

David M. ed. Ecology and Biogegraphy of Pinus. Cambridge; Cambridge 

University.

Lanner, Ronald M.; and Van Devender, Thomas R. 1998. The recent history 

of pinyon pines in the American Southwest. pp 171-182 in Richardson, 

David M., ed. Ecology and Biogegraphy of Pinus. Cambridge; Cambridge 

University.

Little, Elbert L., Jr. 1999. Atlas of United States Trees. United States Geological Survey. Digital representation. http://esp.cr.usgs.gov/data/atlas/little/

vegmaps_met.html

Marshall, Joseph 1957. The Birds of the pine oak woodland in southern Arizona 

and adjacent Mexico. Pacific Coast. Avifauna No.32.
USDA Forest Service Proceedings RMRS-P-67. 2013 203

Biogeography and Diversity of Pines in the Madrean Archipelago Ferguson and others

Marshall, Robert M.; Turner Dale; Gondor Anne; [and others]. 2004. An 

Ecological Analysis of Conservation priorities in the Apache Highlands 

ecoregion. Prepared by The Nature Conservancy of Arizona, Instituto del 

Medio Ambiente y el Desarrollo Sustentable del Estado de Sonora, agency 

and institutional partners. 152 p.

Martin, Paul S.; Yetman, David; Fishbein, Mark; [and others], eds. 1998. 

Gentry’s Río Mayo Plants. The tropical deciduous forest and environs of 

northwest Mexico. Tucson: University of Arizona. 558 p.

McLaughlin, Steven P. 1995. An overview of the flora of the sky islands, 

southeastern Arizona: diversity, affinities, and insularity. pp 60-70 in DeBano, Leonard F.; Ffolliott, Peter F.; Ortega-Rubio, Alfredo; [and others], 

tech. coords. Biodiversity and management of the madrean achipelago; the 

sky islands of southwestern United States and northwestern Mexico; 1994 

Sept. 19-23; Tucson, AZ. Gen Tech. Rep. RM-GTR-264. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest 

and Range Experiment Station. 669 p. 

McLaughlin, Steven P. 2007. Tundra to tropics: The floristic plant geography 

of North America. Sida Botanical Miscellany 30:1-58.

McLaughlin, Steven P. 2008. Vascular flora. pp 120-140 in Ffolliott, P.F.; and 

Davis, O.K. eds. Natural Environments of Arizona. Tucson: University 

of Arizona.

Mearns, Edgar A. 1907. Mammals of the Mexican boundary of the United 

States. United States National Museum Bulletin 56.

Palacio-Prieto, J.L.; Bocco G.; Velazquez, A.; [and others]. 2000. La condicion actual de los recursos forestales en Mexico: resultados del inventario 

forestal nacional 2000. Investigaciones Geograficas, Boletin del Instituto 

de Geografía, UNAM 43: 183-203

Price, R.A; Liston, A; and Strauss, S.H. 1998. Phyologeny and systematics 

of Pinus. pp 49-68 in Richardson, D.M., ed. Ecology and biogegraphy of 

Pinus. Cambridge; Cambridge University. 

Swann, Don E.; Mau-Crimmins, Theresa M.; and Stitt, Eric W. 2005. In 

search of the Madrean Line: biogeography of the herpetofauna in the Sky 

Island Region. in Gottfried, Gerald J.: Gebow, Brooke S.; Eskew, Lane G.; 

and Edminster, Carlton B., compilers. Connecting mountain islands and 

desert seas: biogeography and management of the Madrean Archipelago 

II; 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Reseach Station. 631 p.

Velázquez, A., Mas, J. F.; Saucedo, M.; Palacio-Prieto, J.L. [and others]. 2001. 

El Inventario Forestal Nacional 2000. Ciencias No. 64:12-19. http://www.

ejournal.unam.mx/ciencias/no64/CNS06403.pdf

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
204 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Ecosystems and Diversity of the Sierra Madre Occidental

M. S. González-Elizondo, M. González-Elizondo, L. Ruacho González, I.L. Lopez Enriquez, 

F.I. Retana Rentería, and J.A. Tena Flores

CIIDIR I.P.N. Unidad Durango, Mexico

Abstract—The Sierra Madre Occidental (SMO) is the largest continuous ignimbrite plate on Earth. Despite 

its high biological and cultural diversity and enormous environmental and economical importance, it is yet 

not well known. We describe the vegetation and present a preliminary regionalization based on physiographic, climatic, and floristic criteria. A confluence of three main ecoregions in the area corresponds with 

three ecosystems: Temperate Sierras (Madrean), Semi-Arid Highlands (Madrean Xerophylous) and Tropical 

Dry Forests (Tropical). The Madrean region harbors five major vegetation types: pine forests, mixed conifer 

forests, pine-oak forests, oak forests and temperate mesophytic forests. The Madrean Xerophylous region 

has oak or pine-oak woodlands and evergreen juniper scrub with transitions toward the grassland and 

xerophylous scrub areas of the Mexican high plateau. The Tropical ecosystem, not discussed here, includes 

tropical deciduous forests and subtropical scrub. Besides fragmentation and deforestation resulting from 

anthropogenic activities, other dramatic changes are occurring in the SMO, including damage caused by bark 

beetles (Dendroctonus) in extensive areas, particularly in drought-stressed forests, as well as the expansion 

of chaparral and Dodoneaea scrub at the expense of temperate forest and woodlands. Comments on how 

these effects are being addressed are made.

Introduction

The Sierra Madre Occidental (SMO) or Western Sierra Madre is the 

longest mountain range in Mexico and the largest continuous ignimbrite plate on Earth. It has a surface of 251,648 km2

 (not including the 

Madrean Archipelago), about the size of the State of Wyoming and 

even larger than the United Kingdom. Elevations range from 300 to 

3340 m. With a rugged physiography of highland plateaus and deeply 

cut canyons, it extends more than 1160 km from near the United 

States-Mexico border (30° 35’N) to northern Jalisco (21°00’N) in 

western Mexico. It is linked to the Rocky Mountains through the Sky 

Islands in the north and connected with the Trans-volcanic Belt to the 

south. It is a boundary area for hundreds of species that reach their 

northern or southern range limits; it also contains a high proportion of 

endemic elements. The SMO is the source of environmental services 

for a large area in northwestern and north-central Mexico, includes 

about two thirds of the standing timber in Mexico, and boasts some 

of the richest diversity of habitats and species anywhere in North 

America.

The prominence of the SMO in the biological diversity of Mexico 

has been addressed by Bye (1995), González-Elizondo (1997), Felger 

and Wilson (1995), Felger and others (1997), and Van Devender 

and Reina (2005), among others. Two areas have been recognized 

by the International Union for the Conservation of Nature (IUCN) 

as megacenters of plant diversity: northern Sierra Madre Occidental 

and the Madrean Archipelago (Felger and others 1997) and the Upper 

Mezquital River region (González-Elizondo 1997). Its high diversity 

has been noted by Spellenberg and others (1996) who found 823 

plant species for the Cascada de Basaseachi National Park. Felger 

and others (1997) estimated 4000 species of vascular plants for the 

northern portion of the range, Martin and others (1998) reported more 

than 2800 species for the Rio Mayo region, and Vázquez-García and 

others (2004) listed 2081 species for the Huichol region in the south.

A study on bats (Torres-Morales and others 2010) revealed that 

species richness of the SMO in northwestern Durango (including 

temperate and tropical areas) is comparable to that of the tropical 

rain forests of southeastern Mexico as a result of the great diversity 

of habitats in relatively small areas. This exceptional diversity is mirrored by three physiognomically dominant Madrean genera: Pinus, 

with 24 species (46% of the Mexican total), Quercus with 54 species 

(34%) and Arbutus with 7 species (100%) (González-Elizondo and 

others, in review). Many new Madrean taxa have been described from 

the region during the last 10 years, including one genus (GonzálezElizondo and others 2002) and more than 20 additional species, 

suggesting that numerous undescribed taxa are still to be found. The 

recent discovery of a second population for Pinus maximartinezii 

(González-Elizondo and others 2011) after almost 5 decades since 

its description reveals the probability of deep gaps in knowledge of 

biodiversity in the SMO.

Studies on SMO ecosystems and its surrounding areas started in 

the 1940s and continue to date (Spellenberg and others 1998; Van 

Devender and others 2003; Martínez-Yrízar and others 2010; and 

papers cited above). However, the Sierra Madre Occidental is far from 

being biologically and ecologically well known. We present here an 

outline of the main Madrean vegetation types and a preliminary delimitation of ecoregions of the SMO based on physiographic, climatic, 

and floristic criteria. Vegetation was mapped using ArcGis 9.3 based 
USDA Forest Service Proceedings RMRS-P-67. 2013 205

Ecosystems and Diversity of the Sierra Madre Occidental González-Elizondo and others

on the vegetation and land use map data in vector format Series III 

of INEGI (2002), with adaptations based on field and literature data. 

This delimitation helps to understand how organisms are distributed 

in the SMO; as important, it should be a baseline for future studies 

on taxonomy, biogeography, and ecology. 

Results and Discussion

Ecoregions

Three main ecoregions converge in the SMO: Temperate Sierras, 

Semi-Arid Highlands, and Tropical Dry Forests (CEC 1997). We refer 

to them as Madrean, Madrean Xerophylous, and Tropical in the study 

area.

M—Madrean region occurs on the higher areas of the sierra, following the Continental “backbone,” including highland ridges and 

a highland plateaus (fig. 1 M). It is almost 200 km wide in some 

places, its climates are temperate and semi-cold, and its prevalent 

communities are temperate forests in which several conifers and oaks 

are dominant elements.

MX—Madrean Xerophylous region occupies the eastern foothills 

and northern and eastern branches of the cordillera (fig. 1 MX), with 

semidry temperate or semidry semicold climates. Oak or pine-oak 

woodlands and evergreen scrub are the dominant vegetation.

T—Tropical region enters the SMO through the tributary ravines 

on the steep western flanks that drop to the narrow Pacific coastal 

plain (fig. 1 T). Climates are warm, semi-warm, and dry-warm, with 

a prevalence of tropical deciduous forests and subtropical scrub. 

Two subregions are recognized based on physiographic and floristic 

criteria within the Madrean region:

M—Madrean sensu stricto on the highland ridges and the highland 

plateau.

MT—Madrean-Tropical on the upper western slopes in which 

the climates are still temperate but moister and warmer that on the 

interior of the range. This subregion is recognized by the presence of 

conifer and oak species with strong affinities to the south, as Pinus 

oocarpa, P. douglasiana, P. devoniana, P. luzmariae, P. maximinoi, 

P. praetermissa, P. yecorensis, Quercus candicans, Q. scytophylla, 

Q. subspathulata, and Juniperus flaccida, among others.

The Madrean sensu stricto region can be further divided into 

Northern, Central and Southern ranges (fig. 1):

M1—The Northern range includes eastern Sonora and western 

Chihuahua south to the Urique barranca (between 27° and 28° N). 

With a mean elevation of 2350 m, it has colder and more continental 

climates than the rest of the SMO. Besides the species shared with 

the Madrean Archipelago to the north (e.g. Juniperus scopulorum, 

Quercus gambelii), Pinus yecorensis and Vaccinium chihuahuense 

are characteristic of the zone.

M2—The Central range extends from southwestern Chihuahua to 

southern Durango and eastern Sinaloa, ending with the Quebrada del 

río Mezquital (22°50’-23°25’ N). With a mean elevation is 2650 m, it 

includes peaks above 3200 m. Many species and a genus (Megacorax) 

are restricted here.

M3—The Southern range includes the area from the Río Mezquital 

to northern Jalisco. It is a rugged zone with deep and wide canyons 

through which the tropical and xerophylous zones converge. Many 

species are restricted to the area, e.g., Pinus maximartinezii. An example of the diversity and distribution of plant species by regions is 

given with Arbutus and Agave (table 1).

Because barriers to distribution across the tropical region are low, 

most tropical elements are distributed on a SE to NW continuum along 

the western foothills of the SMO from northern Nayarit and Jalisco 

to Sonora and Chihuahua. Nevertheless, some elements are evolving 

more locally, such as Agave, which show differential distributions 

with some species restricted to the subregions here proposed (table 1). 

As for Arbutus, the SMO harbors the highest diversity worldwide, 

with seven Madrean species. Arbutus arizonica is widely distributed 

and but other species are restricted to some subregions.

Vegetation

The Madrean, Madrean Xerophylous and Tropical regions include 

characteristic ecosystems and vegetation. The Madrean region harbors 

five major types: pine forests, mixed conifer forests, pine-oak forests, 

oak forests, and temperate mesophytic forests as well as communities 

of primary and secondary chaparral and montane meadow vegetation. Madrean Xerophylous includes oak or pine-oak woodland and 

evergreen juniper scrub with transitions toward the grassland and 

xerophylous scrub of the Mexican high plateau and to the subtropical 

scrub at the southern area of the Madrean Archipelago subregion. A 

map (fig. 2) illustrates the Madrean and Madrean Xerophylous vegetation types. The Tropical region has a prevalence of tropical deciduous 

forests and subtropical scrub, with small spots of tropical subdeciduos 

forest. These three, with no Madrean affinities, are not treated here. 

They would occupy the ravines on the western and southern flanks 

of the sierra in the map.

The classification of the vegetation follows Rzedowski (1978), a 

system of basic and excluding categories that is (1) easy to subcategorize and adapt to more specific situations; (2) comparable with 

other systems worldwide; and (3) relatively easy to map. The system 

combines physiognomic, floristic and ecological criteria.

Pine Forests

Between 1600 and 3320 m, these forests cover 12% of the surface 

area in diverse combinations of Pinus. They occur in several environmental conditions, with combinations of mainly P. arizonica, P. 

engelmannii, and P. chihuahuana on the northern and central zones of 

the range; P. durangensis or P. teocote from central Chihuahua to the 

south; and P. leiophylla and P. strobiformis in humid sites. P. cooperi 

is monodominant in valleys with deep soils, while the weeping pine (P. 

lumholtzii) dominates thin, acidic, poor soils. On the western slopes 

(region MT), P. oocarpa, P. yecorensis, P. douglasiana, P. herrerae, 

P. devoniana, and P. maximinoi dominate in subhumid places, P. 

lumholtzii or P. luzmariae on poor soils.

Mixed Conifer Forests

Fir and pine-fir forests cover less than 0.3% of the surface area 

mainly in small patches between 1900 and 3300 m in humid ravines 

and slopes in the north. The physiognomic dominants are Douglas-fir 

(Pseudotsuga menziesii) and species of Abies and Picea associated 

with Pinus and sometimes Quercus. Pseudotsuga predominates, but 

there are also small areas of Abies durangensis and Picea chihuahuana, 

and even smaller areas of Abies concolor, A. neodurangensis, and 

Picea engelmannii var. mexicana.
206 USDA Forest Service Proceedings RMRS-P-67. 2013

González-Elizondo and others Ecosystems and Diversity of the Sierra Madre Occidental

Figure 1—Regions and subregions of the Sierra Madre Occidental. MA = Madrean Archipelago; M = Madrean; MX = Madrean Xerophylous; MT = 

Madrean with tropical affinities; T = Tropical; M1 = North; M2 = Center; M3 = South.
USDA Forest Service Proceedings RMRS-P-67. 2013 207

Ecosystems and Diversity of the Sierra Madre Occidental González-Elizondo and others
208 USDA Forest Service Proceedings RMRS-P-67. 2013

González-Elizondo and others Ecosystems and Diversity of the Sierra Madre Occidental Table 1—Continued.
USDA Forest Service Proceedings RMRS-P-67. 2013 209

Ecosystems and Diversity of the Sierra Madre Occidental González-Elizondo and others

Figure 2—Vegetation of the Sierra Madre Occidental.
210 USDA Forest Service Proceedings RMRS-P-67. 2013

González-Elizondo and others Ecosystems and Diversity of the Sierra Madre Occidental

Pine-Oak Forests

Covering about 30% of the surface, from 1250 to 3200 m, pine-oak 

forests occur with a wide array of species combinations depending on 

elevation, physiography, and climate: Pinus arizonica, P. engelmannii, and P. chihuahuana with Quercus rugosa and/or Q. gambelii in 

the northern portions of the SMO; P. durangensis, P. teocote, and P. 

cooperi with Q. sideroxyla, Q. crassifolia, Q. rugosa, Arbutus bicolor, 

A. madrensis, and A. arizonica in relatively dry places; or A. tessellata

in warmer areas. Diverse species of Quercus may combine with P. 

engelmannii, P. chihuahuana, and A. arizonica in semi-dry temperate 

areas. On thin soil or outcrops of very unsheltered bedrock, forests 

of P. lumholtzii with Q. radiata and a xeric phase of Q. crassifolia

are prominent. Manzanita (Arctostaphylos pungens), Juniperus deppeana, and J. durangensis are among the common shrubs. On the 

western slopes (MT), P. oocarpa is associated with P. devoniana, P. 

douglasiana, Q. viminea, and Q. gentryi, among others. Pinus herrerae 

dominates on ridges and slopes while on lower areas P. luzmariae is 

on mesas and hillsides frequently with P. lumholtzii and Q. radiata. 

Between 1800–2300 m, Q. jonesii mixes with P. lumholtzii, Q. 

resinosa, Q. crassifolia, and Q. viminea. Q. resinosa mixes with Q. 

praineana and P. oocarpa on lower elevations in Durango, Jalisco, 

and Zacatecas.

Oak Forests

These forests are communities with a dominance of Quercus species but with very different composition, structure, and ecological 

affinities. They cover almost 14% of the surface area of the SMO, 

from 340 m on low hills in the middle of tropical deciduous forest, to 

2900 m in semi-cold climates. Communities dominated by Quercus 

in dry temperate climates are described under the oak woodlands. 

Among the main associations are Q. sideroxyla and Q. rugosa, up to 

2900 m and Q. fulva, Q. mcvaughii or Q. scytophylla below. Q. laeta has 

a broad ecological tolerance, occurring on the western and southern 

flanks of the SMO. In the Madrean Tropical subregion there are mesophytic oak forests with Q. candicans, Q. crassifolia, Q. diversifolia, Q. 

scytophylla, and Clethra spp; at lower elevations, Q. subspathulata

enters the subregion. Other oak forests on the western flanks are those 

of Q. viminea or Q. alamosensis to the north and Q. diversifolia and 

Q. gentryi. Q. albocincta occurs at lower elevations. In the south, 

low forests of robles (Q. magnoliifolia, Q. resinosa) sometimes with 

Q. eduardii or Q. viminea, are found in many areas being substituted 

by Dodonaea scrub.

Mesophytic Forests

These forest communities are rich in their diversity and structure. 

They occupy small areas, covering about 0.14% of the surface area 

on the western flank of the sierra, between 1000 and 2350 m. Among 

their characteristic elements are Magnolia pacifica subsp. tarahumara, 

Ostrya virginiana, Tilia americana var. mexicana, Cedrela odorata, 

Styrax ramirezii, several Lauraceae such as Persea liebmannii, P. 

podadenia, Nectandra spp. and Litsea spp and oaks like Q. candicans, Q. crassifolia, Q. castanea, Q. rugosa, Q. scytophylla, and Q. 

splendens. Other species are Arbutus xalapensis, Garrya laurifolia, 

Cornus disciflora, C. excelsa, Clethra spp, Prunus spp, Peltostigma 

pteleoides, Ilex quercetorum, I. toluccana, Cleyera integrifolia, and 

Brahea aculeata. Often pines and other conifers are present including Pinus maximinoi, P. devoniana, P. douglasiana, P. herrerae, P. 

strobiformis, P. durangensis, Abies neodurangensis or Pseudotsuga 

menziesii. Other communities, such as the Madrean montane meadow 

that occurs in small spots between 2300 and 3100 m, include a high 

proportion of endemic species. 

Oak or Pine-Oak Woodlands

These woodlands cover more than 13% of the surface along the 

eastern foothills and isolated outlying ranges of the SMO at 1450-2500 

m. Physiognomic dominants are pinyon (Pinus cembroides) and/or 

several species of Quercus of xerophylous affinities. 

In northern Sonora and Chihuahua, blue oak (Q. oblongifolia), 

Emory oak (Q. emoryi), or Arizona oak (Q. arizonica) are often 

monodominant or associated with Pinus cembroides, P. chihuahuana, Arbutus arizonica, Q. grisea, and Q. chihuahuensis. Quercus 

hypoleucoides occurs in the uplands and to the west is Q. viminea. 

Common associations on the eastern foothills of the sierra are those 

of P. cembroides and Q. grisea or Q. eduardii. At the transition with 

higher elevation forests, Pinus chihuahuensis, P. engelmannii, and Q. 

durifolia are common. Táscate (Juniperus deppeana) is the common 

shrub; J. coahuilensis enters the region where lower elevation juniper 

scrub contacts higher elevation woodlands.

Weaknesses and Strengths of the Present 

Situation of the Madrean Ecosystems in the 

Sierra Madre Occidental

An accelerated deforestation, fragmentation of habitats, and reduction in forest densities are the most dramatic changes occurring in 

the last decades in the Sierra Madre Occidental. Although sustainable 

forest management is promoted and forest certification has become 

well established in many ejidos, communities, and private lands, illegal 

logging still persists as well as clearing of the forest for agriculture 

or cattle grazing. Forests and woodlands of the SMO have been 

becoming more open and replaced by shrubs (inland chaparral), a 

disturbance linked to perturbation by logging, overgrazing, and fire. 

Also, extreme climatic events in the lower peripheral communities 

are driving changes. A striking example of this was the heavy frosts 

of the winter of 1997 that killed extensive areas of Acacia schaffneri 

and Opuntia scrub on the areas flanking the eastern foothills. This 

was followed by an extreme drought and hot temperatures during the 

1998 spring that reduced many of the oak woodlands at the eastern 

lower slopes.

Other changes occurring in the vegetation cover are (1) the expansion 

of the shrubs Arctostaphylos pungens and Quercus depressipes driven 

by disturbance; (2) the dwindling of oak woodlands that are being 

replaced by hopbush (Dodonaea viscosa), an invader from warmer 

areas; and (3) the effects of bark beetle (Dendroctonus) infestation that 

have killed extensive areas of pines and also affected fir, Douglas-fir 

and spruce that were likely already stressed by drought.

The implementation of sustainable management in recent years is 

allowing conservation of the biodiversity in some places of the sierra. 

CONAFOR’s National Forest Commission programs on payment for 

environmental services (including hydrological, carbon sequestration, 

and conservation of biodiversity) and protection and conservation 

of soils, reforestation and watershed restoration involve ejidatarios, 

comuneros, and private owners in the processes of protection and 

management of ecosystems. Additionally, many communities are 

betting on ecotourism, as well as rural and adventure tourism, which 

represent an alternative to logging and provide time for the forest to 

recover, bringing hope to the ecosystems and people of the SMO.
USDA Forest Service Proceedings RMRS-P-67. 2013 211

Ecosystems and Diversity of the Sierra Madre Occidental González-Elizondo and others

Conclusions

Despite its high biological and cultural diversity, the SMO is 

ironically far from being well known. New records, new species, and 

new genera are still being discovered in areas of the Sierra Madre 

Occidental. Much of the biological treasures of this mountain range 

remain to be discovered and much more exciting exploration is yet 

to be done to find them.

Acknowledgments

Thanks to COFAA IPN for this research stimulus. This work was 

developed with financial support of the projects SIP-20110681 and 

SIP-20120650. We appreciate the suggestions and corrections made 

by Celia López González and Richard Spellenberg on the first version 

of this paper.

References

Bye, R. 1995. Prominence of the Sierra Madre Occidental in the Biological 

Diversity of Mexico. pp. 19-27 in: DeBano, L.F.; Ffolliott, P.F.; Ortega 

Rubio, A; [and others], tech. coords. 1995. Biodiversity and management 

of the Madrean archipelago: The sky islands of southwestern United States 

and northwestern Mexico. Gen. Tech. Rep. RM GTR 264. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest 

and Range Experiment Station. 

CEC (Commission for Environmental Cooperation). 1997. Ecological regions 

of North America: Toward a common perspective. Quebec. 71 p.

Felger, R.S.; Wilson, M.F. 1995. Northern Sierra Madre Occidental and its 

Apachian outliers: A neglected center of biodiversity, pp. 36 59 in: DeBano, 

L.F.; Ffolliott, P.F.; Ortega Rubio, A; [and others], tech. coords. 1995. 

Biodiversity and management of the Madrean archipelago: The sky islands 

of southwestern United States and northwestern Mexico. Gen. Tech. Rep. 

RM GTR 264. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range Experiment Station. 

Felger, R.; Nabhan, G.; Bye, R. 1997. The Apachian/Madrean region of 

southwestern North America and U.S.A. pp. 172-180 in: S.D. Davis, 

V.H. Heywood, O. Herrera-McBryde, J.Villa-Lobos, and A.C. Hamilton, 

eds. Centres for Plant Diversity: A guide and strategy for their conservation, Vol. III, The Americas. Cambridge, UK: The World Wide Fund for 

Nature & International Union for the Conservation of Nature, The World 

Conservation Union.

González-Elizondo, M.; González-Elizondo, M.S.; Ruacho González, L; 

Molina Olvera, M. 2011. Pinus maximartinezii Rzed., Primer registro para 

Durango, Segunda localidad para la especie. Acta Bot. Mex. 96: 33-48.

González-Elizondo, M.S. 1997. Upper Mezquital River Region, Sierra 

Madre Occidental, Mexico. pp. 157-160 in: S.D. Davis, V.H. Heywood, 

O. Herrera-McBryde, J.Villa-Lobos, and A.C. Hamilton, eds. Centres for 

Plant Diversity: A guide and strategy for their conservation, Vol. III, The 

Americas. Cambridge, UK: The World Wide Fund for Nature & International 

Union for the Conservation of Nature, The World Conservation Union.

González-Elizondo, M.S.; López Enriquez, I.L.; Wagner, W.L. 2002. Megacorax gracielanus (Onagraceae), a new genus and species from Durango, 

Mexico. Novon 12: 360 365.

González-Elizondo, M.S; González-Elizondo, M.; Márquez Linares, M.A. 

2007. Vegetación y Ecorregiones de Durango. Plaza y Valdés EditoresIPN. México, D.F. 219 pp.

González-Elizondo, M.S.; González-Elizondo, M.; Tena-Flores, J.A.; RuachoGonzález, L.; López-Enriquez, I.L. (in rev.). Vegetación de la Sierra Madre 

Occidental, México. Una Síntesis. Acta Bot. Mex.

INEGI. 2002. Conjunto de datos vectoriales de la carta de uso del suelo y 

vegetación esc. 1:250 000. Serie III. Instituto Nacional de Estadística y 

Geografía, Aguascalientes.

Luquín S., H.; Nieves H., G.; Ayala R., J.M. 2004. Vegetación del Norte de 

Jalisco, pp. 77-83 in: Vázquez-García, J. A., M. de J. Cházaro B., G. Nieves 

H., Y. L. Vargas-Rodríguez, M. Vázquez G. y A. Flores M. (eds.) Flora del 

Norte de Jalisco y Etnobotánica Huichola. Serie Fronteras de Biodiversidad 

1. Universidad de Guadalajara (CUCBA-CUSH), Guadalajara.

Martin, P. S.; Yetman, D.; Fishbein, M.; Jenkins, P.; Van Devender, T. R.; 

Wilson, R.K. 1998. Gentry ́s Río Mayo Plants: The Tropical Deciduous 

Forest and Environs of Northwest Mexico. The University of Arizona 

Press. Tucson, AZ. 558 p.

Martínez-Yrízar, A.; Felger, R. S.; Búrquez, A. 2010. Los Ecosistemas de 

Sonora: un diverso capital natural. En: Diversidad biológica de Sonora. F. 

Molina Freaner y T. Van Devender (eds.). Pp. 129-156. UNAM/CONABIO, México.

Rzedowski, J. 1978. Vegetación de México. Editorial Limusa. México, D. 

F. 432 p.

Spellenberg, R., Lebgue, T.; Corral-Díaz, R. 1996. A specimen-based, annotated 

checklist of the vascular plants of Parque Nacional “Cascada de Basaseachi” 

and adjacent areas, Chihuahua, Mexico. México, DF. Listados Florísticos 

de México XIII. UNAM-Instituto de Biología. 72 p.

Torres-Morales, L.; García-Mendoza, D. F.; López-González, C.; MuñizMartínez, R. 2010. Bats of northwestern Durango, Mexico: Species richness 

at the interface of two biogeographic regions. The Southwestern Naturalist 

55(3): 347-362.

Van Devender, T.; Reina, A.L. 2005. Floristic comparison of an Arizona Sky 

Island and the Sierra Madre Occidental in eastern Sonora: the Huachuca 

Mountains and the Yécora Area. pp. 154-157 in: Gottfried, G. J., Gebow, 

B. S., Eskew, L. G., and Edminster, C. B. (comps). Connecting mountain 

islands and desert seas: biodiversity and management of the Madrean 

Archipelago II. 2004 May 11-15; Tucson, AZ. Proceedings RMRS-P-36. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station.

Van Devender, T.R., A.L. Reyna G., M.C. Peñalba G. and C.I. Ortega R. 2003. 

The Ciénega de Camilo: a threatened habitat in the Sierra Madre Occidental 

of eastern Sonora, Mexico. Madroño 50(3): 187-195.

Vázquez-García, J. A., M. de J. Cházaro B., G. Nieves H., Y. L. VargasRodríguez, M. Vázquez G. y A. Flores M. 2004. Flora del Norte de Jalisco 

y Etnobotánica Huichola. Serie Fronteras de Biodiversidad 1. Universidad 

de Guadalajara (CUCBA-CUSH), Guadalajara. 182 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
212 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Wide Ranges of Functional Traits in the Flora from the 

Central Region of Sonora: A Diversity to be Explored

César Hinojo-Hinojo, Alejandro E. Castellanos, and Jose M. Llano Sotelo

Laboratorio de Ecofisiología Vegetal, Departamento de Investigaciones Científicas y Tecnológicas de la 

Universidad de Sonora, Hermosillo, Sonora, Mexico

Abstract—Although the Sonoran Desert does not have the highest plant species richness, it has been 

documented with the highest growth form diversity from the North American deserts. It is not known if 

this high growth form diversity could also harbor a high functional diversity. In this study we characterize 

the ecophysiological functional traits of photosynthetic capacity, stomatal conductance, transpiration and 

resource use efficiency (water and nitrogen), as well as morphological and structural traits of specific leaf 

area and nitrogen content in 52 species that inhabit the arid and semiarid Central Region of Sonora. For all 

species and analyzed traits, we found a larger range of values compared to those reported from a recent 

world survey, which represent a very high functional diversity. Such large range and amplitude in functional 

trait values has not been reported for other regions of the world. This study presents evidence of the high 

functional diversity of the Sonoran Desert plants waiting to be described and documented.

Introduction

	 Great efforts have been made to know the functional traits of speciesthat inhabit the different biomes of the world during the past two

decades. This information is important because it allows us to know

and understand the biodiversity, not just from a taxonomic point of

view but as functional diversity or, in other words, the functions,

properties, and possible effects and responses of the species to the

environment. Leaf traits are the most common and important plant

functional traits because they are strongly related to various traits of

other parts of the plants, to species ecological strategy, and even to

various aspects of the ecosystem functioning (Cornwell and others

2008; Craine and others 2005; Díaz and others 2004; Vendramini and

others 2002; Westoby 1998).

	 At a global level, the major effort so far to assess functional traits

of plantspeciesistheTRYdatabase (online in http://try-db.org/) with

over 3million traitrecordsfrom69,000 speciesfromallmajor biomes

in the world. Kattge and others (2011) suggest that it is evident that

plant traits from the warm deserts and, in general, most of the arid

and semiarid zones floras of the world are still poorly represented in

this database.

	 Arid and semiarid zones are important worldwide because they

represent almost 50% of the terrestrial area (without considering the

polar caps). Shreve (1942) and Stebbins (1952) recognized the large

growth forms and trait diversity that existed in the plants from these

ecosystems and the different ways in which they deal with extreme

conditions of water scarcity, high temperatures, and high irradiance

that characterize them. This can mean that arid and semiarid zones

harbor a high functional diversity; however, this issue has been little

explored.

	 The Sonoran Desert, as delimited by Shreve and Wiggins (1964),

shows some climatic and floristic particularities compared to other

deserts oftheworld.It possesses a bi-seasonalrainfall pattern formost

of its extension, with abundant monsoon rains in summer and long

duration and highly unpredictable low intensity rains during winter

(Brito-Castillo and others 2010; Shreve and Wiggins, 1964; Schmidt

1989). Its flora has neartic and neotropical biogeographic influences

(Castellanos 1992), which result in a great number of species that

reach its most northern, southern, or western distribution limits in

this region (Castellanos 1992; Castellanos and others 2010; Van Devender and others 2010). In the Central Region of Sonora, however,

its major affinity is with the highly seasonal neotropical flora from the

tropical deciduous forest ecosystems of Mexico (Rzedowski, 1973).

Moreover, it has been found that the Sonoran Desert possesses the

highest growth form diversity of the NorthAmerican deserts(Shreve

1942; Cody 1991).

	 Due to the poor representation of warm deserts in plant functional

trait studies, and to the climatic and floristic particularities above

mentioned of the Sonoran Desert, the objective of this research was

to conduct a characterization of leaf traits of species that inhabit the

arid and semiarid area of the Central Region of Sonora. We assessed

these through a large sampling in terms of number ofspecies, growth

forms, habitats, and habits. It is in this Central Region of Sonora,

which comprises a gradient of arid and semiarid zones, where desert

and subtropical vegetation integratesin ecotones creating interesting

mosaics in terms of biota composition and physiognomy of plant

communities.

	 The large sampling undertaken in this study was conducted where

both “soft” leaf traits as the specific leaf area, leaf nitrogen content,

and “hard” as various ecophysiological traitsrelated to the photosynthesis and gas exchange were considered (Lavorel and Garnier, 2002)
USDA Forest Service Proceedings RMRS-P-67. 2013 213

Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: . . . Hinojo-Hinojo and others

to represent the functional diversity of the flora from this region in

a variety of leaf traits. It was expected that the species would show

wide ranges in the leaf traits due to the great variety of strategies

and traits of plants that inhabit these environments can develop to

deal with the extreme conditions in the arid and semiarid zones, as

suggested in Stebbins (1952).

Methodology

Study Sites

	 To include the more representative plant communities from the

Central Region of Sonora, 14 localities were selected following a

gradient in vegetation, aridity and disturbance level (table 1). These

sites were located near the limits of the Plains of Sonora and Foothills

Thornscrub subdivisions of the Sonoran Desert (Shreve and Wiggins

1964).

Species Leaf Traits Characterization

	 Samplings in the 14 sites were carried out in September and the

first half of October in 2010 to determine some species leaf traits.An

effort was made to include a wide variety of species, growth forms,

habitats, and habits. The sampling period was in the rainy season to

avoid the limiting effects of the water stress on the photosynthetic

capacity withmeasurementsmade between the 8:00AM and 12:00 PM.

	 On each site and for each species, photosynthesis of leaves at high

irradiances (generally 1500-2000 μmol photons m-2 s-1) was determined at field natural conditions as a measure of the photosynthetic

capacity(Aarea),stomatal conductance towatervapor(gs),transpiration

rate (E), water use efficiency (EUA = Amount of fixed CO2 per unit

of transpired water), and intercellular concentration of CO2 using a

portable photosynthesis system LCi (ADC BioScientific, England)

provided with the broad leaf chamber. Also, the dark respiration rate

(R)was determined forsome species by putting the leavesin complete

darkness during the measurement period. At least three plants per

species were measured, one leaf per plant, with three measurements

per leaf, except in rare occasions where it was not possible to sample

enough individuals. Each measured leaf was collected in paper envelopes and stored in ice for its transportation. The real area used in

the measurements of gas exchange was determined using the image

analyzingsoftware ImageJ1.43u(NationalInstitutesofHealth,USA).

This value was used to correct the gas exchange measurements.

	 After the measurement of leaf area, each leaf was oven-dried at 70

oC for 2 daysto determine leaf dry weight. Using the leaf area and dry

weight, the specific leaf area (SLA) was calculated. For those same

leaves, nitrogen content per leaf weight (Nmass) was determined by

the fenate method for Kjeldahl total nitrogen using a Digestor 2020

(Tecator, Spain) and a rapid flow analizerRFA(ALPKEM).The measure ofspecific leaf area was used as a conversion unit for expressing

some leaf traits on a mass and area basis (mass and area subscripts).

The nitrogen use efficiency was calculated for each measure of Aarea

as the Aarea/Narea quotient. The complete list of leaf traits measured

and their corresponding units are listed in the table 2.

Data Analysis

	 All the data were screened and those that showed any evidence

of stress limitation weren’t included in the analysis (e.g., stomatal

conductance values below 0.05 mol H2O m-2 s-1 (Flexas and others

2006) or individual trait values far from the trend of the rest of the

values for the same species). A mean value for each measured leaf

trait was obtained with the remaining data for each species. These

mean values are those reported in the paper and compared with all

of the world data (TRY database in Kattage and others 2011).

Results and Discussion

Leaf Traits of Species in the Central Region of 

Sonora

	 A total of 52 species was sampled that represent a wide variety

of growth forms (with the exception of plants with CAM photosynthetic pathway) including those species that are the most commonly

found in the flora of the Central Region of Sonora. All species were

sampled under natural conditions except for Cucurbita digitata, C. 

palmeri, Jatropha cinerea and the exotic ruderal Ricinus communis,

which were sampled under irrigated conditions. Photosynthesis and

gas exchange data were obtained for 51 species. Only Ipomoea nil

Table 1—Geographic location, vegetation type, and annual rainfall for the study sites.

Site Latitude Longitude Vegetation typea Annual rainfallb

1 N 29.023° O 111.137° Abandoned agricultural field 200-300 mm

2 N 28.966° O 110.961° Subtropical scrub 200-300 mm

3 N 28.700° O 110.540° Subtropical scrub 400-500 mm

4 N 28.710° O 110.540° Induced buffel savanna 400-500 mm

5 N 29.563° O 111.011° Disturbed mezquital 300-400 mm

6 N 29.684° O 110.145° Disturbed mezquital 500-600 mm

7 N 29.559° O 110.123° Disturbed mezquital 500-600 mm

8 N 29.474° O 110.217° Subtropical scrub 500-600 mm

9 N 29.465° O 110.248° Subtropical scrub 500-600 mm

10 N 29.439° O 110.315° Riparian vegetation 400-500 mm

11 N 29.435° O 110.334° Mezquital 500-600 mm

12 N 29.346° O 110.497° Desert microphyllous scrub 400-500 mm

13 N 29.327° O 110.561° Riparian vegetation 400-500 mm

14 N 29.240° O 110.726° Mezquital 300-400 mm

a

Following the INEGI's land use and vegetation classification for Mexico

b

Following the isohyets map of Mexico by CONABIO. Online at: http://www.conabio.gob.mx/informacion/gis/
214 USDA Forest Service Proceedings RMRS-P-67. 2013

Hinojo-Hinojo and others Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: . . .

did not give reliable data because plants were understress. However,

its leaves were included in the data for specific leaf area (SLA) and

nitrogen content (N). Data were obtained on dark respiration (R) for

21 species. The SLAcould be determined for all the species, while N

and nitrogen use efficiency (NUE) could be determined for 48 and 46

species respectively. Compared to previous data (Wright and others

2004; Kattge and others 2011), this study had the largest number of

species with an ecophysiological characterization from an arid or

semiarid region and from the Sonoran Desert.

	 Trait mean values for all species grouped by growth form are

shown in table 3. Evergreen trees presented the lowest values of gas

exchange and SLA traits, herbaceous and grass species had the highest gas exchange values, vines had the highest SLA and high Amasa,

Nmass and gs but low Aarea, , and shrubs had intermediate values on all

traits. These patterns had the most traits that agree with previously

reported valuesfor these groups of plants(Larcher 1995; Castellanos

1991; Castellanos and others 1989).

Comparison of Leaf Traits From Species of the 

Central Region of Sonora with Those of the 

World

	 Severalstudies have addressed the issue ofthe leaffunctional traits

of species in different biomes of the world and, therefore, may serve

to give a meaning to the values found in the species of the Central

Region of Sonora. For example, the mean values of SLAand N of the

species of the present study (fig. 1) agreed with values reported for

species with the lowest leaf life spans (between 1-4 months) (Reich

and others 1992; 1997), which may be related to the short seasonal

character of species in the Central Region of Sonora. These values

may be considered high in a global context of the species and in particular for those that could be expected in an arid region where the

trend for species that inhabit the warmest and driest regions have the

lowest SLA (Wright and others 2004) in contrast with our findings

in this study. Wright and others (2005) report that species with the

highest irradiances and lowest annual rainfall habitat conditions had

highest values in Narea, which agrees with the high values found in

the studied species, although in this study we also report high values

of Nmass (fig. 1).

	 It is evident that all leaftraitsfrom the speciesthat we studied had a

similar or larger range of valuesthan those reported for all the species

in the world that have been studied (fig. 1) in comparing the data for

the species that we studied with those of the global compendium of

the TRY database functional traits (Kattge and others 2011). In fact,

the maximum value in all the trait ranges exceeds the value of the

97.5% quantile from the data of TRY database. Ranges this wide are

not common and we have not found such amplitude in the literature

from other ecosystems or regions of the world. According to Tilman’s (2001) definition of functional diversity, those components of

biodiversity that influence how an ecosystem operates or functions

that we found for both ecophysiological and morphostructural traits

or “hard” and “soft” traits sensu (Lavorel and Garnier 2002) in species that inhabit the arid and semiarid zones of the Central Region

of Sonora represent, without any doubt, the high functional diversity

that exists in the flora of the southwestern Sonoran Desert.

	 Previous studies that have documented a high biological diversity

differentfromthe taxonomic diversity in the SonoranDesert are those

of Shreve (1942) and more recently Cody (1991, which point out the

highestgrowthformdiversityoftheNorthAmericandeserts.Nilsenand

others(1984)documentedawidediversityinecophysiologicstrategies

in water use within a single functional type, the phreatophytes of the

Sonoran Desert. Stebbins (1952) explains that the plant diversity in

sites with aridity may be high, where an important factor is the high

environmental(resources) andtopographicalheterogeneity,butplants

show a wide variety of ways (strategies) and structures (organs) that

can make it possible for a plant to inhabit arid conditions, which can

be interpreted as a high functional diversity.

	 Afew effortsto understand the causes of high biological and functionaldiversityoftheSonoranDeserthavebeenreportedwhile amajor

emphasis has gone to describe its taxonomic diversity (Shreve and

Wiggins,1964;Turner and1995;Molina andVanDevender2010).We

propose that the high diversity (e.g., functional diversity and growth

form diversity) is found in the Central Region of Sonora may be due

a number of factors including (1) the floristic affinity, influence and

confluence of different contiguous biogeographic (Neartic and Neotropical) regions (Rzedowski, 1973; Castellanos, 1992; Castellanos

Table 2—Leaf traits characterized for the species, its abbreviations and measurement units.

Leaf trait Abbreviation Units

Specific leaf area SLA m2 kg-1

Photosynthetic capacity per leaf area Aarea μmol CO2 m-2 s-1

Photosynthetic capacity per leaf dry mass Amass nmol CO2 g-1 s-1

Transpiration rate E mmol H2O m-2 s-1

Stomatal conductance to water vapor gs mol H2O m-2 s-1

Dark respiration per leaf area Rarea μmol CO2 m-2 s-1

Dark respiration per leaf dry mass Rmass nmol CO2 g-1 s-1

Leaf nitrogen content per leaf area Narea g N m-2

Leaf nitrogen content per leaf dry mass Nmass %

Nitrogen use efficiency NUE μmol CO2 g-1 N s-1

Water use efficiency WUE μmol CO2 mmol-1 H2O
USDA Forest Service Proceedings RMRS-P-67. 2013 215

Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: . . . Hinojo-Hinojo and others
216 USDA Forest Service Proceedings RMRS-P-67. 2013

Hinojo-Hinojo and others Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: . . .

Figure 1—Comparison of leaf trait ranges between the species of the Central Region of Sonora (CRS) and the the world species included in 

the TRY database (TRY). Data points for the TRY species correspond to the 2.5% quantil (lowest point), mean value (mid point), and 97.5% 

quantil (highest point), and for the CRS species are the minimum (lowest point), mean (mid point) and maximum values (highest point).
USDA Forest Service Proceedings RMRS-P-67. 2013 217

Wide Ranges of Functional Traits in the Flora from the Central Region of Sonora: . . . Hinojo-Hinojo and others

and others 2010; Van Devender, 2010); (2) a large environmental

heterogeneity in a short distance (Castellanos and others 1992); and

(3) the interspecific positive ecological interactions(e.g., facilitation,

nursing, symbiosis, mutualism, fertility islands, etc.) that can allow

that species with different ecological strategies and stress tolerance

levels to coexist in the same ecosystem.

Conclusions

	 Ecophysiologic and morphostructural leaftraits of 52 species were

characterized,whichmakesthisstudythelargestfortheecophysiologic

characterization from an arid and semiarid and from the Sonoran

Desert species. The ranges found in all leaf traits characterized for

the studied species of the Central Region of Sonora had a similar and

even larger range of values than those reported for all the species in

the world. This wide amplitude in the ranges of leaf trait values is an

indicator of a very high functional diversity in the speciesthat inhabit

the arid and semiarid gradient of the Central Region of Sonora. This

work is evidence ofthe high functional diversity in theSonoranDesert

plants waiting to be described and documented.

Acknowledgments

	 We want to acknowledge and thank the National Institute of Ecology and British Embassy in Mexico (BIO3 project) and CONACYT

(61865) for their support (A. C.) and in the form of a scholarship

(C. H.) during the course of these studies. Also, thanks to Dra. Clara

Tinoco Ojanguren for her support during the study and her review

of this paper.

References

Brito-Castillo, L.; Crimmins, M. A.; Díaz, S. C. 2010. Clima. In: MolinaFreaner, F.E. andVan Devender,T.R, eds, DiversidadBiológica de Sonora.

México D. F. UNAM: 73-96.

Castellanos, A. E. 1991. Photosynthesis and gas Exchange of vines. In: Putz,

F. E.; Mooney, H. A. (eds.), The Biology of Vines. Cambridge. Cambridge

University Press. 171-214.

Castellanos,A. 1992.Ecología, utilizacion y conservacion de las comunidades

vegetales enel estadodeSonora:unanalisis.In:Moreno,J.L., ed. Ecología,

Recursos Naturales y Medio Ambiente en Sonora. Hermosillo. Secretaria

de Infraestructura Urbana y Ecología: 25-37.

Castellanos, A. E.; Mooney, H. A.; Bullock, S. H.; [and others] 1989. Leaf,

stem, and metamer characteristics of vines in a tropical deciduous forest

in Jalisco, Mexico. Biotropica. 21(1): 41-49.

Castellanos-Villegas, A. E.; Bravo, L. C.; Koch, G. W.; [and others]. [In

press]. Impactos ecológicos por el uso del terreno en el funcionamiento de

ecosistemas áridos y semiáridos. 157-186 p.In: Molina-Freaner, F.E. yVan

Devender,T.R., eds. DiversidadBiológicadeSonora.MéxicoD.F.UNAM.

Cody, M. L. 1991. Niche theory and plant growth form. Plant Ecology.

97(1):39-55.

Cornwell, W. K.; Cornelissen, J. H. C.; Amatangelo, K.; [and others]. 2008.

Plantspeciestraits are the predominant control on litter decomposition rates

within biomes worldwide. Ecology Letters. 11: 1065-1071.

Craine, J. M.; Lee, W. G.; Bond, W. J.; Williams, R. J.; Johnson, L. C. 2005.

Environmental constraints on a global relationship among leaf and root

traits of grasses. Ecology. 86(1): 12-19.

Díaz, S.; Hodgson, J. G.; Thompson, K.; [and others]. 2004. The plant traits

that drive ecosystems: Evidence from three continents. Journal of Vegetation Science. 15: 295-304.

Flexas,J.;Bota,J.;Galmés,J.;Medrano,H.;Ribas-Carbó,M. 2006.Keeping a

positive carbonbalanceunder adverse conditions:responsesofphotosynthesis and respiration to waterstress. Physiologia Plantarum. 127(3): 343-352.

Kattge, J.; Díaz, S.; Lavorel, S.; [and others]. 2011. TRY—a global database

of plant traits. Global Change Biology. 17(9): 2905-2935.

Larcher, W. 1995. Physiological Plant Ecology. 3rd ed. Berlin-Heidelberg:

Springer-Verlag.

Lavorel, S.; Garnier, E. 2002. Predicting changes in community composition

and ecosystem functioning from plant traits: revisiting the Holi Grail.

Functional Ecology. 16(5): 545-556.

Molina-Freaner, F. E. y Van Devender, T. R.2010. Diversidad Biológica de

Sonora. México D. F. UNAM.

Nilsen,E.T.;Sharifi,M.R.;Rundel,P.W.1984.Comparativewaterrelationsof

phreatophytesintheSonoranDesertofCalifornia.Ecology.65(3):767-778.

Reich, P.B.,Walters, M.B.; Ellsworth, D. S. 1992. Leaflife-span in relation to

leaf, plant, and stand characteristics among diverse ecosystems. Ecological

Monographs. 62(3): 365-392.

Reich, P. B., Walters, M. B.; Ellsworth, D. S. 1997. From tropics to tundra:

Global convergence in plant functioning. Proceedings of the National

Academy of Sciences. 94: 13730-13734.

Rzedowski, J. 1973. Geographical relationships of the flora of mexican dry

regions. 61-72 p. En: Grahan,A(Ed.), Vegetation and Vegetational History

of Northern Latin America. Elsevier. Amsterdam.

Shreve, F. 1942. The desert vegetation of North America. The Botanical

Review. 4(3): 195-246.

Shreve, F.; Wiggins, I.L. 1964. Vegetation and Flora of the Sonoran Desert.

Part I: Vegetation of the Sonoran Desert. Original Edition. Stanford: Stanford University Press.

SchmidtR.H.,Jr. 1989.The arid zones ofMexico: climatic extremes and conceptualization ofthe Sonoran Desert.Journal ofArid Environments. 16: 241-256.

Stebbins, G. L. 1952. Aridity as a stimulus to plant evolution. American

Naturalist. 86: 33-44.

Tilman, D. 2001. Functional diversity. In: Levin, S. A., ed. Encyclopedia of

Biodiversity. Vol. 3. San Diego: Academic Press: 109-120.

Turner, R. M.; Bowers, J. E.; Burgess, T. L.; Hastings, J. R. 1995. Sonoran

DesertPlants:AnEcologicalAtlas.Tucson.TheUniversityofArizonaPress.

Van Devender,T. R.; Felger, R. S.; Fishbein, M.; [and others]. 2010. Biodiversidad de plantas vasculares. In: Molina-Freaner, F. E. and Van Devender, T.

R., eds. Diversidad Biológica de Sonora. México D. F.: UNAM: 229-255.

Vendramini,F.;Díaz,S.;Gurvich,D.E.;Wilson,P.J.;Thompson,K.;Hodgson,

J. G. 2002. Leaf traits as indicators of resource-use strategy in floras with

succulent species. New Phytologist. 154: 147-157.

Westoby, M. 1998. A leaf-height-seed (LHS) plant ecology strategy scheme.

Plant and Soil. 199: 213-227.

Wright, I. J., Reich, P. B.; Westoby, M.; [and others]. 2004. The worldwide

leaf economics spectrum. Nature. 428: 821–827.

Wright,I.J.,Reich, P.B.;Cornelissen,J. H.C.;[and others]. 2005.Modulation

of leaf economic traits and trait relationships by climate. Global Ecology

and Biogeography. 14: 411–421.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
218 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Cytogeography of Larrea tridentata at the ChihuahuanSonoran Desert Ecotone

Robert G. Laport and Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

Abstract—The long separation of the Chihuahuan and Sonoran Deserts is reflected in the high species richness 

and endemism of their floras. Although many endemic species from both deserts reach their distributional 

limits where the Sierra Madre Occidental massif fragments into smaller mountain complexes in northern 

Mexico and adjoining areas of the United States, indicator species approaches to identify the boundary between the Chihuahuan and Sonoran Deserts are complicated by differences in physiology and morphology 

among species that independently influence their local occurrences. Here we use an alternate approach that 

examines distributional differences within a single species that is widespread in both deserts. We find that 

diploid and tetraploid Larrea tridentata occur sympatrically at an abrupt boundary in the low elevation areas 

of the Gila and San Pedro Rivers of southeastern Arizona. This cytotype boundary is correlated with climatic 

and vegetational transition, is consistent with a boundary between the Chihuahuan and Sonoran Deserts first 

suggested by Shreve, and raises interesting questions about the historical persistence of this contact zone.

Introduction

The Chihuahuan and Sonoran Desert biomes of North America 

harbor species-rich, highly endemic floras that reflect a long isolation 

maintained along most of their boundary by the high elevations of 

the Sierra Madre Occidental. At present, species from both deserts 

come into contact only in the lowland areas where the Sierra Madre 

Occidental massif fragments into smaller mountain complexes in 

northern Mexico and adjoining areas of the United States (Warshall 

1994). Species of plants (Henrickson and Straw 1976; Mabry and others 1977; McLaughlin 1986; Shreve 1942, 1951), mammals (Riddle 

1998; Riddle and others 2000), reptiles (Morafka 1977), and bees 

(Danforth 1994; Rozen 1992) from each desert have been documented 

to reach their distributional limits in this area. These biological discontinuities coupled with abrupt changes in several climatological 

measures (Ackerman 1941; MacEwen and others 2005; Russell 1931; 

Schmidt 1979) support the presence of a transition zone between the 

Chihuahuan and Sonoran Deserts.

Although there is a general consensus that the boundary between the 

Chihuahuan and Sonoran Deserts occurs where vegetation transitions 

to a high elevation grassland community in southeastern Arizona, 

southwestern New Mexico, and northeastern Sonora, uncertainty persists over the formal location of the eastern boundary of the Sonoran 

Desert within this region (Gehlbach 1967; Schmidt 1979; Thornthwaite 1948; Weaver and Clements 1938). This uncertainty stems in 

part from the use of species assemblages, or select plant species to 

identify a boundary. However, different physiological and morphological traits among species invariably result in local differences in their 

occurrence, which are accentuated by the complex climatological and 

topographic variability of the “Sky Islands” area. Thus, the location 

of the eastern boundary of the Sonoran Desert has depended on the 

choice of indicator species and/or the definition of desert employed. 

For example, Shreve (1942, 1951) argued that the distributional limits of Saguaro Cactus (Carnegiea gigantea) and Ironwood (Olneya 

tesota) were closely associated with the edge of the Sonoran Desert 

as it transitions to the intervening grasslands currently separating the 

Chihuahuan and Sonoran Deserts. In contrast, Schmidt (1979) used 

measures of aridity to suggest the Chihuahuan and Sonoran Deserts 

were connected and shared a boundary in southeastern Arizona.

An approach not previously used to investigate the ChihuahuanSonoran Desert boundary is to examine genetic or distributional 

discontinuities within a single species of plant that inhabits both 

deserts. Single-species analyses should generally enjoy the advantage 

of controlling for the variable environmental effects on distantly 

related indicator species (Gleason 1926, 1939). Abrupt breaks in 

the distribution of a species, or discontinuities in DNA molecular 

markers, may indicate significant geological or environmental shifts. 

For example, Riddle (2000) demonstrated that species from multiple 

taxonomic groups in the North American deserts comprised divergent 

mitochondrial DNA lineages. Divergence of these phylogroups was 

found to correlate with past inundations that bisected the peninsula 

for species with distributions on the Baja Peninsula. We adopted 

a similar approach in this study using cryptic genetic variation to 

identify distributional shifts in the North American creosote bush 

(Larrea tridentata (DC) Coville; Zygophyllaceae).

Larrea tridentata is a ubiquitous shrub found throughout the 

Chihuahuan, Sonoran, and Mojave Deserts (Barbour 1969; Benson 

and Darrow 1981; Hunter and others 2001; Laport and others 2012; 

Lewis 1980; Turner and others 1995; Yang 1967, 1970; Yang and 

Lowe 1968). The distribution of L. tridentata has been used to define 

the extent of these deserts (Hastings and Turner 1965; Hunziker and 

others 1977; Shreve 1942) and its macrofossils are used to infer past 

desert conditions (Betancourt and others 1990). Yang (1967, 1970) 
USDA Forest Service Proceedings RMRS-P-67. 2013 219

Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone Laport and Minckley

documented the existence of three chromosome races of creosote bush 

in North America. Recent analyses using guard cell measurements 

(Hunter and others 2001) and flow cytometry (Laport and others 

2012) demonstrated that populations of tetraploids (2n = 4x = 52) 

and hexaploids (2n = 6x = 78) intermingle in the northern Sonoran 

Desert, while diploids (2n = 2x = 26) are limited to the Chihuahuan 

Desert and only hexaploids are known from the Mojave Desert.

As part of a broader study of cytotype distributions, Laport and 

others (2012) identified an abrupt transition between diploids and 

tetraploids near the classically defined boundary of the Sonoran 

Desert. However, it remains unclear if the diploid and tetraploid 

cytotypes are restricted to the Chihuahuan and Sonoran Deserts, 

respectively, due to the low resolution of prior sampling. Here, we 

describe additional sampling across this di ploid-tetraploid cytotype 

transition and examine evidence for the presence of a concordant 

ecological gradient supporting a transition from the Chihuahuan to 

Sonoran Desert. We measured plant community composition at sites 

spanning the cytotype transition and used multi-decadal climate data 

to determine if the cytotype transition is concordant with biotic and 

abiotic change. Finally, we constructed Ecological Niche Models, a 

widely used method for generating hypotheses of species distributions 

(Hijmans and others 2003; Kozak and Weins 2006; Oberhauser and 

Peterson 2003; Roura-Pascual and others 2004) to test if bioclimatic 

data predict the boundary between L. tridentata cytotypes. These data 

were then used to discuss the potential relationship of this transitional 

area to the historical extent of the Chihuahuan and Sonoran Deserts.

Methods

Details of the cytotye distributions discussed here can be found 

in Laport and others (2012). Here, we employed 15 diploid and 13 

tetraploid sites from areas surrounding the Gila and San Pedro River 

valleys of southeastern Arizona, USA (table 1; fig. 1). These sites, as 

reported in Laport and others (2012), were established in spring 2007 

and 2008. The cytotype of 5-50 individuals per site was inferred using 

flow cytometry (Laport and others 2012). Discovery of the transition between diploid and tetraploid Larrea tridentata prompted the 

establishment of seven additional sites (T1Q1, T1Q2, T1Q3, T1Q4, 

T1Q5, T6Q1, T6Q2; table 1) along the Gila and San Pedro Rivers in 

2008, creating two sampling transects between known diploid and 

tetraploid sites. Each plant (650 in total) was permanently marked 

with a numbered aluminum tag, and the GPS coordinates of each site 

were recorded (table 1).

To examine if other elements of the plant community changed 

in conjunction with the transition from diploid to tetraploid Larrea 

tridentata, we measured vegetation attributes at five sites along the 
220 USDA Forest Service Proceedings RMRS-P-67. 2013

Laport and Minckley Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone

San Pedro River and four sites along the Gila River (tables 1, 2). At 

each site, a representative 25 m x 25 m sampling plot was established 

in spring 2010 and the total number of each woody perennial species 

present was recorded (table 2). The vegan (v2.0.3) CRAN package 

for community ecology analyses implemented in the R statistical 

package (www.r-project.org) was used to conduct Nonmetric Multidimensional Scaling (NMDS) ordinations on the species density data. 

NMDS ordinations were implemented on square-root transformed 

and Wisconsin double standardized data. A two-dimensional solution 

with stress = 0.05192 was obtained using Bray-Curtis dissimilarities 

and 20 random starts. This solution was selected after inspection of 

a dimensionality vs. stress plot revealed that stress values reached 

Figure 1—Ecological Niche Model projections of (A) tetraploid and (B) diploid L. tridentata based upon collections near the Chihuahuan/Sonoran 

Desert transition. Relief map (C) displaying L. tridentata collections in the San Pedro River and Gila River drainages of SE Arizona. Diploid sites are 

represented by white squares; tetraploid sites are represented by black circles. Warmer colors in the niche model projections represent higher predicted 

suitability. The northern range of L. tridentata (D) adapted from Hunziker et al. 1977. Star denotes location of study region.
USDA Forest Service Proceedings RMRS-P-67. 2013 221

Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone Laport and Minckley

an asymptote at solutions with higher dimensionality (McCune and 

Grace 2002).

To test if climate differed between diploid and tetraploid sites, we 

obtained climate values from the closest (mean 21.4 km, range 1–45 

km) National Oceanic and Atmospheric Administration (www.noaa.

gov) weather stations to 12 diploid and 13 tetraploid sites (table 1). 

Some sites could not be paired with unique weather stations and were 

excluded from climate analyses. We obtained 30-year averages (longest 

available for all sites) for January maximum temperature, January 

minimum temperature, July maximum temperature, July minimum 

temperature, and mean annual precipitation for Analysis of Variance 

(ANOVA) models and Principal Components Analysis (PCA) using 

the JMP statistical package (v9; SAS Institute, Cary, North Carolina, 

USA). Altitudes were obtained from the USGS National Map (www.

nationalmap.gov), as were political boundaries and bedrock geology 

for qualitative assessments of cytotype distributions in Quantum GIS 

(www.qgis.org) (fig. 1).

We modeled cytotype distributions using the occurrence data in 

table 1 to examine if niche models supported the observed boundary of diploid and tetraploid Larrea tridentata. Altitude and climate 

variables at 30 arcsecond resolution (“bioclim”) covering the 

geographic area of the L. tridentata localities listed in table 1 were 

obtained from Worldclim (www.worldclim.org; Hijmans and others 

2005). To reduce inclusion of climate parameters from areas outside 

the distribution of L. tridentata and over-prediction of occurrence, 

the bioclim data were trimmed to encompass only the distribution 

of sampling localities detailed in table 1 (Pearson and others 2007; 

Stohlgren and others 2011). To minimize the high correlation among 

the 19 bioclim variables we reduced them to the six most dominant 

variables via Principal Component Analysis (PCA) on site-specific 

bioclim data (obtained using the “query” tool in Quantum GIS). 

The six variables retained were those most closely associated with 

the first six Principal Components: mean temperature of the wettest 

quarter, annual precipitation, mean temperature of the coldest quarter, precipitation of the driest quarter, temperature annual range, and 

seasonality of precipitation, respectively.

We used the program Maxent (v3.3.0b; www.cs.princeton.

edu/~schapire/maxent/) with default parameter settings to construct 

a probabilistic niche model prediction for each cytotype (Phillips 

and others 2006). We ran 20 replicate models that were averaged via 

the crossvalidation method. Niche models for diploid and tetraploid 

Larrea tridentata were projected onto a broader region encompassing 

the “Sky Islands” and surrounding areas, and qualitatively reviewed in 

Quantum GIS (fig.1). Following Warren and others (2008) and Glor 

and Warren (2011), we also assessed the consistency and accuracy 

of the averaged predictions via the receiver operating characteristic 

(ROC) and the area under the ROC curve (AUC). High AUC values 

are generally indicative of models that predict occurrences well (Phillips 

and others 2006).

Results

Diploid and tetraploid sites were found in both the Gila and San 

Pedro River valleys. All individuals in four of the added transect sites 

(T1Q1, T1Q2, T1Q5, T6Q2) were diploid and all individuals in two 

of the sites (T1Q4, T6Q1) were tetraploid (table 1). A mixed site of 

36 diploid and 14 tetraploid individuals was found near the center of 

the San Pedro River transect (T1Q3). Excluding this mixed-cytotype 

site (T1Q3), diploid and tetraploid sites were separated by ca. 7 km 

along the San Pedro River transect while diploid and tetraploid sites 

were separated by ca. 18 km along the Gila River transect.

All plant species recorded in the Gila and San Pedro River transect 

sites had Sonoran Desert affinities (table 2) but vegetation ordination 

separated diploid and tetraploid sites (fig. 2). Tetraploid sites were 

associated with high densities of Parkinsonia spp., Cylindropuntia 

spp., L. tridentata, and Prosopis velutina. Diploid sites were associated 

with high densities of Calliandra eriophylla, Carnegiea gigantea, 

Encelia farinosa, Ephedra spp, Fouqueria splendens, Acacia greggii, 

Opuntia spp., Prosopis velutina, and Simmondsia chinensis. Diploid 

sites tended to have higher densities of Arizona Upland species than 

tetraploid sites.

Multiple ANOVA (F = 3.257, P < 0.001) and PCA using all climate 

data suggested diploid and tetraploid sites differed significantly (fig. 3). 

Figure 3—PCA of NOAA 30-year average climate data collected for diploid 

and tetraploid sites. Diploid sites denoted as open squares, tetraploid 

sites denoted as black circles.

Figure 2—Non-metric Multi-Dimensional Scaling species density ordination of diploid and tetraploid sites. Diploid sites denoted by open squares, 

tetraploid sites denoted by black circles, the single mixed-cytotype site 

is denoted by a black square.
222 USDA Forest Service Proceedings RMRS-P-67. 2013

Laport and Minckley Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone

PC1 was associated with July maximum temperature (61.3% of 

variation), and PC2 was associated with mean annual precipitation 

(21.7% of variation). The first four Principal Components—July 

maximum temperature, mean annual precipitation, altitude, and 

January maximum temperature, respectively—accounted for 98.1% 

of the variation. Diploid sites tended to be at higher elevations (F = 

31.6203, P < 0.001) and to have lower average temperatures (January 

minimum temperature, F = 9.8483, P = 0.005) (fig. 4).

Qualitatively, the niche models for diploid and tetraploid Larrea 

tridentata corresponded to the reported ranges for the individual 

cytotypes (Barbour 1969; Felger 2000; Hunter and others 2001; 

Hunziker and others 1977; Laport and others 2012; Turner and others 

1995; Yang 1967; 1970). High average AUCs were obtained for both 

diploid (0.888, 0.178 s.d.) and tetraploid (0.893, 0.067 s.d.) niche 

models. The mean temperature of the wettest quarter made the largest 

contribution as an explanatory variable in the diploid model (39.8%), 

and mean temperature of the coldest quarter made the next largest 

contribution (29.8%). In the tetraploid model, mean temperature of 

the coldest quarter made the largest contribution (61.2%) and mean 

temperature of the wettest quarter made the next largest contribution 

(33.6%). Although diploids were predicted to have suitable habitat 

in some of the higher elevations of the Sonoran Desert, tetraploid 

suitability was essentially restricted to the Sonoran Desert (fig. 1). 

Nevertheless, diploid and tetraploid L. tridentata were predicted to 

share a boundary in the low elevations of the Gila and San Pedro River 

drainages, the same area where this transition is actually observed 

(fig. 1).

Discussion

Here, we documented an abrupt transition between diploid and 

tetraploid Larrea tridentate, a characteristic desert species, over the 

span of approximately 7-18 kilometers in the San Pedro and Gila 

River valleys of southeastern Arizona (fig. 1). A single site where 

diploids and tetraploids co-occur was discovered in the San Pedro 

River valley, an observation not previously reported. Although Yang 

(1970) reported the discovery of diploids and tetraploids within a. 

5-mile radius near Mountain View, Arizona, it is not clear if they were 

growing sympatrically.

Differences in plant community suggest that our Gila and San 

Pedro River sites occurred near the eastern boundary of the Sonoran 

Desert (Bahre 1995; Betancourt and others 1990; McL aughlin 1986; 

Schmidt 1979; Shreve 1942, 1951; Turner and others 1995). Vegetation 

ordination results indicate diploid and tetraploid sites differ somewhat, 

though not strongly, in species composition. For example, tetraploid 

sites had higher densities of Larrea tridentata, while diploid sites 

tended to have higher densities of Arizona Upland species such as

Calliandra eriophylla, Encelia farinosa, and Simmondsia chinensis. 

The mixed-cytotype site grouped with sites where tetraploids occur 

(fig. 2). Surprisingly, diploid sites also had higher densities of the 

Sonoran Desert endemic Carnegiea gigantea, a species typically used 

as an indicator of Sonoran Desert habitat (Shreve 1951; Soule and 

Lowe 1970). However, the study area is located only a few kilometers 

north and west of the documented range limit of C. gigantea, as the 

Chihuahuan endemic Flourensia cernua (Benson 1969; Benson and 

Darrow 1981; Brown 1994; Laport personal observation; Zimmerman 

1969). These species have been used variously to denote the boundaries 

of the Sonoran and Chihuahuan Deserts, respectively (Shreve 1951; 

Zimmerman 1969). The higher densities of C. gigantea in diploid 

sites may be attributable to local variation in their occurrence, or the 

placement of sampling plots, which could be corrected with more 

intensive vegetation sampling. Nevertheless, these observations can 

be interpreted as indicating that the study area resides in an Arizona 

Upland plant association (sensu Brown 1994) near the boundary 

between the Sonoran Desert and a higher elevation grassland community (sensu Burgess 1995), and that the diploid cytotype may not 

be entirely restricted to the Chihuahuan Desert and its surrounding 

semidesert habitat.

Altitude, precipitation, and temperature data in addition to our 

ecological niche models constructed from bioclim variables indicate 

abiotic differences occur at or near the range limits of diploid and 

tetraploid Larrea tridentata. The northwestern part of the study area 

is at lower elevation than the southeastern part of the study area. 

Tetraploid L. tridentata predominate in the northern and western 

portion of the study region while diploids were found only on the 

southeastern portion of the study area. Although our niche models 

predict some overlap between the diploid and tetraploid ranges (figs. 

1, 2, 3), the overall pattern obtained from the altitude/climate data 

and niche models suggests areas inhabited by these two cytotypes 

are abiotically unique with warmer temperatures prevailing in areas 

where tetraploids are found. The differences observed here presumably 

reflect more general climatic differences between the eastern Sonoran 

Desert and the adjacent high elevation grassland biomes, and concur 

remarkably well with prior investigations of climatic differences at 

the boundary of the Sonoran and Chihuahuan Deserts (Ackerman 

1941; MacEwen and others 2005; Russell 1931; Schmidt 1979). 

Despite high AUCs indicative of “good” models, the diploid model 

under-predicts the actual range of L. tridentata in SE Arizona and SW 

New Mexico (fig. 1B). Pearson and others (2007) suggested that niche 

models with limited sampling might conservatively be interpreted as 

indicating areas of similar climatic conditions to the known localities, 

rather than a model of species range limits. Nevertheless, a broader 

scale analysis of L. tridentata spanning all three North American 

warm deserts recapitulates the niche model differences reported here, 

suggesting the limited sampling of this study does not overtly bias 

niche model construction and projection (Laport et al. unpublished).

The results of our analyses contribute to the body of literature 

investigating the limits of the Chihuahuan and Sonoran Deserts. The 

transition between diploid and tetraploid Larrea tridentata over a few 

kilometers is generally concordant with the distributional limits of 

some characteristic Chihuahuan and Sonoran endemic species (i.e., 

Figure 4—Boxplots of altitude and NOAA 30-year average climate data 

for diploid and tetraploid L. tridentata sites.
USDA Forest Service Proceedings RMRS-P-67. 2013 223

Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone Laport and Minckley

F. cernua, C. gigantea) as well as a subtle climatic gradient (Benson 

and Darrow 1981; Schmidt 1979). Given the patterns observed in this 

relatively small-scale analysis, it would be informative to include additional sites and their physical features in the transition area for future 

analyses. We doubt soils explain cytotype distributions in this study 

because of the predominance of unconsolidated alluvial substrates 

in these river valleys. However, prior studies have demonstrated 

significant effects of soil on the distribution of L. tridentata (Ignace 

and Huxman 2009; McAuliffe 1994). Moreover, limestone-containing 

soils are common throughout the Chihuahuan Desert (Brown 1994; 

Wentworth 1985) and many of the diploid sites in this study appear to 

occur upon underlying limestone-containing sedimentary series. This 

includes the unsampled, but probable, boundary between diploids and 

tetraploids (fig. 1) in the Needle’s Eye Canyon along the Gila River 

(Richard and others 2000; www.nationalmap.gov).

If the within-taxon analysis presented here is a reasonable indicator 

of where the Chihuahuan and Sonoran Desert boundary occurs, the 

distributions of diploid and tetraploid Larrea tridentata (along with 

other plants) suggest the current boundary between these deserts 

occurs within the Gila and San Pedro River valleys.

The boundary between diploids and tetraploids identified in this 

study may imply that a previously unrecognized zone of contact has 

long persisted in the low elevations of the Gila and San Pedro Rivers. 

No paleovegetation records have been reported for low elevations in 

this area, though some historical records and photographs indicate 

limited pockets of desert vegetation occurred among grasslands in 

these areas within the last 200 years (Hastings and Turner 1965; 

Humphrey 1987). The steep alluvial slopes near the cytotype boundary in the Needle’s Eye Canyon of the Gila River (Mescal Mts., Ryan 

1985) are very similar to northern extralimital populations of Larrea 

tridentata that grow on the alluvial slopes of the Rio Grande valley 

south of Albuquerque, New Mexico (Wislizenus 1848). These climatic 

conditions have likely been present for thousands of years raising the 

possibility that L. tridentata has persisted for an extended period in 

this ancient drainage. Future efforts to obtain paleovegetation data 

from pack rat (Neotoma) middens at low elevations in the eastern 

Catalina, northwestern Galiuro, and southern Mescal Mountains 

would be particularly informative for determining ancient botanical 

communities of the area, which may help elucidate the importance 

and persistence of the Gila and San Pedro Rivers as corridors between 

the Chihuahuan and Sonoran Deserts.

References

Ackerman, E.A. 1941. The Köppen classification of climates in North America. 

Geographical Review 31: 105-111.

Bahre, C.J. 1995. Human Impacts on the Grasslands of Southeastern Arizona. 

In McClaran, M. P., and T.R. Van Devender, Eds., The Desert Grassland. 

Tucson: The University of Arizona Press, pp. 230-264.

Barbour, M.G. 1969. Patterns of genetic similarity between Larrea divaricata

of North and South America. American Midland Naturalist 81: 54-67.

Brown, D.E., Ed. 1994. Biotic Communities: Southwestern United States and 

Northwestern Mexico. Salt Lake City: University of Utah Press.

Benson, L. 1969. The Cacti of Arizona. 3rd Ed. Tucson: University of Arizona 

Press.

Benson, L., and R.A Darrow. 1981. Trees and Shrubs of the Southwest Deserts. 

Tucson: University of Arizona Press.

Betancourt, J.L., T.R. Van Devender, P.S. Martin, Eds. 1990. Packrat Middens: 

The Last 40,000 Years of Biotic Change. Tucson: University of Arizona Press.

Burgess, T.L. 1995. Desert Grassland, Mixed Shrub Savanna, Shrub Steppe, 

or Semidesert Scrub? In McClaran, M. P., and T.R. Van Devender, Eds., 

The Desert Grassland. Tucson: The University of Arizona Press, pp. 31-67.

Danforth, B.N. 1994. Taxonomic review of Calliopsis subgenus Hypomacrotera

(Hymenoptera; Andrenidae), with special emphasis on the distributions and 

host plant associations. Pan-Pacific Entomologist 70: 283-300.

Felger, R.S. 2000. Flora of the Gran Desierto and Rio Colorado of Northwestern Mexico. Tucson: University of Arizona Press.

Gehlbach, F.R. 1967. Vegetation of the Guadalupe escarpment, New MexicoTexas. Ecology 48: 393-419.

Gleason, H.A. 1926. The individualistic concept of the plant association. 

Bulletin of the Torrey Botanical Club 53: 7-26.

Gleason, H.A. 1939. The individualistic concept of the plant association. 

American Midland Naturalist 21: 92-110.

Glor, R.E., and D. Warren. 2011. Testing ecological explanations for biogeographic boundaries. Evolution 65: 673-683.

Hastings, J.R., and R.M. Turner. 1965. The changing mile. An ecological study 

of vegetation change with time in the lower mile of an arid and semiarid 

region. Tucson: University of Arizona Press.

Henrickson, J.R. and R.M. Straw. 1976. A Gazeteer of the Chihuahuan Desert 

Flora. Los Angeles: California State University.

Hijmans, R.J., M. Jacobs, J.B. Bamberg, D.M. Spooner. 2003. Frost tolerance 

in wild potato species: Assessing the predictivity of taxonomic, geographic, 

and ecological factors. Euphytica 130, 47-59.

Hijmans, R.J., S.E. Cameron, J.L. Parra, P.G. Jones, A. Jarvis. 2005. Very high 

resolution interpolated climate surfaces for global land areas. International 

Journal of Climatology 25: 1965-1978.

Hunter, K.L., J.L. Betancourt, B.R. Riddle, T.R. Van Devender, K.L. Cole, 

and W.G. Spaulding. 2001. Ploidy race distributions since the Last Glacial 

Maximum in the North American desert shrub, Larrea tridentata. Global 

Ecology & Biogeography 10: 521-533.

Hunziker, J.H., R.A. Palacios, L. Poggio, C.A. Naranjo, and T.W. Yang. 

1977. Geographic distribution, morphology, hybridization, cytogenetics, 

and evolution. In Creosote Bush: Biology and Chemistry of Larrea in 

New World Deserts, US/IBP Synthesis Series 6. Stroudsburg: Dowden, 

Hutchison & Ross, Inc.

Humphrey, R.R. 1987. 90 Years and 535 Miles: Vegetation Changes Along the 

Mexican Border. Albuquerque: University of New Mexico Press.

Ignace, D.D., Huxman, T.E. 2009. Limitations to photosynthetic function across 

season in Larrea tridentata (creosotebush) growing on contrasting soil 

surfaces in the Sonoran Desert. Journal of Arid Environments 73: 626-633.

Kozak, K.H., J.J. Wiens. 2006. Does niche conservatism promote speciation? 

A case study in North American salamanders. Evolution 60, 2604-2621.

Laport, R.G., R.L. Minckley, J. Ramsey. 2012. Phylogeny and cytogeography 

of the North American Creosote Bush (Larrea tridentata; Zygophyllaceae). 

Systematic Botany 37: 153-164.

Lewis, W.H. (Ed.) 1980. Polyploidy, Biological Relevance. New York: 

Plenum Press.

Mabry, T.J., J.H. Hunziker, D.R. Difeo Jr. 1977. Creosote Bush: Biology and 

Chemistry of Larrea in New World Deserts, US/IBP Synthesis Series vol. 

6. Stroudsburg: Dowden, Hutchinson & Ross, Inc.

MacEwen, R., R.S. Mann, A.M. Castillo, D.P. Guertin. 2005. Defining boundaries across borders: A case study extending a major land resource area into 

Mexico. RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station.

McAuliffe, J.R. 1994. Landscape evolution, soil formation, and ecological 

patterns and processes in Sonoran Desert bajadas. Ecological Monographs

64: 112-148.

McCune, B., and J.B. Grace. 2002. Analysis of Ecological Communities. Glen 

Eden Beach: MjM Software Design.

McLaughlin, S.P. 1986. Floristic analysis of the southwestern United States. 

Great Basin Naturalist 46: 46-65.

Morafka, D.J. 1977. Is there a Chihuahuan Desert? A quantitative evaluation 

through a herpetofaunal perspective. In Wauer, R.H. and D.H. Riskind, 

Eds., Transactions of the Symposium on the Biological Resources of the 

Chihuahuan Desert Region, U.S. and Mexico. Alpine, Texas. USDI National 

Park Service Transactions and Proceedings Series 3, pp. 437-454.

Oberhauser, K., A.T. Peterson. 2003. Modeling current and future potential 

wintering distributions of eastern North American monarch butterflies. Proceedings of the National Academy of Sciencesof the USA 100, 14063-14068.
224 USDA Forest Service Proceedings RMRS-P-67. 2013

Laport and Minckley Cytogeography of Larrea tridentata at the Chihuahuan-Sonoran Desert Ecotone

Pearson, R.G., Raxworthy, C.J., Nakamura, M., Peterson, A.T. 2007. Predicting 

species distributions from small numbers of occurrence records: a test case 

using cryptic geckos in Madagascar. Journal of Biogeography 34: 102-117.

Phillips, S.J., R.P. Anderson, R.E. Schapire. 2006. Maximum entropy modeling 

of species geographic distributions. Ecological Modeling 190: 231-259.

Richard, S.M., S.J. Reynolds, J.E. Spencer, and P.A. Pearthree, comps., 2000, 

Geologic Map of Arizona: Arizona Geological Survey Map M-35, 1 sheet, 

scale 1:1,000,000.

Riddle, B.R. 1998. The historical assembly of continental biotas: Late Quaternary range-shifting, areas of endemism, and biogeographic structure in 

the North American mammal fauna. Ecography 21: 437-446.

Riddle, B.R., D.J. Hafner, L.F. Alexander, and J.F. Jaeger. 2000. Cryptic vicariance in the historical assembly of a Baja California Peninsular Desert 

biota. Proceedings of the National Academy of Sciences of the USA 97: 

14438-14443.

Rozen, Jr., J.G. 1992. Systematics and host relationships of the cuckoo bee 

genus Oreopasites (Hymenoptera: Anthopheridae: Nomadinae). American 

Museum Novitates 3046: 1-56.

Roura-Pascual, N., A.V. Suarez, C. Gomez, P. Pons, Y. Touyama, A.L. Wild, 

A.T. Peterson. 2004. Geographical potential of Argentine ants (Linepitherma humile Mayr) in the face of global climate change. Proceedings of 

the Royal Society of London Series B-Biological Sciences 271, 2527-2534.

Russell, R.J. 1931. Dry Climates of the United States. Berkeley: University 

of California Press Publications in Geography 5:1-41 + map.

Ryan, G.S. 1985. Mineral investigation of the Needle’s Eye Wilderness Study 

Area, Gila County, Arizona. U.S. Bureau of Mines Mineral Land Assessment MLA 9-85. Denver: U.S. Department of the Interior Bureau of Mines.

Schmidt, Jr., R.H., 1979. A climatic delineation of the ‘real’ Chihuahuan 

Desert. Journal of Arid Environments 2: 243-250.

Shreve, F. 1942. The desert vegetation of North America. The Botanical 

Review 8: 195-246.

Shreve, F. 1951. Vegetation and Flora of the Sonoran Desert. Volume I. Vegetation. Carnegie Institution of Washington Publications 591.

Soule, O.H., and C.H. Lowe. 1970. Characteristics of tissue fluids in the Sahuaro Giant Cactus (Cereus giganteus). Annals of the Missouri Botanical 

Garden 57: 265-351.

Stohlgren, T.J., C.S. Jarnevich, W.E. Esaias, J.T. Morisette. 2011. Bounding 

species distribution models. Current Zoology 57: 642-647.

Thornthwaite, C.W. 1948. An approach toward a rational classification of 

climate. Geographical Review 38: 55-94.

Turner, R.M., J.E. Bowers, T.L. Burgess. 1995. Sonoran Desert Plants: An 

Ecological Atlas. Tucson: University of Arizona Press.

Warshall, P. 1994. The Madrean Sky Island Archipelago: A planetary overview. Gen. Tech. Rep. RM-GTR264. Fort Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station.

Warren, D.L., R.E. Glor, M. Turelli. 2008. Environmental niche equivalency 

versus conservatism: Quantitative approaches to niche evolution. Evolution 62: 2868-2883.

Weaver, J.W. and F.E. Clements. 1938. Plant Ecology. New York: McGrawHill Book Co.

Wentworth, T.R. 1985. Vegetation on limestone in the Huachuca Mountains, 

Arizona. The Southwestern Naturalist 30: 385-395.

Wislizenus, A. 1848. Memoir of a tour to northern Mexico, connected with Col. 

Doniphan’s Expedition, in 1846 and 1847. Washington: Tippin & Streeper.

Yang, T.W. 1967. Chromosome numbers in populations of creosote bush (Larrea divaricata) in the Chihuahuan and Sonoran subdivisions of the North 

American Desert. Journal of the Arizona Academy of Science 4: 183-184.

Yang, T.W. 1968. A new chromosome race of Larrea divaricata in Arizona. 

Western Reserve Academy Natural History Museum Special Publication 

2: 1-4. Hudson, OH: Western Reserve Academy.

Yang, T.W. 1970. Major chromosome races of Larrea divaricata in North 

America. Journal of the Arizona Academy of Science 6: 41-45.

Yang, T.W., and C.H. Lowe. 1968. Chromosome variation in ecotypes of 

Larrea divaricata in the North American Desert. Madroño. 19: 161-164.

Yang, T.W., J.H. Hunziker, L. Poggio, and C.A. Naranjo. 1977. Hybridization 

between South American jarilla and North American diploid creosote bush 

(Larrea, Zygophyllaceae). Plant Systematics and Evolution 126: 331-346.

Zimmerman, R.C. 1969. Plant ecology of an arid basin, Tres Alamos-Redington 

area southeastern Arizona. U.S. Geological Survey Professional Paper, 

485-D.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 225

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Chihuahuan Desert Flora of La Calera, Municipio de Agua 

Prieta, Sonora, Mexico

Ana Lilia Reina-Guerrero and Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Abstract—A total of 555 plant collections were made on 20 trips in 2002-2008 to La Calera area in the Sierra 

Anibácachi, Municipio de Agua Prieta, 11.3 km south of the Arizona border (31°13’59”N 109°37’53”W, 

elevation range from 1220 m to 1539 m) in northeastern Sonora. Chihuahuan desertscrub on limestone 

substrates is dominated by creosotebush (Larrrea divaricata), Chihuahuan whitethorn (Acacia neovernicosa), 

mariola (Parthenium incanum), and tarbush (Flourensia cernua). Riparian vegetation along a rocky bedrock/

gravel wash includes desert willow (Chilopsis linearis), netleaf hackberry (Celtis reticulata), desert hackberry 

(C. pallida), woolly buckthorn (Sideroxylon lanuginosum), soaptree yucca (Yucca elata), Arizona walnut 

(Juglans major), and Coahuila juniper (Juniperus coahuilensis). The flora is diverse with 350 taxa in 60 

families and 222 genera in 25 km2 (2.5 km2 extensively inventoried). Only 5.1% of the taxa are non-native. 

The most numerous plants are in the Poaceae (57 taxa), Asteraceae (53), Fabaceae (26), Malvaceae (22), 

and Euphorbiaceae (21) in the genera Euphorbia (11), Abutilon and Bouteloua (7 each), Eragrostis and 

Muhlenbergia (6 each), and Dalea, Ipomoea, and Sphaeralcea (5 each). Twenty-six taxa are probably the 

first records for Sonora. Another 21 taxa are noteworthy state records (rare or significant range extensions)..

Introduction

The Chihuahuan Desert is the largest of the three creosotebushdominated warm deserts in North America (Shreve, 1942). It primarily 

occurs from New Mexico and Texas southeast through Chihuahua and 

Coahuila to San Luis Potosí (Lowe, 1964; Rzedowski, 1978; Brown, 

1982). Most (90%) is in Mexico on the Mexican Plateau bounded to 

the east by the Sierra Madre Oriental and to the west by the Sierra 

Madre Occidental. Most of this area is above 1000 m elevation and 

regular incursions of frigid Arctic air from the north have shaped the 

evolution of the flora. More than 80% of Chihuahuan desertscrub is 

found on limestone and the gray gravel of this substratum give the 

characteristic view of the landscape (Lowe, 1964; Brown, 1982). 

Throughout the area, there is a mosaic of Chihuahuan desertscrub on 

limestone bedrock and desert grassland in fine-soiled valley bottoms. 

In the Santa Catalina Mountains of Arizona, Shreve (1922) and Whittaker and Niering (1968) concluded that vegetation is more xeric and 

open and that both vegetation and species reach higher elevations on 

limestone. We have noticed that this effect is most prevalent in drier 

Chihuahuan Desert areas, but hardly noticeable in foothills thornscrub 

on limestone in east-central Sonora southward. 

The Chihuahuan Desert reaches its westernmost extent in southeastern Arizona and northeastern Sonora. In this paper, we study 

the flora in Chihuahuan desertscrub on Rancho La Calera in Sierra 

Anibácachi, southwest of Agua Prieta, Sonora.

Study Area

Rancho La Calera, is an isolated hill northeast of Sierra Anibácachi, 

ca. 10 km (by air) southwest of Agua Prieta, 11.3 km (by air) south of 

the United States/Mexico border 31°13’59”N 109°37’53”W (fig. 1). 

The vegetation on limestone substrates at 1220-1539 m elevation is 

Chihuahuan desertscrub. The flora is representative of an area of 25 

km2, with only 2.5 km2 sampled intensively. The Sierra Anibácachi 

is formed by Lower Cretaceous limestone in the Bisbee Group, a 

southern extension of thick and well-exposed marine sediments in 

the Mule Mountains (McKee and others 2005).

The climate of the area is arid, with biseasonal rainfall, hot summers, and cold winters. Weather stations are located in Agua Prieta 

and Naco, Sonora. The mean annual rainfall is 353.5 mm in the 

Municipio of Agua Prieta, with 40-47% during the summer monsoon 

season in July-September. The mean annual temperature is 17.3 °C, 

with a summer maximum of 43 °C in June 1994 and July 1989; a 

winter minimum of –14 °C and wind chill of –24 °C was registered 

in February 2011. Low winter temperatures limit biological activities. 

Another climatic factor is prevailing winds from the south for more 

than 60% of the year, with a strength of 5.40 to 8.49 meters/second 

recorded in the international airport in Douglas, Arizona (Reza, 2011; 

CONAGUA, 2010; Universidad Autónoma de Tamaulipas, 2003).

The Sierra Anibácachi is in the Río Agua Prieta drainage basin, 

which is a tributary of the greater Río Yaqui. It begins as White Water Draw in Arizona before crossing the border into Sonora in Agua 

Prieta. The watershed area in Arizona is 2,650 km2. From the border 

south to its junction with Río Fronteras, the Río Agua Prieta basin has 

an area of 1,444 km2. Downstream this drainage coalesces into the 

Río Bavispe upstream of Lázaro Cárdenas (La Angostura) reservoir 

(CONAGUA 2010).
226 USDA Forest Service Proceedings RMRS-P-67. 2013

Reina-Guerrero and Van Devender Chihuahuan Desert Flora of La Calera, Municipio de Agua Prieta, Sonora, Mexico

Methods

Plant Collections 

Between September 2002 and May 2008, we made 20 trips to the 

study area, both during the summer rainy season in August-October, 

and the dry spring in March-June. We surveyed a broad range of habitats, including rocky slopes, peaks, cliffs, alluvial bajadas, arroyos, 

and cattle tanks. Most of our efforts were concentrated on the slopes 

and arroyo close to the La Calera ranch headquarters, and the quarry 

(calera) owned by Grupo México. Whenever possible we recorded 

plant common names from local cowboys. 

We collected 590 specimens under the SEMARNAT permit to J. 

Jesús Sánchez-E., Curator of the Universidad de Sonora Herbarium 

(USON). Specimens were deposited into herbaria at the University 

of Arizona (ARIZ), USON, Arizona State University (ASU), and 

the Universidad Nacional Autónoma de México (MEXU). Although 

there were some previous collections from the Municipio of Agua 

Prieta, the Calera area had not been collected before. Specimens were 

identified in the University of Arizona herbarium or by specialists 

in other herbaria using duplicates or images. Nomenclature mostly 

follows the list in Van Devender and others (2010), which is available 

and updated in the Madrean Archipelago Biodiversity Assessment 

(MABA)/Southwest Environmental Information Network (SEINet) 

online database (Madrean.org). Plant collections, observations, and 

photographs are available in this database. A complete list of the La 

Calera flora will be available as a Research Species list in the database. 

Results

Vegetation 

The vegetation in the La Calera study area is Chihuahuan desertscrub, 

which occurs on limestone bedrock, limestone-derived alluvial 

bajadas, and the lower fine-soiled flats. The limestone bedrock on a 

northeast slope is dominated by Fouquieria splendens, in association 

with the shrubs Acacia neovernicosa, Calliandra eriophylla, Senna 

wislizenii, and Vauquelinia californica. Additional shrubs include A. 

millefolia, Aloysia wrightii, Brickellia baccharidea, Dalea formosa,

Eysenhardtia orthocarpa, Fraxinus gooddingii, Mimosa biuncifera, 

Mortonia scabrella, and Tecoma stans. On upper parts of ridges, 

Quercus pungens and Bernardia myricifolia are locally dominant.

Succulents on bedrock slopes, including Agave palmeri, Dasylirion 

wheeleri, Opuntia phaeacantha, and Yucca baccata, are important 

in the community, as are the perennial herbs Anemone tuberosa, 

Astrolepis cochisensis, Physaria fendleri, P. purpurea, and Salvia 

parryi. Perennial bunch grasses, notably Aristida purpurea var. nealleyi, Bouteloua curtipendula, Muhlenbergia arizonica, and Panicum 

hallii, are common. 

On the alluvial slopes, the dominants are Acacia neovernicosa, 

Larrea divaricata, and Parthenium incanum, with Krameria erecta 

and Tiquilia canescens. These plants are also common in flatter areas 

along with Flourensia cernua and Prosopis velutina. On benches above 

the arroyo, we found Anisacanthus thurberi, Condalia warnockii, 

Forestiera angustifolia, F. phillyreoides, Koeberlinia spinosa, and 

Ziziphus obtusifolia. Along a gravelly/rocky wash incised through the 

alluvium to the limestone bedrock, Baccharis salicifolia, Gutierrezia 

microcephala, and Hymenoclea monogyra, are common in riparian 

vegetation. In the wash, B. sarothroides, Celtisreticulata, C. pallida, 

Chilopsis linearis, Sideroxylon lanuginosum, and Yucca elata are 

occasional, and Juglans major and Juniperus coahuilensis are rare.

Flora Composition

We documented 350 taxa in 60 families and 222 genera in Chihuahuan desertscrub at La Calera. The families with more species 

are Poaceae (57 taxa, 16.3%), Asteraceae (53 taxa, 15.1%), Fabaceae 

(26 taxa, 7.4%), Malvaceae (22 taxa, 6.3%), Euphorbiaceae (21 taxa, 

6.0%), Solanaceae (15 taxa, 4.3%), and Cactaceae (12 taxa, 3.4%). 

The genera with the most species are Euphorbia (11 taxa), Abutilon 

and Bouteloua (7 taxa each), Eragrostis and Muhlenbergia (6 taxa 

each), Dalea, Ipomoea, and Sphaeralcea (5 taxa each), Acacia, Boerhavia, Brickellia, Lepidium, Opuntia, and Sporobolus (4 taxa each). 

Only 18 species are non-native (5.1%), including Cynodon dactylon, 

Eragrostis lehmanniana, Pennisetum ciliare, Sorghum halepense, and 

Tamarix chinensis. Although those species sometimes are invasive in 

other areas, they do not represent an ecological threat at La Calera. 

The life forms of La Calera are herbs (70.0%, including grasses), 

woody plants (24.9%), and succulents (5.1%). The dicot herbs (188 

taxa) are perennial (97 taxa) and annual (91 taxa). The grasses and 

sedges (57 taxa) are perennial (38 taxa) and annual (19 taxa). 

Noteworthy Records—About 26 species in the La Calera area 

are probably the first records for Sonora, with no previous records 

in the MABA/SEINet database. These include Abutilon coahuilae, 

Acourtia nana, Amaranthus blitoides, Asclepias macrotis, Bothriochloa laguroides var. laguroides, Chamaesaracha sordida, Cryptantha albida, Cylindropuntia leptocaulis X O. spinosior, Cyphomeris 

gypsophiloides, Dyssodia papposa, Fumaria parviflora, Gaillardia 

pulchella, Heliotropium greggii, H. pringlei, Hybanthus verticillatus, 

Ibervillea tenuisecta, Nolina texana, Opuntia macrocentra (most 

other Sonoran records are misidentified O. gosseliniana), Physaria 

fendleri, Physalis lagascae, Quercus pungens, Ruellia parryi, Simsia 

lagascaeformis, Sphaeralcea polychroma, Stipa eminens, and Vicia 

ludoviciana. Most of these records represent southern extensions 

from the United States.

An additional 28 taxa are considered rare in Sonora: Abutilon parvulum, A. wrightii, Asclepias nyctaginifolia, Bernardia myricifolia, 

Clerodendron coulteri, Conoclinium dissectum, Cryptantha pusilla, 

Erioneuron avenaceum, Euphorbia exstipulata, E. stictospora, Polygala scoparioides, and Vauquelinia californica ssp. pauciflora. Other 

species are rarely collected in Sonora: Dalea formosa, Flourensia 

cernua, Mortonia scabrella, Phyllanthus polygonoides, and Thamnosma texana. The annual Streptanthus carinatus had canary yellow 

sepals in limestone cliffs and white sepals in sand in the arroyo. The 

yellow form was formerly S. arizonicus S. Watson var. luteus KearFigure 1—Map of the Rancho Calera (CA) area.
USDA Forest Service Proceedings RMRS-P-67. 2013 227

Chihuahuan Desert Flora of La Calera, Municipio de Agua Prieta, Sonora, Mexico Reina-Guerrero and Van Devender

ney & Peebles. It was synonomized into S. carinatus ssp. arizonicus 

by Kruckeberg and others (1982), but warrants reevaluation. Again 

most of these species are southern range extensions, but Abutilon 

abutiloides, Physalis lagascea, and Ipomoea aristolochiifolia (Austin, 

2006) are northern extensions of tropical species not recorded in the 

United States.

Discussion

After 7 years of plant collections, the flora of the La Calera area 

is well represented. This is a limestone flora rich in plant species 

and the first Chihuahuan desertscrub flora studied in Sonora. The 

only North American desert characterized by trees is the subtropical 

Sonoran Desert, and not the more temperate Great Basin, Mohave 

and Chihuahuan Deserts. Only 1.7% of the La Calera flora are trees. 

Of these, Celtis reticulata, Chilopsis linearis, Juglans major, and 

Juniperus coahuilensis are riparian along the wash. Prosopis glandulosa and P. velutina are trees in other areas, but shrubs in La Calera. 

Vauquelinia californica is a tall shrub that is locally dominant on 

steep slopes. This species is endemic to central Arizona and adjacent 

Sonora, and does not occur in the main Chihuahuan Desert in New 

Mexico, Texas, Chihuahua, or Coahuila. The subspecies pauciflora

is found on limestone in southeastern most Arizona and few places 

in Sonora. Quercus pungens is typically a shrub on limestone, but 

becomes a tree of 5-6 m height in western Texas.

Of the 85 species of native annuals, 62 (72.9%) are summer annuals, 20 (23.5%) spring annuals, and three active in either season. 

The abundance of summer annuals (72.9%) is interesting because to 

the west in the Sonoran Desert, and more so in the Mohave Desert, 

winter annuals are dominant. 

The flora of the San Pedro Riparian National Conservation Area 

(SPRNCA) in Cochise County, Arizona, supports a flora of 625 taxa 

(56% greater than La Calera) in an area of 19,291 ha (8 times larger 

than La Calera; Makings, 2006). The flora of the Huachuca Mountains 

in Cochise County supports a flora of 917 taxa (61% greater than La 

Calera) in an area of 31,600 ha (12 times larger than La Calera; Bowers and McLaughlin, 1996). According to Bowers and McLaughlin 

(1996), species diversity is best correlated with elevational range and 

habitat diversity, since both temperature and precipitation vary with 

elevation. The La Calera flora is interesting because habitat diversity 

and elevational range are minimal but the flora is diverse. Wentworth 

(1982) documented the distributions of plant species on limestone 

bedrock and calcareous soils at 1400 to 1900 m elevations in the Mule 

Mountain. Of 232 species in these habitats, 147 are shared with La 

Calera. The 85 species not in la Calera were in the mesic limestone 

habitat in oak woodland zone, which is 400 m above the La Calera 

study area; i.e., Bouvardia ternifolia, Cercocarpus montanus, Garrya wrightii, Quercus arizonica, Pinus discolor, etc. It is especially 

interesting that La Calera had 203 taxa not recorded in the Mule 

Mountains, only 39 km west-northwest of La Calera.

The distributions of the plants in the La Calera flora are predominately southwestern United States and northwestern Mexican, with 

biotic affinities to the Great Plains grasslands and the Chihuahuan 

Desert, which form a mosaic in most areas. Fouquieria splendens, 

Larrea divaricata, Tecoma stans, Opuntia phaeacantha, and Yucca 

elata are widespread. Bernardia myricifolia, Acacia neovernicosa, 

Flourensia cernua, Mortonia scabrella, Nolina texana, Panicum 

hallii, Parthenium incanum, Quercus pungens, Salvia parryi, Senna 

wislizeni, and Tiquilia canescens are Chihuahuan species mostly 

found on limestone or calcareous soils. Opuntia chlorotica primarily 

occurs in Arizona and adjacent southern California, southwestern New 

Mexico, and northern Sonora, but not in the Chihuahuan Desert to 

the east.

Three species in La Calera are considered “in risk” or “candidate” 

species according to the Official Mexican Regulation NOM-059-

ECOL-2002 (D.O.F. 2002), which establishes the degree of protection for species: Amoreuxia palmatifida and Peniocereus greggii var. 

greggii are listed as Pr [special protection] and Juglans major is listed 

as A [threatened]. None of the plants from La Calera is listed either 

in Arizona or the United States.

We conclude that the La Calera is a very interesting area with a rich, 

regionally unique flora in the westernmost Chihuahuan Desert. With 

records of about 40 species new to or rare in Sonora in the northwestern 

borderlands of the country, the flora is very important for Mexico. 

The Grupo México quarry operation has severely disturbed slopes 

on the La Calera ridge just northeast of the study area. Protection of 

a representative area of this habitat and vegetation is warranted. . 

Acknowledgments

We want to thank the 35 specialists who helped with identifications, 

especially Charlotte and John Reeder, Paul Fryxell, Billie Turner, 

Alfonso Delgado, and Victor W. Steinmann. And thanks to the many 

botanists who joined us in the field, especially Greta Anderson, John 

Anderson, Dixie Z. Damrel, Shannon Doan, George Ferguson, Zhiwei 

Liu, and John Wiens. We thank Stephen P. McLaughlin and Richard 

Worthington for their careful review.

References

Austin, Daniel F. 2006. Noteworthy distributions and additions in Southwestern 

Convolvulaceae. Canotia, 2: 79-106

Bowers, Janice E.; and McLaughlin, Steven P. 1996. Flora of the Huachuca 

Mountains, a botanically rich and historically significant sky island in 

Cochise County, Arizona. Journal of the Arizona-Nevada Academy of 

Science, 29: 66-107. 

Brown, David E. 1982. Chihuahuan Desertscrub. In Brown, David E. ed. Biotic Communities of the American Southwest – United States and Mexico. 

Desert Plants. Special Issue. 4 (1-4): 169-179.

CONAGUA. 2010. Determinación de la disponibilidad de agua en el acuífero 

2629, río Agua Prieta, estado de Sonora. 30 p.

D.O.F. 2002. NOM-059-ECOL-2001. Protección ambiental - especies nativas 

de México de flora y fauna silvestres - Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio - Lista de especies en riesgo. 

Diario Oficial de la Federación, 6 March 2002.

Kearney, Thomas H.; and Peebles, Robert H. 1964. Arizona Flora. Berkeley: 

University of California Press, 1085 p.

Kruckeberg, A.R.; Rodman, J.E.; and Worthington, Richard D. 1982. Natural 

hybridization between Streptanthus arizonicus and S. carinatus (Cruciferae). 

Systematic Botany, 7: 291-299. 

Lowe, Charles H. 1964. Arizona’s Natural Environment. Landscapes and 

Habitats. Tucson: The University of Arizona Press.136 p.

Makings, Elizabeth. 2006. Flora of the San Pedro Riparian National Conservation Area. Deserts Plants. Special Issue. 22. 104 p.

McKee, J.W.; McKee, M.B.; and Anderson, T.H. 2005. Mesozoic basin formation, mass-gravity sedimentation, and inversion in northeastern Sonora and 

southeastern Arizona. In Anderson, T.H.; Nourse, J.A.; McKee, J.W.; and 

Steiner, M.B. eds. The Mohave-Sonora Megashear Hypothesis: Development, Assessment, and Alternatives. The Geological Society of America. 

Special Paper 393. 481-508.

Reza, Francisco; and López, Gerardo. Sonora se congela: -16 oC. El Imparcial. Diario Independiente de Sonora: Hermosillo, México. Febrero 4, 

2011. p. A1, col. 1.

Rzedowski, Jerzy. 1978. Vegetación de México. México: Editorial Limusa. 

432 p.
228 USDA Forest Service Proceedings RMRS-P-67. 2013

Reina-Guerrero and Van Devender Chihuahuan Desert Flora of La Calera, Municipio de Agua Prieta, Sonora, Mexico

Shreve, Forrest. 1922. Conditions indirectly affecting vertical distribution on 

desert mountains. Journal of Ecology, 3: 269-274

Shreve, Forrest. 1942. The desert vegetation of North America. Botanical 

Review, 8: 195-246

Universidad Autónoma de Tamaulipas. 2003. Manifestación de impacto ambiental modalidad particular. Comisión Federal de Electricidad. Proyecto 

ciclo combinado Agua Prieta II. 151 p.

Van Devender, Thomas. R.; Felger, Richard F.; Fishbein, Mark; MolinaFreaner, Francisco; Sánchez-Escalante, J. Jesús; and Reina-Guerrero, Ana 

L. 2010. Biodiversidad de las plantas vasculares. Pp. 229-262 in MolinaFreaner, Francisco; and Van Devender, Thomas R. eds., Diversidad Biológica de 

Sonora, Universidad Nacional Autónoma de México, Hermosillo. http://

www.skyislandalliance.org/Sonoran%20vascular%20plants-VanDevender%20et%20al.2010.pdf

Wentworth, Thomas R. 1982. Vegetation and flora of the Mule Mountains, 

Cochise County, Arizona. Journal of the Arizona-Nevada Academy of 

Science, 17: 29-44.

Whittaker, Robert H.; and Niering, William A. 1968. Vegetation of the Santa 

Catalina Mountains, Arizona. IV. Limestone and acid soils. Journal of 

Ecology, 56: 523-544.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 229

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Flora of Chihuahuan Desertscrub on Limestone in 

Northeastern Sonora, Mexico 

Thomas R. Van Devender and Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona

J. Jesús Sánchez-Escalante

Departamento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Hermosillo, Sonora, 

Mexico

Abstract—Transects were done in desertscrub on limestone to characterize the flora of the westernmost 

Chihuahuan Desert. Most of the sites (15) were in the Municipios of Agua Prieta and Naco in northeastern 

Sonora, with single sites near Ascensión, northwestern Chihuahua and east of Douglas in southeastern 

Arizona. A total of 236 taxa were recorded on transects. Dicot perennial herbs (66 species) were the most 

numerous life forms, followed by annual herbs (36 species), subshrubs (36 species), and woody shrubs (33 

species). The most diverse limestone floras were on Rancho La Morita in the Municipio of Naco (87 to 119 

taxa/transect), Rancho La Calera (88 taxa), and Cerro El Caloso and Cerrito Los Janitos (84 taxa each) in the 

Municipio of Agua Prieta. Chihuahuan desertscrub at its western limits is a mixture of typical Chihuahuan 

species that reach Texas and Coahuila with western and tropical species in associations that are regionally 

distinct from other parts of the Chihuahuan Desert.

Introduction 

	 The Chihuahuan Desert is the largest of the three creosotebushdominated warm desertsin NorthAmerica (Shreve, 1942). It primarily occurs from San Luis Potosí northwest through Chihuahua and

Coahuila to Texas and New Mexico (Lowe, 1964; Rzedowski, 1978;

Brown, 1982). Most (90%) is in Mexico on the Mexican Plateau

bounded to the east by the Sierra Madre Oriental and to the west

by the Sierra Madre Occidental. Most of this area is above 1000 m

elevation and regular incursions of frigid Arctic air from the north

have shaped the evolution of the flora. More than 80% of Chihuahuan desertscrub is found on limestone, and the gray gravel of this

substratumgive the characteristic viewofthe landscape (Lowe, 1964;

Brown, 1982). Throughout the area, there is a mosaic of Chihuahuan

desertscrub on limestone bedrock and desert grassland in fine-soiled

valley bottoms. In the Santa Catalina Mountains of Arizona, Shreve

(1922) and Whittaker and Niering (1968) concluded that vegetation

is more xeric and open and that both vegetation and species reach

higher elevations on limestone than on other substrates. We have

noticed that this effect is most prevalent in northern areas with drier

climates, but hardly noticeable in foothillsthornscrub on limestone in

east-central Sonora and tropical deciduousforest in southern Sonora.

	 During the spring of 2007, we attempted to relocate the endangered

Cochise pincushion cactus (Coryphantha robbinsorum) reported

from Sonora by Lopresti (1984). Although C. robbinsorum was not

found, local floras on calcareous substrates were surveyed in northeastern Sonora and adjacent Arizona and Chihuahua (Van Devender

and Reina-G., 2007). Here we use the results of these inventories to

characterize Chihuahuan desertscrub at its westernmost extension.

Methods 

	 We visited a C. robbinsorum population on Permian limestone on

the Magoffin Ranch in Cochise County, Arizona, to refine our search

image for the plant and its habitat. Potential areasfor C. robbinsorum 

were identified in Sonora based on the presence of Paleozoic or lower

Cretaceous limestone substrates on Instituto Nacional de Estadística

y Geografía (INEGI) geological maps, the proximity to the Arizona

C. robbinsorum populations, and habitat descriptions in Lopresti

(1984). Seven areas in Sonora in the Municipios de Agua Prieta and

Naco and one in Chihuahua were selected for plant inventories(table 1,

fig. 1).

	 The climate of the area is arid, with biseasonal rainfall, hot summers, and cold winters. The weather stations nearest all transects are

located in Agua Prieta and Naco, Sonora. The mean annual rainfall

is 353.5 mm in the Municipio of Agua Prieta, with 40-47% during

the summer monsoon season in July-September. The mean annual

temperature is 17.3 °C, with a summer maximum of 43 °C in June

1994 and July 1989; a winter minimum of –14 °C and wind chill of

–24 °C was registered in February 2011. Low winter temperatures

limit biological activities. Another climatic factor is the prevailing

wind from the south for more than 60% of the year, with a strength

of 5.4 to 8.5 meters/second recorded in the international airport in

Douglas,Arizona (Reza et al., 2011;CONAGUA, 2010; Universidad

Autónoma de Tamaulipas, 2003).
230 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico Table 1—Localities for limestone transects in Arizona, Chihuahua, and Sonora. Identification Location Elevation Description Arizona MG1: Cochise County: limestone hills on Magoffin Ranch 31°22’13”N 109°08’18”W, 1284 m elevation to 31°22’00”N 109°24”W, 1311 m

(Coryphantha robbinsorum population). Open, grassy Chihuahuan desertscrub-desert grassland mix on very gentle slope with gray limestone bedrock on upper portions of the hill.

Chihuahua: Municipio de AscensiónCH1

Cerro de Cal (La Biznaga on geology map), west of Ascensión; two areas combined

31°07’04”N 108°05’23”W, 1380 m elevation and 31°06’35”N 108°05’04”W, 1322 m elevation

Open Chihuahuan desertscrub on dark gray limestone with white and orange secondary veins.

Sonora: Municipio de Agua PrietaAL1

Ridge above El Álamo, ca. 21 km south of the Cerro La Minita transect on Rancho Puerta Blanca, north of the Sierra Pitáicachi, ca. 2 km west of Cerro Agua Zarca, 7 km northwest of Rancho El Capadero, 9.7 km (by air) southeast of La Cieneguita (= Oquita Montenegro) in Cajón Bonito

1239 to 1289 m elevation Chihuahuan desertscrub on west-facing, gray, fossiliferous limestone; good grass, not heavily grazed.

AP1 ca. 15 km east of Agua Prieta on north side of MEX 2; 31°19’03”N 109°22’36”W 1318 m elevation

Chihuahuan desertscrub on gentle hill, south and northeast slopes, highly modified Cretaceous limestone with bands of whitish quartz; southern slope very grassy, northeast slope dominated by Flourensia cernua and Acacia neovernicosa.

CA1

Rancho La Calera, ca. 10 km (by air) southwest of Agua Prieta, outlier ridge of Sierra Anibácachi

31°14’07”N 109°37’40”W, 1289 m elevation

Chihuahuan desertscrub on whitish gray limestone upper northeast-facing slope, alluvium below.

CA2

Rancho La Calera, ca. 10 km (by air) southwest of Agua Prieta, outlier ridge of Sierra Anibácachi

31°13’59”N 109°37’37”W, 1297 m elevation

Chihuahuan desertscrub on whitish gray limestone upper northwest-facing slope, alluvium below

CL1 Cerro Caloso (= Calichoso), ca. 8 km (by air) east of Cabullona

lower end: 31°08’23”N 109°28’15”W, 1215 m elevation, upper end: 31°08’14”N 109°27’59”W, 1272 m

Chihuahuan desertscrub on east-northeast facing limestone ridge with most layers nearly vertical, but with lots of bedrock exposed, dark gray (to purple) with lots of secondary white calcite veins.

JN1

Cerrito Los Janitos, east of the north end of the Sierra Anibácachi, ca. 1.5 km (by air) west-southwest of Agua Prieta

lower end: 31°15’57”N 109°40’15”W, 1334 m elevation, upper end: 31°15’56”N 109°40’27W, 1429 m

Chihuahuan desertscrub on light gray limestone (similar to CA1 and CA2); mostly on east-facing slope.
USDA Forest Service Proceedings RMRS-P-67. 2013 231

Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico Van Devender and others Table 1—Continued Identification Location Elevation Description PB1 Ca. 40 km east of Agua Prieta on Rancho Puerta Blanca, north of Arroyo Guadalupe and MEX 2, just south of Arizona border (below transect PB2) upper end: 31°18’56”N 109°06’30”W, 1294 m elevation Chihuahuan desertscrub on south-southwest slope; lower part limey quartzite, upper gray limestone PB2 Ca. 40 km east of Agua Prieta on Rancho Puerta Blanca, north of Arroyo Guadalupe and MEX 2, just south of Arizona border (above transect PB1) upper end: 31°19’22”N 109°06’37”W, 1386 m elevation Chihuahuan desertscrub on south-southwest slope; gray limestone. PB3 Ca. 37 km east of Agua Prieta on Rancho Puerta Blanca, south of MEX 2, east of Arroyo Guadalupe 31°17’58”N 109°08’40”W, 1277 m elevation Chihuahuan desertscrub on top of gentle north-south limestone conglomerate ridge PB4 Ca. 37 km east of Agua Prieta on Rancho Puerta Blanca, south of MEX 2, east of Arroyo Guadalupe 31°18’04”N 109°07’36”W, 1355 m elevation Chihuahuan desertscrub on gentle west-facing slope, mostly loose limestone weathering from limestone conglomerate ridge. Sonora: Municipio de Naco MO1 Hill west of Arroyo La Bellota, Rancho La Morita, ca. 25 km west of Agua Prieta lower end: 31°08’23”N 109°28’15”W, 1215 m elevation, upper end: 31°08’14”N 109°27’59”W, 1272 m Excellent gray bedrock limestone on gentle slopes (all directions) and large flat area on top; grasses common. MO2 Hill west of Arroyo La Bellota, Rancho La Morita, ca. 25 km west of Agua Prieta lower end: 31°15’14”N 109°49’05”W, 1438 m elevation; upper east end: 31°15’21”N 109°48’50”W, 1486 m; upper north end: 31°15’27”N 109°48’55”W, 1486 m Chihuahuan desertscrub. Transect began on west-facing slope with scattered gray bedrock limestone on alluvium; then to north slope bedrock on more gentle northeast slopes and hill tops. MO3 Cerro La Bruja (north), Rancho La Morita, ca. 23 km west, 5.5 km south (by air) of Agua Prieta lower end: 31°14’39”N 109°48’25”W, 1496 m elevation; upper end: 31°14’42”N 109°48’22”W, 1543 m Chihuahuan desertscrub on excellent gray bedrock limestone on steep south-facing slope. MO4 Cerro La Bruja (north), Rancho La Morita, ca. 23 km west, 5.5 km south (by air) of Agua Prieta (just west of MO3 transect) lower end: 31°14’40”N 109°48’57”W, 1477 m elevation; upper end: 31°14’46”N 109°48’47”W, 1547 m Chihuahuan desertscrub on excellent gray bedrock limestone on steep south-facing, open grassy slope. MO5 Cerro La Bruja, Rancho La Morita, ca. 23 km west, 5.5 km south (by air) of Agua Prieta (across Arroyo La Bruja, southeast of MO3 and MO4 transects) lower northwest end: 31°14’17”N 109°47’38”W, 1509 m elevation; east end: 31°14’17”N 109°47’340”W, 1520 m Chihuahuan desertscrub on excellent gray bedrock limestone on gentle slopes on north-facing slope and east-west ridgetop; grasses common.
232 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico

	 All of the study area except the site in Chihuahua is in the Río

Agua Prieta drainage basin, a tributary of the greater Río Yaqui that

begins asWhiteWaterDrawinArizona before crossingtheborderinto

Sonora in Agua Prieta. The watershed area in Arizona is 2,650 km2.

Fromthe bordersouth to itsjunction withRío Fronteras, theRíoAgua

Prieta basin has an area of 1,444 km2. Downstream these tributaries

coalesce into the Río Bavispe upstream of the Lázaro Cárdenas (La

Angostura) reservoir (CONAGUA2010). The Río Yaqui itself forms

when the Ríos Bavispe and Áros join north of Sahuaripa.

Plant Collections and Relevés

	 During the spring of 2007, plants were recorded in the seven study

areas on March 23-27, April 3-19, and May 2-5 (fig. 1). A total of

17 transects were done on limestone bedrock and limestone derived

alluvial surfaces. We surveyed a broad range of habitats, including

rocky slopes, peaks, cliffs, alluvial bajadas, arroyos, and cattle tanks.

We collected 216 specimens, which were deposited in herbaria in

Mexico and the United States, including the University of Arizona

(ARIZ), Universidad de Sonora (USON), Arizona State University

(ASU), Universidad Autónoma Nacional de México (UNAM), and

others. Collections were sponsored under a Secretaría de Medio

Ambiente y Recursos Naturales (SEMARNAT) permit to J. Jesús

Sánchez-Escalante (USON). Semi-quantitative relevés were done to

characterizethevegetationandassessspeciesabundance.Relevéswere

about 0.5 hectares(50 x 100 m = 5,000 m2) in area (Mueller-Dombois

andEllenberg,1974). Allperennialplantswere recordedandassigned

relative abundances of rare (1-2 individuals), uncommon, common,

or abundant (the most numerous); annual species were registered as

present, whether alive or dead. This is a rapid assessment method

readilycomparable amongdifferenttransects.Plant collections,observations from the relevés, and photographs are available online in the

Madrean Archipelago Biodiversity Assessment (MABA)/Southwest

Environmental Information Network (SEINet) database (Madrean.

org).Specimenswere identifiedintheUniversityofArizonaherbarium

or duplicates were provided to specialists. Nomenclature mostly follows Van Devender et al. (2010) and updates in SEINet. Only taxa

identified to the species level were used to compare among transect

floras. Plant collections, observations, and photographs are available

in the SEINet/MABA database. A complete list of the transect floras

will be available as a Research Species list in the database.

Results

Vegetation

	 The vegetation on all transects was Chihuahuan desertscrub with

varying importance of desert grassland plants. Two simple measures

of importance were used: (1) the number of transects a species occurred on, and (2) a relative abundance of abundant on at least one

transect. A total of 47 species occurred on 10 or more of the 17 transects, and 15 species were abundant on at least one transect (table 2).

The data indicate that the shrubs Acacia neovernicosa, Calliandra 

eriophylla, Fouquieria splendens, Larrea divaricata, Menodora scabra, Parthenium incanum, Rhus microphylla, and Senna wislizenii;

the succulents Agave palmeri, Cylindropuntia spinosior, Dasylirion 

wheeleri, Echinocereus fendleri, and Opuntia phaeacantha; and perennial grasses(seven species) are characteristic plants on limestone.

Additional typical limestone plantsinclude the shrubs Dalea formosa 

and Flourensia cernua, the perennial herbs Astrolepis cochisensis, 

Bahia absinthifolia, Chamaesaracha sordida, Dalea wrightii, Physaria 

fendleri, Polygala macradenia, and Tiquilia canescens, the perennial grass Panicum hallii; and the annuals Hedeoma nanum, Linum 

puberulum, and Thymophylla pentachaeta.

Figure 1—Map of study areas: AL = El Álamo, AP = Agua Prieta, CA = 

Rancho la Calera, CL = Cerro Caloso, JN = Cerrito Los Janitos, MG = 

Magoffin Ranch, PB = Rancho Puerta Blanca.

Table 2—Plants on ten or more transects on bedrock limestone in 

Arizona, Chihuahua, and Sonora. Species abundant on at least 

one transect in bold. NT = number of transects.

Species NT Species NT

SHRUBS SUCCULENTS (cont’d)

Fouquieria splendens 17 Yucca baccata 12

Acacia neovernicosa 16 Opuntia engelmannii 11

Rhus microphylla 15 N = 9

Aloysia wrightii 13 PERENNIAL HERBS

Larrea divaricata 12 Astrolepis cochisensis 16

Condalia warnockii 11 Croton pottsii 15

Senna wislizenii 11 Chamaesaracha sordida 13

Eysenhardtia orthocarpa 10 Bahia absinthifolia 12

Flourensia cernua 10 Physaria fendleri 12

 N = 9 Tiquilia canescens 11

SUBSHRUBS Dalea wrightii 10

Calliandra eriophylla 16 Polygala macradenia 10

Menodora scabra 16 N = 8

Parthenium incanum 16 PERENNIAL GRASSES

Zinnia acerosa 15 Bouteloua curtipendula 16

Dalea formosa 14

Aristida purpurea var. 

nealleyi 13

Hibiscus coulteri 14 Heteropogon contortus 13

Hibiscus denudatus 13 Bouteloua eriopoda 12

Gutierrezia microcephala 11 Dasyochloa pulchella 11

 N = 8 Panicum hallii 11

WOOODY VINE Aristida ternipes var. ternipes 10

Cottsia gracilis 14 N = 7

 N = 1 ANNUALS

SUCCULENTS Hedeoma nanum 16

Cylindropuntia spinosior 15 Thymophylla pentachaeta 14

Dasylirion wheeleri 15 Ditaxis neomexicana 13

Echinocereus fendleri 15 Linum puberulum 12

Opuntia phaeacantha 15 Galium proliferum 10

Agave palmeri 14 N = 5

Mammillaria grahamii 14 Total: 47 taxa

Cylindropuntia leptocaulis 12
USDA Forest Service Proceedings RMRS-P-67. 2013 233

Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico Van Devender and others

	 Other typical limestone plants found on fewer than 10 transects or

not scored abundant were the shrubs Fraxinus gooddingii, Mortonia 

scabrella, Quercus pungens, andRhus virens subsp. choriophylla; the

subshrubsCyphomeris gypsophiloides, Mentzelia oligosperma,Ruellia 

parryi, Salvia parryi, and Thymophylla acerosa; the perennial herbs

Dalea pogonathera, Euphorbia villifera, Phyllanthus polygonoides, 

Polygala scoparioides, Senna bauhinioides, S. lindheimeriana, 

Thamnosma texana, and Verbesina longipes; and the annuals Dalea 

nana var. carnescens and Iva ambrosiifolia. Vauquelinia californica 

var. pauciflora is a prominent shrub in the Magoffin Hills that is rare

in Sonora. It was only found in the Rancho La Calera area, where it

was a local dominant.

Flora Composition

	 We documented 236 taxa in 170 genera and 54 familiesin Chihuahuan desertscrub on the 17 transects. The families with most species

wereAsteraceae(36taxa,15.2%),Poaceae(33taxa,13.9%),Cactaceae

(21taxa,8.8%),Fabaceae (18,taxa,7.6%),Malvaceae andPteridaceae

(8 taxa each, 3.3%), and Euphorbiaceae (7 taxa, 2.9%). The genera

with the most species were Opuntia (6 taxa), Dalea, Bouteloua, 

Echinocereus, and Polygala (5 taxa each), Cylindropuntia, Abutilon, 

Aristida, and Muhlenbergia (4 taxa each), and Brickellia, Eragrostis, 

Physaria, and Senna (3 taxa each). The life forms on transects were

herbs (57.2%, including grasses), woody plants (31.3%), and succulents (11.4%). Of the dicot herbs (102 taxa), 66 taxa are perennial

and 36 taxa annual, while 27 of 33 grass taxa were perennial and six

annual taxa. The sampling yielded 51 to 119 taxa/transect (average

73.4 taxa/transect).

Non-Native Species—Floras on limestone in the study area

only had six (2.5%) non-native species: Eragrostis lehmanniana, 

Eragrostis cilianensis, Erodium cicutarium, Fumaria parviflora, 

Salsola tragus, and Sisymbrium irio. None of them are invasive in

these areas. Eragrostis lehmanniana is an African bunchgrass that is

a serious invasive that displaces native species in desert grassland in

southeastern Arizona but less so in adjacent Sonora.

	 Severalinvasive species not encounteredon the limestone transects

are expanding into nearby areas.TheAfrican buffelgrass(Pennisetum 

ciliare) has been widely planted in the Sonoran Desert inArizona and

Sonora. This subshrubby grass is an aggressive competitor for water

and nutrients and introduces fire as an ecological processinto non-fire

adapted desertscrub, thornscrub, and tropical deciduous forest. It is a

seriousinvasive speciesin areas mostly below 1060 m elevation. Van

Devender andDimmitt(2006)reportedthe expansionofbuffelgrassto

elevations of 1195 to 1395 m as far east as 44 km east of Agua Prieta

in northeastern Sonora. In August 2007, it was found at 1565 m in

desert grassland 46.5 km east of Agua Prieta. In September 2006, it

was found at 1287 m elevation within a kilometer of the La Calera

limestone transects. Amodest-sized plant appeared in a small arroyo

that had been surveyed a number of times previously. Thislocation is

well away from a paved road and the nearest plants observed along

the Agua Prieta-Nacozari highway are ca. 5.5 km to the east. Also,

two very small individuals were found in desert grassland on a new

dirt pile on Rancho El Porvenir, east of Cabullona (close to the El

Caloso transect), where dirt roads were recently bulldozed as part

of mineral prospecting activities. Either seeds were transported to

the area by vehicles, or blown from the highway about 6.5 km to

the west. Although buffelgrass is a serious invasive in Chihuahuan

desertscrub in the Big Bend of Texas, it mostly does not occur away

frompaved highwaysin northeastern Sonora.Increased temperatures

(global warming) could change itsstatusin northeastern Sonora from

non-threatening to invasive.

	 Saharamustard(Brassica tournefortii)is aNorthAfrican-European

spring annual that is a serious invasive in low elevation (below ca.

800 m) Sonoran desertscrub in northwestern Sonora, southwestern

Arizona, and southeastern California (Dimmitt and Van Devender,

2009). Its arrival and expansion in other parts of Sonora is more

recent, as it disperses mostly along major highways. It was present

in agricultural areas near Hermosillo in central Sonora in 1995 and

west of Bacobampo in southern Sonora in 1993. Between 2003 and

2005, itspread southward along Mexican Federal Highway 15 (MEX

15) about 180 km from Magdalena de Kino and Santa Ana south to

Hermosillo. By 2008, it was in Guaymas, another 128 kilometers to

the south.In northernSonora,itwasfound in 2005 at 1109melevation

in Cíbuta, 29 km south of Nogales on the Arizona border. By 2007,

it was 25-41 km east of Agua Prieta along MEX 2 (156-172 km east

of Nogales) at 1126-1295 m in cottonwood-willow riparian forest,

Chihuahuan desertscrub, and desert grassland. By 2010, it was 48.5

km east ofAgua Prieta at 1455 m elevation in desert grassland. MEX

2 is the major truck route between Baja California and Chihuahua,

which crossesthe Lower Colorado River Valley Sonoran desertscrub

areas invaded by B. tournefortii.

	 PriortoKearney and Peebles(1960), theEuropeanEruca vesicaria

ssp. sativa was known in Arizona from a few urban and vegetable

garden settingsin the Tucson and Phoenix areas. Since 1960, arugula

has been known from the agricultural areas in the Sonoran Desert at

Gila Bend. In the spring of 2005, it was the most abundant annual

for 70 km along Interstate 8 westward from Gila Bend in an area of

several hundred square kilometers (Mark A Dimmitt, pers. comm.,

2006).The plantsreached about a metertall and carpeted undisturbed

valleys and rocky slopes in open Sonoran desertscrub. Alarms were

raised concerning its invasive potential in the Sonoran Desert, especially in Organ Pipe Cactus National Monument. In March 2012,

it was observed just across the border in Sonoyta, Sonora. Another

aspect of its ecological potential was uncovered with its discovery

in 2007 along MEX 2 east and west of Agua Prieta, in Chihuahuan

desertscrub at 1215 to 1321 m elevation. In 2009-2010, it was found

51.2 to 53.5 km east of Agua Prieta at 1495-1499 m elevation in

desert grassland. In 2007, it was common in desert grassland at 1360

to 1533 m elevation in the Janos area in northwestern Chihuahua.

These observations suggest that this mustard is potentially invasive

in higher elevation habitats. A 2012 collection near Moctezuma in

central Sonora in foothills thornscrub at 637 m elevation suggests

that it may expand its range into more tropical areas as well.

Bromus rubens is aNorthAfrican-Eurasian spring annual grassthat

is a serious invasive in California and Arizona. Fires fueled by dense

spring growth are devastating to Mohave and Sonoran desertscrub,

whose dominants are not well adapted to fire. It is not common or

invasive in Sonora. Prior to this study, it was previously known from

the Sierra Pinacate in northwestern Sonora (Felger 2000) and the Sierra el Humo south of Sásabe. In 2007-2008, it was found in Arroyo

Guadalupe on Rancho Puerta Blanca, Municipio of Agua Prieta. In

May 2010 it was found 41.3 km east of Agua Prieta along MEX 2

at 1323 m in desert grassland. This species appears to be spreading

from disturbed areas along the border roads in Arizona into the Río

Yaqui drainages and south into Sonora. 

Biogeography

	 Our collections of Cyphomeris gypsophiloides, Echinocereus 

fasciculatus var. fasciculatus, Euphorbia villifera, Mentzelia oligosperma, Polygala lindheimeri var. parvifolia, Quercus pungens, and
234 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico

Zephyranthes longifolia areprobablythefirstrecordsforthespeciesfor

Sonora. The observation of Ferocactus wisizenii at Cerro de Cal may

be the firstrecord forChihuahua. Manymore species were previously

known from Sonora from only a few records, including Bernardia 

myricifolia, Castilleja lanata, Cheilanthes villosa, Linum lewisii, 

Mammilaria lasiacantha, Notholaena aschenborniana, Petrophytum 

caespitosum, Phyllanthus polygonoides, Physaria fendleri, Polygala 

scoparioides, Ruellia parryi, Salvia parryi, Senna lindheimeriana, 

and Vauquelinia californica ssp. pauciflora. 

	 Anumber of plants on the transects are mostly western speciesthat

reach as far east as the El Paso, Texas, area including Agave palmeri, 

Aristida ternipes var. ternipes, Bebbia juncea, Gutierrezia microcephala, Mammillaria grahamii, and Yucca baccata. Cylindropuntia 

spinosior is replaced to the east by the closely related C. imbricata. 

Othertransectspecies clearly are notin themainChihuahuan desertto

the east, including Echinocereus pseudopectinatus, Encelia farinosa, 

Hermannia pauciflora, and Vauquelinia californica. Cylindropuntia 

thurberi and Eysenhardtia orthocarpa are tropical/madrean species

reaching their northern range limits in northeastern Sonora or southeastern Arizona.

	 Wentworth (1982) documented the distributions of plant species

on limestone bedrock and calcareous soils at 1400 to 1900 m elevations in the Mule Mountain. Of 232 species in these habitats, 127 are

shared with the transect floras of this study. The 72 species not on

the transects,includingBouvardia ternifolia, Cercocarpus montanus, 

Garrya wrightii, Quercus arizonica, and Pinus discolor were in this

mesic limestone habitat in oak woodland zone, which is 400 m above

the La Calera study area.

Discussion

	 The Chihuahuan Desert in Sonora is represented by isolated areas

in the northeastern corner of the state, mainly in the municipios of

Agua Prieta, Naco, Fronteras, Bavispe and Nacozari de García. The

transition to the Sonoran Desert is in the west side of the Sierra El

Tigre along Presa Angostura on the Río Bavispe. Brown and Lowe

(1980) mapped the vegetation as an eastern patch of the Arizona

Upland subdivision of the Sonoran Desert, but in actuality, it is a

distinct area, perhapstheElTigre subdivision.FartherwestinArizona

(Vekol and Waterman Mountains) and northwestern Sonora (Sierra

del Viejo) at lower elevations, limestone substrates support Sonoran

desertscrub rather than Chihuahuan desertscrub

	 The vegetation and geology on most of the areas surveyed in this

study were basically similar i.e., exposed limestone bedrock and

Chihuahuan desertscrub with mixed desert grassland elements with

the shrubs Acacia neovernicosa, Calliandra eriophylla, Fouquieria 

splendens, Parthenium incanum, andperennialgrasses.Theabundance

of grasses on the bedrock limestone habitats varied, and generally

increased with elevation. The areas with the most grasses were the

Magoffin Hills and Rancho La Morita habitats at 1400 to 1500 m

elevation. The abundance of grasses also reflected grazing pressure.

Grasseswere lush on portions ofRancho PuertaBlanca on theCuenca

los Ojos Foundation reserve south of MEX 2 where livestock were

removed in about 2002, but were sparse on adjacent grazed areas on

Ejido Guadalupe.

	 While recognizing the basic similarities among all limestone

habitatssurveyed, there were local differencesin each area surveyed.

The desertscrub on the Cerro de Cal in Chihuahua was the most xerophytic, likely because the Sierra San Luis blocks winter rains and

it was heavily grazed. Yet Eriogonum wrightii was only seen on the

Magoffin Hills and Cerro de Cal transects. Vauquelinia californica 

ssp. pauciflora wasseen on the Magoffin Hills andRancho LaCalera,

where the vegetation was much more open, with less grass and more

limestone species, and much steeper slopes. On Cerrito Los Janitos,

the vegetation and substrate were very similar to the nearby Rancho

La Calera but without V. californica, and with Mimosa biuncifera. 

Quercus pungens is a shrubby oak found on limestone substrates

from Texas to southeastern Arizona. Although not previously known

from Sonora, it was found on eight areas in this study, all on steep

limestone slopes and ridge tops. Koeberlinia spinosa was only seen

on two adjacent transects on Rancho Puerta Blanca north of Arroyo El Guadalupe. Bernardia myricifolia was only found on the

two Rancho La Calera transects. Dodonaea viscosa was common

on the El Álamo transect, but not found elsewhere. Bebbia juncea, 

Crossosoma bigelovii, Cylndropuntia thurberi, Encelia farinosa, 

Hermannia pauciflora, and Lantana macropoda were only found on

Cerro El Caloso east of Cabullona. Dalea nana, Elytraria imbricata, 

Euphorbia villifera, Petrophytum caespitosum, Polygala lindheimeri, 

and Senna lindheimeriana were only found on the Rancho La Morita

transects.

	 In summary, the species composition of Chihuahuan desertscrub

at its western limits is a mixture of typical Chihuahuan species that

reachTexas andCoahuilawithwesternandtropicalspeciesinassociations that are regionally distinct from other parts of the Chihuahuan

Desert. This should be considered when evaluating these areas for

conservation needs.

Acknowledgments

	 We thank Mima Falk, Marc A. Baker, Don Jesús Barra-Moreno,

DoñaAlicia Oroz-Gámez, Don Pedro García, Octavio García,John F.

Wiens, Jeffrey Moore, and Manuel Espericueta for help in the field;

Section 6 Grant from U.S. Fish and Wildlife Service. We thank Valer

and JosiahAustin andBobMinckley for accessto private lands owned

by the Cuenca Los Ojos Foundation. We thank Elizabeth Makings

and C. David Bertelesen for their careful review of the paper.

References

Brown, David E. 1982. Chihuahuan Desertscrub. In Brown, David E. ed. Biotic Communities of theAmerican Southwest – United States and Mexico.

Desert Plants. Special Issue. 4 (1-4): 169-179.

Brown, David E.; and Charles H. Lowe. 1980. Biotic Communities of the

Southwest.Gen.Tech.Rep.RM-78(map).U.S.DepartmentofAgriculture,

Forest Service, Rocky Mountain Forest and Range Experiment Station,

Fort Collins, Colorado.

CONAGUA. 2010. Determinación de la disponibilidad de agua en el acuífero

2629, río Agua Prieta, estado de Sonora. 30 p.

Dimmitt, Mark A.; and Van Devender, Thomas R. 2009. Sahara mustard

(Brassica tournefortii):Anew,seriousthreat to lowdesert ecosystemsin the

southwestern United States and northwestern Mexico. Pg. 241-246 in Van

Devender,ThomasR.;Espinosa-García,Francisco J.;Harper-Lore,Bonnie

L.; and Hubbard, Tani eds., Invasive Plants on the Move. Controlling them

in NorthAmerica. Proceedings ofWeedsAcrossBorders 2006Conference,

Hermosillo, Sonora, May 25-28, 2006; Tucson, Arizona.

Felger, Richard S. 2000. Flora of the Gran Desierto and Río Colorado of

northwestern, Mexico. University of Arizona Press, Tucson, AZ. 673 p.

Kearney,Thomas H.; and Peebles, Robert H. 1960.Arizona Flora, 2nd edition

with supplement by J. H. Thomas, J. H.; and E. McClintock, E. University

of California Press, Berkeley. 1085 p.

Lopresti, Vincent 1984. Coryphantha robbinsorum en México. Cactáceas y

Suculentas Mexicanas, 29: 81-83.

Lowe, Charles H. 1964. Arizona’s Natural Environment. Landscapes and

Habitats. The University of Arizona Press.136 p.
USDA Forest Service Proceedings RMRS-P-67. 2013 235

Flora of Chihuahuan Desertscrub on Limestone in Northeastern Sonora, Mexico Van Devender and others

Mueller-Dombois, D., and Ellenberg, H. 1974. Aims and Methods of Vegetation Ecology. John Wiley & Sons, New York.

Reza, Francisco; and López, Gerardo. Sonora se congela: -16 oC. El Imparcial. Diario Independiente de Sonora: Hermosillo, México. Febrero 4,

2011. p. A1, col. 1.

Rzedowski, Jerzy. 1978. Vegetación de México. Editorial Limusa. México,

D.F. 432 p.

Shreve, Forrest. 1922. Conditions indirectly affecting vertical distribution on

desert mountains. Journal of Ecology, 3: 269-274

Shreve, Forrest. 1942. The desert vegetation of North America. Botanical

Review, 8: 195-246

Shreve, Forrest and Wiggins, Ira L.1964.Vegetation and Flora of the Sonoran

Desert.Volume one. Stanford University Press. Stanford,California. 840 p.

Universidad Autónoma de Tamaulipas. 2003. Manifestación de impacto ambiental modalidad particular. Comisión Federal de Electricidad. Proyecto

ciclo combinado Agua Prieta II. 151 p.

VanDevender,Thomas.R.;Felger,RichardF.;Fishbein,Mark;Molina-Freaner,

Francisco; Sánchez-Escalante, J. Jesús; and Reina-Guerrero, Ana L. 2010.

Biodiversidad de las plantas vasculares. Pp. 229-262 in Molina-Freaner,

Francisco; andVanDevender,ThomasR. eds.,DiversidadBiológica de Sonora,UniversidadNacionalAutónoma deMéxico,Hermosillo. http://www.

skyislandalliance.org/Sonoran%20vascular%20plants-VanDevender%20

et%20al.2010.pdf

Van Devender, Thomas R.; and Reina-G,Ana Lilia. 2007. Final report on surveys of Coryphantha robbinsorum and other rare plantsin northern Sonora

and adjacent Chihuahua. U.S. Fish andWildlife Service Section 6 contract.

VanDevender,ThomasR.; andDimmitt,MarkA. 2006. FinalReport on “Conservation ofArizona Upland Sonoran Desert Habitat. Status and Threats of

Buffelgrass (Pennisetum ciliare) in Arizona and Sonora.” National Plant

Conservation Initiative Project #2004-0013-003.

Wentworth, Thomas R. 1982. Vegetation and flora of the Mule Mountains,

Cochise County, Arizona. Journal of the Arizona-Nevada Academy of

Science, 17: 29-44.

Whittaker, Robert H. and. Niering, William A. 1968. Vegetation of the Santa

Catalina Mountains, Arizona. IV. Limestone and acid soils. Journal of

Ecology, 56: 523-544

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
236 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Preliminary Flora of the Sierra Bacadéhuachi, Sonora, 

Mexico

 Thomas R. Van Devender and Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona

George M. Ferguson

University of Arizona, Tucson, Arizona

George Yatskievych

Missouri Botanical Garden, St. Louis, Missouri

Beatriz E. Loyola-Reina

JRM Consultores, Hermosillo, Sonora, Mexico

Gertrudis Yanes-Arvayo, and Maria de la Paz Montañez-Armenta

Universidad de la Sierra, Moctezuma, Sonora, Mexico

John L. Anderson

Bureau of Land Management, Phoenix, Arizona

Stephen F. Hale

EcoPlan Associates, Inc., Mesa, Arizona

Sky Jacobs

Tucson, Arizona

Abstract—The Sierra Bacadéhuachi in east-central Sonora is the westernmost mountain range in the Sierra 

Madre Occidental (SMO), located east of Bacadéhuachi, Municipio de Bacadéhuachi, 34 km east of the 

Chihuahua border, and 165 km south of the Arizona border. The vegetation ranges from lowland foothills 

thornscrub up through desert grassland to oak woodland and pine-oak forest. The flora was sampled in 

December 1995, July 2008, June, August, and September 2011, and March 2012. The flora totals 442 taxa 

in 297 genera and 96 families. The most species-rich families and genera are Asteraceae (53 taxa), Fabaceae 

(48 taxa), Poaceae (46 taxa), Quercus (11 species), Cheilanthes (8 species), Muhlenbergia (7 species), 

Bouteloua (6 species), and Pinus (5 species). Twenty-two species are non-native (5.0%), 10 of them grasses. 

No species are endemic. Although tree composition and structure of the upland woodlands and forests are 

similar to the Yécora, Sonora, area to the south, the preliminary Bacadéhuachi flora is less diverse.

Introduction

Rzedowski (1978) pointed out that in Mexico in general, the greatest plant species diversity was in pine-oak forest at high elevation.

Conservation International in 2007 named the Madrean Pine-oak

Woodlands as a global biodiversity hotspot. This area included the

SierraMadreOriental in easternMexico, the SierraMadreOccidental

(SMO) in western Mexico, and the Sky Island ranges north of them

into Arizona, New Mexico, and Texas. The Sierra Madre Occidental

extends in western Mexico from Zacatecas and Jalisco north to Chihuahua andSonora,Mexico(Rzedowski1978).Highest elevationsfor

this cordillera exceed 2800m, andits continuity provides animportant

route for fauna and flora dispersing between tropical and temperate

pine forests, andbetween tropicalforests andnorthern grasslands.The

continental divide follows the Sierra Madre northward to the upper

Río Gavilán-Sierra Huachinera on theChihuahua-Sonora border, and

then through the isolated Sierra Púlpito and Sierra San Luisin Sonora

and the Animas Mountains in New Mexico.

	 Van Devender and others (2010) summarized the flora of Sonora.

The current flora documented with voucher specimens is 3672 taxa.

Extensive inventoriesin the SMO in east-central Sonora documented

1774 taxa (48.6% of the state flora) in the Municipio de Yécora

(3300 km2; Van Devender and others 2005; Van Devender and ReinaGuerrero, unpublished).We estimate that the number of plantspecies

in the northern SMO and Sky Island ranges in Sonora to be at least

2600 species (70.8% of the state flora). In this paper we present a
USDA Forest Service Proceedings RMRS-P-67. 2013 237

Preliminary Flora of the Sierra Bacadéhuachi, Sonora, Mexico Van Devender and others 

preliminaryflora oftheSierraBacadéhuachiin thewesternmostSMO

to further document the biodiversity in this regional hotspot.

Methods

Study Area

	 The Sierra Bacadéhuachi isthe westernmost mountain range in the

Sierra Madre Occidental, located east of Bacadéhuachi, Municipio

de Bacadéhuachi, 34 km east of the Chihuahua border, and 165 km

south of theArizona border. Field activities were centered on Rincón

de Guadalupe (29°50’40”N 108°58’37”W, 1680 m elevation) owned

by the Catholic Archdiocese of Hermosillo. The southern portion of

the Sierra Bacadéhuachi is in the Ríos Nácori Chico and Riíto drainages, with the Río Bacadéhuachi on the north, and looping around

to west, all part of the greater Río Yaqui watershed. Elevations range

from 700 m at Bacadéhuachi to 2400 m on the highest peak, an

elevational range of about 1700 meters. Plants were observed and

collected in Bacadéhuachi and the nearby arroyo, along the old and

new roads between Bacadéhuachi and Nácori Chico, and the main

Sierra Bacadéhuachi along the road to Rincón de Guadalupe and the

high peaks above.

Plant Collections

	 Plants were collected under the SEMARNAT permit to J. Jesús

Sánchez-Escalante,Curator ofthe Universidad de Sonora Herbarium

(USON). Specimens were deposited into herbaria at the University

of Arizona (ARIZ), USON, Arizona State University (ASU), and the

Universidad Nacional Autónoma de México (MEXU). A few early

plant records from the Bacadéhuachi area were found in the Southwest Environmental Information Network (SEINet) online database.

In July 1940, Stephen S. White (1948), as part of his flora of the Río

Bavispe region, made collections in La Palmita and the Aguage de

Bacatejaca between Granados and Bacadéhuachi, and at La Galera

(5 km N of Bacadéhuachi, 29°51’12”N 109°07’57”W, 715 m elevation). Between Bacadéhuachi and Nácori Chico, Joe T. Marshall

collected Quercus tuberculata in June 1953 and Joe T. Marshall and

Richard F. Felger collected Q. emoryi in June 1960. These were part

of Marshall’s general tree inventories for his 1957 book “Birds of the 

Pine-oak Woodland in southern Arizona and Adjacent Mexico.”

	 Otherwise, the Sierra Bacadéhuachi plant collections and observations reported here were by the authors. Ferguson along with Mark

Kaib, H. Mata-Mangueros, andA. Valencia-Cordoba visited the area

inDecember1995.VanDevender,Reina-Guerrero, andLoyola-Reina

did plant transects in 11 thornscrub (FTS) areas along the route of

the new road (completed in 2011) between Bacadéhuachi and Nácori

Chico, all on the southern slopes of the Sierra de Bacadéhuachi. All

authors participated in a Madrean Archipelago Biodiversity Assessment (MABA) Expedition trip to the Sierra Bacadéhuachi in August

2011. Van Devender and Reina-Guerrero, with the help of David G.

Bygott, Marisa Rice, and Robert Villa, observed plants in the same

area in June and September 2011 and March-April 2012. Nomenclature mostly follows the list in Van Devender and others (2010),

which is available and updated in the Madrean Archipelago BiodiversityAssessment (MABA)/Southwest Environmental Information

Network (SEINet) online database (Madrean.org). Plant collections,

observations, and photographs from the study area are available in

this database. A complete list of the Sierra Bacadéhuachi flora will

be available as a Research Species list in the database.

Results

Vegetation

	 The vegetation on the southern lower flanks of the Sierra Bacadéhuachi is foothills thornscrub (FTS). This is the northern part of

a very large expanse of FTS southward to where the Ríos Áros and

Bavispe join to become the Río Yaqui, and merges into tropical deciduous forest (TDF) near San Nicolás.

	 In theSierraBacadéhuachi, oakwoodland is above desert grassland

or FTS. Quercus chihuahuensisisthe most common oak, with rare Q. 

emoryi and Q. tuberculata. With increasing elevation Q. arizonica,

Q. oblongifolia. Quercus viminea, Pinus chihuahuana, and Arbutus 

arizonica become more common. At higher elevations, there is a

mosaic of oak woodland on drier slopes and pine-oak forest in more

mesic areas. With the addition of Q. hypoleucoides, Q. jonesii (= Q. 

coccolobifolia), Q. mcvaughii, and Q. tarahumara, the oak diversity

in higher woodlands is comparable to the Yécora area in the SMO

(166 km south), with Q. durifolia notably absent. Pine-oak forest is

dominated by Pinus engelmannii, with P. arizonica and P. yecorensis 

locally common, and mixed oaks, Arbutus xalapensis, and Juniperus 

deppeana var. patoniana. At the highest elevations, P. strobiformis 

is present, and Q. rugosa is rare, with no presence of a mixed conifer

zone (no Abies or Pseudotsuga).

	 Riparian vegetation in drainagesislinear, transecting other upland

vegetation types.In the lowland tropical zone,riparian habitats are not

well developed because FTS extendsinto the bottom of the drainages

in most areas. Populus monticola and Salix bonplandiana were seen

at Rancho el Carricito (916 m). The absence of Guazuma ulmifolia

is of interest. This riparian tree is common and widespread in tropical southern Sonora, but apparently reaches its northern limit north

of Tepache (53 km west-southwest) and on Rancho el Bábaco north

of Sahuaripa (45 km south). White (1948) collected P. fremontii at La

Galera (715m) along Río Bacadéhuachi at FTS elevation. The arroyo

at Rancho Agua Nueva (10 km east of Bacadéhuachi) in the desert

grassland-FTS transition has P. fremontii and Platanus wrightii (1145

m). Salix gooddingii was only seen in Arroyo la Matancita (1600 m)

in oak woodland near Rincón de Guadalupe. Riparian treesinArroyo

Campo los Padres at Rincón de Guadalupe are Alnus oblongifolia,

Cupressus arizonica, and P. wrightii. We follow the classification

used in Martin and others (1998) for vegetation names.

Flora

	 We identified 442 plant taxa in 298 genera and 96 families in the

Sierra Bacadéhuachi area. The families with the most species are

Asteraceae (53 taxa, 12.0%), Fabaceae (48 taxa, 10.9%), Poaceae

(46 taxa, 10.4%), Euphorbiaceae (18 taxa, 4.1%), Pteridaceae (15

taxa, 3.4%), and Cactaceae (14 taxa, 3.2%).The genera with the most

species are Quercus (11 taxa), Cheilanthes (8 taxa), Muhlenbergia

(7 taxa), Bouteloua (6 taxa), Asclepias, Acacia, Euphorbia, Mimosa,

and Opuntia, and Pinus (5 taxa each), and Ambrosia, Salvia, and

Solanum (4 taxa each). The life forms in the Bacadéhuachi flora are

herbs (49.5%, including grasses and sedges), woody plants (35.3%),

and succulents (5.0%). Trees with 51 species (11.5%) are important

in the flora and except for desert grassland, all of the vegetation types

in the study area. The dicot herbs (219 taxa) are perennial (148 taxa)

and annual (71 taxa). The grasses and sedges (46 taxa) are perennial

(25 taxa) and annual (21 taxa).

	 Only22speciesarenon-native(5.0%),includingtengrasses.Melinis 

repensis anAfro-Asian bunchgrass presentin desert grassland,which

has the potential to dominate and displace native perennial grasses.
238 USDA Forest Service Proceedings RMRS-P-67. 2013

 Van Devender and others Preliminary Flora of the Sierra Bacadéhuachi, Sonora, Mexico 

Brassica tournefortii, Cynodon dactylon, Pennisetum ciliare, and

Sorghum halepense are invasive elsewhere in Sonora (Van Devender

andothers 2009), butnotintheBacadéhuachi area.Pennisetum ciliare

is often planted in cleared desertscrub, thornscrub, and TDF in other

areas in Sonora. Surprisingly, the only observation of it in the study

area was 10.2 km east-southeast of Bacadéhuachi. Disturbance along

the new highway will probably allow B. tournefortii, P. ciliare, and

other non-native species to disperse into relatively intact FTS. Marrubium vulgare was locally common at Agua Nueva. Horehound or

marrubio is a North African-Eurasian perennial herb that is almost

always found in Sonora around stock corrals. It persists long after

ranches are abandoned, but seldom spreads into natural habitats.

	 Asingle B. tournefortii was collected in the town ofBacadéhuachi.

Sahara mustard is a North African-European spring annual that is a

seriousinvasive in lowelevationSonoran desertscrub in northwestern

Sonora,southwesternArizona, andsouthwesternCalifornia (Dimmitt

and Van Devender 2009). Its arrival and expansion in other parts of

Sonora is more recent. It was present in agricultural areas near Hermosillo in central Sonora in 1995 and west ofBacobampo in southern

Sonora in1993.Itsdispersalmostlyfollowsmajorhighways.Between

2003 and 2005, it spread southward about 180 km along Mexican

Federal Highway 15 (MEX 15) from Magdalena de Kino and Santa

Ana south to Hermosillo. By 2008, it was in Guaymas, another 128

km to the south. In northern Sonora, it wasfound at 1109 m elevation

in Sierra Cíbuta, 29 km south of Nogales on the Arizona border in

2005. By 2007, it was 25 to 41 km east of Agua Prieta along MEX 2

(156-172 km east of Nogales) at 1126-1295 m in cottonwood-willow

riparianforest,Chihuahuandesertscrub, anddesertgrassland.By2010,

B. tournefortii was 48.5 km east of Agua Prieta at 1455 m elevation

in desert grassland. Our 2012 collections of it in Baviácora in the Río

Sonora valley,Granadosin theRíoBavispeValley, andBacadéhuachi

in the Río Yaqui drainage demonstrate that its dispersal into FTS in

eastern Sonora is just beginning. These localities are 97 to 190 km to

the northeast and east-northeast ofthe previous Hermosillo localities.

Discussion

	 In comparison with the intensely inventoried flora of the Municipio de Yécora, Sonora (36 trips in 1995-2008 by Van Devender

and Reina-Guerrero) with 1774 plant taxa in 3300 km2, the flora of

the Sierra Bacadéhuachi with 442 taxa is preliminary. The study area

with an elevational range of 1700 m and a maximum elevation of

2400 m comparable to those of the Yécora area (1660 m range, 2140

maximum elevation) potentially has a much richer flora. Its affinities

are strongly with the main SMO and lowland New World tropics of

northwestern Mexico. Desmanthus illinoensis was collected on a

disturbed roadside in FTS at 692 m elevation near Bacadéhuachi in

2008. This prostrate perennial herb with white flowers is widespread

in the eastern United States asfar west as New Mexico, but has not yet

been collected inArizona.The Lupinus aff. argenteuswas collected in

pine-oakforest at ca.2300melevationnearCerrolaPlaca inDecember

1995. The nearest known populations are on the Continental Divide

in the northern SMO of Chihuahua (70 km NNW) and in the Santa

Catalina Mountains in Arizona (335 km north-northwest), but the

high-elevation Madrean lupines in Sonora need taxonomic revision.

Hexalectris revoluta was collected in oak woodland at 1680 m elevation at Rincón de Guadalupe in April 2012. This rare orchid, which

was described from near Galeana, Nuevo León, is known fromTexas,

New Mexico, and Arizona. The nearest locality is in the Peloncillo

MountainsofsouthwesternNewMexico(260kmnorth).Ribes dugesi 

is a Madrean pine-oak forest shrub previously only known in Sonora

on Mesa el Campanero nearYécora (166 km south). Adolphia infesta 

collected in FTS at 1120 m at Rancho el Saucito (17 km southeast of

Bacadéhuachi) in July 2008 is a rare spiny shrub in Sonora. Tithonia 

fruticosa is a large shrub/tree sunflower collected on Rancho el Embudo (10.3 km east of Bacadéhuachi) at 1215 m elevation in desert

grassland in March 2012. Previously Carl V. Hartman collected it at

Huehuerachi (Huepaverchi?) near Nácori Chico in December 1890

(NY) on the Lumholtz Expedition (Lumholtz, 1902) and Felger and

Marshall collected it northeast of Nácori Chico in June 1960. This is

the northernmost location for this tropical species.

	 The vegetation of the Sierra Bacadéhuachi is similar to that along

the MEX 16 highway in the Yécora area, except that desert grassland

is present and TDF is not. The northernmost TDF is in the Sierra San

Javier at ca. 28°45’N. The FTS in the Sierra Bacadéhuachi is the

transition fromthe NewWorld tropicsto the northern temperate zone.

To the north, FTS is replaced by desert grassland as winters become

colder and periodic fires become ecological processes. The northern

limits of FTS in Sonora are at about 30°11’N east of Sinoquipe in

the Río Sonora Valley and 30°26’N at Presa Angostura on the Río

Bavispe at the southern end of the Sierra El Tigre.

	 Desert grassland is a vegetation type found from the Mexican

Plateau in Coahuila, Chihuahua, and Durango north to southeastern

New Mexico and westernTexas and west to southeasternArizona and

northwestern Sonora. In Sonora, it occurs as far west as the Sásabe

plains between the Sierra El Humo and the Baboquivari Mountains

in Arizona. In northeastern Sonora, it forms a mosaic with Chihuahuan desertscrub as far south as Colonia Morelos and Carretas on

the north and east sides of the Sierra el Tigre (White 1948). On the

southwestern end of the Sierra Bacadéhuachi there is a large area

of desert grassland dominated by Prosopis velutina at 1150 to ca.

1850 m elevation above FTS. This is one of the southernmost desert

grassland areas in Sonora.

	 The flora of the SMO is not well known. The Yécora area was included in the broad Río Mayo flora area in Martin and others (1998),

but the only detailed publication is on the grasses(Van Devender and

others 2005).The only published partial flora of a Sonoran Sky Island

Mountain range is for the Sierra El Tigre as part of White’s (1948)

flora of the Río Bavispe region. With the MABA/SEINet database,

plant collections and observations from the SMO and various Sky

Island ranges are becoming more available, but are not yet summarized in the literature. The Sierra Bacadéhuachi flora is an important

contribution to the understanding of the biodiversity of the northern

SMO and the Madrean Archipelago.

Acknowledgments

	 We thank Mike Bauer for his loving and meticulous identification

of grasses. Enrique Yescas of SonoraEs magazine greatly facilitated

fieldwork. Father Varela gave permission to stay atRincón de Guadalupe. Reviews by Socorro González-Elizondo and Mark A. Dimmitt

improved the paper.

References

Dimmitt, Mark A.; and Van Devender, Thomas R. 2009. Sahara mustard

(Brassica tournefortii):Anew,seriousthreat to lowdesert ecosystemsin the

southwestern United States and northwestern Mexico. Pg. 241-246 in Van

Devender,ThomasR.;Espinosa-García,Francisco J.;Harper-Lore,Bonnie

L.; and Hubbard,Tani. eds., Invasive Plants on the Move. Controlling them

in NorthAmerica. Proceedings ofWeedsAcrossBorders 2006Conference,

Hermosillo, Sonora, May 25-28, 2006; Tucson, Arizona.

Lumholtz, Carl S. 1902. Unknown Mexico:ARecord of Five Years’Exploration among the Tribes of the Western Sierra Madre; in the Tierra Caliente
USDA Forest Service Proceedings RMRS-P-67. 2013 239

Preliminary Flora of the Sierra Bacadéhuachi, Sonora, Mexico Van Devender and others 

of Tepic and Jalisco; and among the Tarascos of Michoacán. New York:

Charles Scribner’s Sons.

Marshall,JoeT.,Jr. 1957. Birds of the pine‐oak woodland in southernArizona

and adjacent Mexico. Pacific Coast Avifauna, 32: 1‐125.

Martin, Paul S.; Yetman, David A.; Fishbein, Mark; [and others], eds. 1998.

Gentry’s Río Mayo Plants. The Tropical Deciduous Forest and Environs

of Northwest Mexico. Tucson: University of Arizona Press.

Rzedowski, Jerzy. 1978. Vegetación de México. México: Editorial Limusa,

México.

White, Stephen S. 1948. The vegetation and flora of the region of the Río de

Bavispe in northeastern Sonora, Mexico. Lloydia, 11: 229‐302.

VanDevender,Thomas.R.;Felger,RichardF.;Fishbein,Mark;Molina-Freaner,

Francisco; Sánchez-Escalante, J. Jesús; and Reina-Guerrero, Ana L. 2010.

Biodiversidad de las plantas vasculares. Pp. 229-262 in Molina-Freaner,

Francisco; andVanDevender,ThomasR. eds.,DiversidadBiológica de Sonora,Hermosillo:UniversidadNacionalAutónomadeMéxico.http://www.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

skyislandalliance.org/Sonoran%20vascular%20plants-VanDevender%20

et%20al.2010.pdf

Van Devender, Thomas R.; Felger, Richard S.; Reina-Guerrero, Ana L.; and

Sánchez–Escalante,J.Jesús. 2009.Sonora:Non-native and invasive plants.

Pp. 85-124 in Van Devender, Thomas R.; Espinosa-García, Francisco J.;

Harper-Lore, Bonnie L.; and Hubbard, Tani. eds., Invasive Plants on the

Move. Controlling them in North America. Proceedings of Weeds Across

Borders 2006 Conference, Hermosillo, Sonora, May 25-28, 2006; Tucson,

AZ.http://www.skyislandalliance.org/WAB06-c19-Van%20Devender%20

et%20al-Sonoran%20non-natives-Jan08.pdf

Van Devender, Thomas R.; Reeder,John R.; Reeder, Charlotte G.; and ReinaGuerrero, Ana L. 2005. Distribution and diversity of grasses in the Yécora

region of the Sierra Madre Occidental of eastern Sonora, Mexico. Pp.

107-121 in Cartron, Jean-Luc E.; Ceballos, Gerardo; and Felger, Richard

S. eds. Biodiversity, Ecosystems, and Conservation in northern Mexico.

New York: Oxford University Press.
240 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Comparison of the Tropical Floras of the Sierra la Madera 

and the Sierra Madre Occidental, Sonora, Mexico 

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona

Gertrudis Yanes-Arvayo

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Ana Lilia Reina-Guerrero

Sky Island Alliance, Tucson, Arizona

Melissa Valenzuela-Yánez, Maria de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Abstract—The floras of the tropical vegetation in the Sky Island Sierra la Madera (SMA) near Moctezuma in 

northeastern Sonora (30°00’N 109°18’W) and the Yécora (YEC) area in the Sierra Madre Occidental (SMO) 

in eastern Sonora (28°25’N 109°15”W) were compared. The areas are 175 km apart. Tropical vegetation 

includes foothills thornscrub (FTS) in both areas and tropical deciduous forest (TDF) in the Yécora area. A 

total of 893 vascular plant taxa are known from these areas with 433 taxa in FTS and 793 in TDF. FTS in SMA 

and YEC (near Curea) had 220 and 298 taxa, with most of them also in TDF (69.5% and 82.9%). Only 83 

taxa in TDF were shared between SMA and YEC (37.7% and 27.9% of the floras). The 49 FTS species in SMA 

but not YEC were not in TDF either, reflecting biotic influences from the Sonoran Desert (10), southwestern 

United States (8), Madrean Archipelago (6), and a few from oak woodland and tropical western Mexico. 

One species (Pseudabutilon thurberi) is endemic to central Sonora and adjacent Arizona. Affinities to the 

New World tropics are very strong in both areas. The structural dominants that define FTS are widespread, 

but composition varies greatly locally. 

Introduction

	 The Madrean Archipelago is the area of isolated Sky Island (SI)

mountain ranges between the Sierra Madre Occidental (SMO) in

Sonora and Chihuahua and the Mogollon Rim in New Mexico and

Arizona. It is an area with high biodiversity in a convergence zone

of five biotic provinces with diverse topography, microclimates, and

habitats:tropicaltemperateSMO,temperateRockyMountains,Great

Plains-Chihuahuan Desert, Sonoran Desert, and the western Mexico

lowlands tropics. The transition between the New World tropics and

the northern temperate zone is at about 29°N-30°30’N in east-central

Sonora, in northwestern Mexico.

	 Here we present results on preliminary floristic inventories in the

tropical lowlands around the Sierra la Madera and compare themwith

the tropical vegetation in the Municipio de Yécora on the western

base of the Sierra Madre Occidental. The results help characterize

foothills thornscrub vegetation in Sonora.

Study Areas

	 The two study areas are in east-central Sonora in the Río Yaqui

drainage basin (fig. 1).

Sierra la Madera

	 The Sierra la Madera is a Sky Island mountain range located

(ca. 29°51’N to 30°10’N, 109°25’W to 109°42’W) between Óputo

(Villa Hidalgo) and Huásabas along the Río Bavispe to the east and

between Cumpas and Moctezuma along the Río Moctezuma to the

west (Yanes-Arvayo and others 2011). Portions of the range are in

the Municipios of Cumpas, Granados, Huásabas, Moctezuma, and

Villa Hidalgo. Elevationsrange from 615 m at Huásabas and 660 m at

Moctezuma to over 2,300 m on the highest peak, an elevational range

of about 1,685 meters. It is one of the higher ranges in Sonora. Most

of the area is privately owned; the higher elevations are in Fracción

V of the Área Natural ProtegidaAjos Bavispe. The vegetation ranges

from foothillsthornscrub (FTS) in the lowlandsto oak woodland and

pine-oak forest at higher elevations. There is cottonwood-willow

riparian forest along theRíosBavispe and Moctezuma.We follow the

classification used in Martin and others (1998) for vegetation names.

Yécora Area

The Municipio de Yécora is an area of 3,300 km2 on the Pacific

slope of the Sierra Madre Occidental eastern Sonora. Elevation in
USDA Forest Service Proceedings RMRS-P-67. 2013 241

Comparison of the Tropical Floras of the Sierra la Madera and the Sierra Madre . . . Van Devender and others

the Municipio rangesfrom 480 m near Curea to 2,140 m on Mesa del

Campanero, an increase of 1,660 meters in 27 km (by air). Mexican

Federal Highway 16 (MEX 16) is one of the few highways that

crosses the SMO, providing a west-east elevational transect from

foothills thornscrub near Curea (460-550 m) and tropical deciduous

forest (TDF, 500-1,160 m) up through oak woodland (1,050-1,700

m) to pine-oak forest (1,220-2,240 m). Grassland occurs in high

valleys (1,200-1,700 m) within oak woodland or pine-oak forest.

Mixed-conifer forest (1,900-2,100 m) is present in Barranca El Salto

on Mesa del Campanero.

Inventories

	 The Universidad de la Sierra (UNISIERRA) in Moctezuma has a

long-term project to document the flora of Sierra la Madera (YanesArvayo and others 2011). Only a few plants were collected in this area

before thefloristic effortssummarizedhere.TheLumholtzExpedition

to the Sierra Madre Occidental followed the Río Bavispe from Óputo

(Villa Hidalgo) to Huásabas and Granados in September-October

1890 (Lumholtz, 1902). The Swedish botanist Carl V. Hartman, on

the Expedition, collected plants along the eastern base of the Sierra

la Madera. Stephen S. White of the University of Michigan visited

the Moctezuma area in 1938 as part of his floristic study of the Río

Bavispe region in northeasternSonora (White, 1948).StephenS.Hale

made a few noteworthy collectionsin a survey oftheTarahumara frog

(Rana tarahumarae) in 1981 (Hale and May, 1983).

	 In August 2010, the Madrean Archipelago Biodiversity (MABA)

Project led an expedition to the Sierra la Madera. A group of 38 scientists, researchers, students, volunteers, and Comisión Nacional de

ÁreasNaturalesProtegidas(CONANP)madebiologicalobservations,

plant collections, and images at ranchos la Barchata, las Bateas, la

Cieneguita (and areas above), Mesa Quemada, and San Fernando.

In September 2011, Van Devender and Reina-Guerrero observed

the flora on the east slope of the Sierra la Madera west of Huásabas.

Melissa Valenzuela-Yánez (2012) as part of her undergraduate thesis

at UNISIERRA collected plants in the Ejido de Tonibabi area east of

Moctezuma on eight trips in April-December 2011. Plant collections

were deposited into the Universidad de Sonora (USON), University

of Arizona (ARIZ), and other herbaria. All plant observations and

collections, and many images are available online in the MABA

database (madrean.org).

Results

	 We documented 220 taxa in 57 families in FTS in the Sierra la

Madera. The families with the most species are Fabaceae (16.4%),

Asteraceae (11.4%), Poaceae (5.9%), and Cactaceae and Malvaceae

(4.5% each). The genera with the most species are Acacia, Euphorbia, and Ipomoea (6 taxa each), and Bouteloua (5 taxa). Only seven

species are non-native (3.2%): i.e., Cynodon dactylon, Marrubium 

vulgare, Melinis repens, Mollugo verticillata, Nasturtium officinale, 

Nicotiana glauca, andPennisetum ciliare.OnlyP. ciliare (buffelgrass)

isinvasive in FTS, although berro (N. officinale) can be very common

in permanent streams. The life forms of the Sierra la Madera FTS

flora are herbs (53.2%, including grasses and sedges), woody plants

(40.9%), and succulents (5.5%). The herbs (103 taxa) are nearly

equally perennial (54 taxa) and annual (49 taxa), with only 14 grass/

sedge taxa. It is clear that there needs to be additional documentation

of annuals and grasses. Although the larger woody plants and succulents are the structural dominantsin FTS, their life forms are not as

common as herbs: i.e., trees (10.5%), shrubs (15.9%), and subshrubs

(8.7%).

	 The floristic affinities of the Sierra la Madera FTS are mostly

tropical. Of the 49 species in the Sierra la Madera FTS that were

not in Yécora FTS or TDF, 43 are native, terrestrial species. Most of

them occur in the Sonoran Desert and/or desert grassland (30 each),

or are widespread from the Sonoran Desert through desert grasslands

to western Texas (16 species), with a total of 36 species (83.7%) in

this group. The affinities are less strong with the SMO/SI (7 taxa).

In the Yécora area, we documented 298 taxa in 61 families in the

FTS flora. The families with the most species are Fabaceae (11.7%),

Asteraceae and Poaceae (10.1% each), Euphorbiaceae (5.4%), and

Convolvulaceae (4.7%).Thegenerawiththemostspecies are Ipomoea

(10), Euphorbia (9), Physalis (6), and Cyperus and Muhlenbergia

(5 each). There are 24 non-native species (8.0%). The life forms of

the Yécora area thornscrub flora are herbs (68.8%), woody plants

(29.5%), and succulents (2.0%). Twice as many herbs are annuals

(111 taxa) as perennials (57 taxa). Annuals and grasses/sedges (37

taxa) are betterrepresented than in the Sierra laMadera.Trees(7.4%),

shrubs (11.4%), and subshrubs (7.0%) are the structural dominants. 

Discussion

	 The flora of FTS has not previously been studied. We identified

220 plant taxa in FTS in the Sierra la Madera and 298 taxa in the

Yécora (35% richer), with 433 taxa in both areas. Most FTS species

also occur inTDF in theYécora area (69.5% and 82.9% of the floras),

which, with 793 taxa, is much more diverse than FTS. But as pointed

out by Rzedowski (1978) for the Sierra Madre Occidental in general,

oak woodland (809) and pine-oak forest (1081) at higher elevations

on the Yécora gradient are more diverse than tropical lowland communities (Van Devender and Reina-Guerrero, unpublished).

	 Floristically, there are 83 taxa shared between the Sierra la Madera

and Yécora FTS floras (37.7% and 27.9%, respectively). The differences between FTS in the Sierra la Madera and the Yécora area are

mostly in herbs. The numbers of structurally dominant woody plants

Figure 1—Study areas. 
242 USDA Forest Service Proceedings RMRS-P-67. 2013

Van Devender and others Comparison of the Tropical Floras of the Sierra la Madera and the Sierra Madre . . .

in the two FTS floras are remarkably similar; i.e., trees (23, 24 taxa),

shrubs (35, 34 taxa), subshrubs (19, 21 taxa), and woody vines (12,

11 taxa). Important FTS trees are Acacia occidentalis, Bursera fagaroides, B. laxiflora, Fouquieria macdougalii, Ipomoea arborescens, 

Jatropha cordata, Lysiloma watsonii, and Parkinsonia praecox,

plus the shrubs Erythrina flabelliformis, Eysenhardtia orthocarpa,

and Mimosa distachya. Woody vines with 20 species are much more

diverse in TDF than FTS, and steadily decline to the west as aridity

increases in the Sonoran Desert.

Deep History of Thornscrub

	 The uplift of the Sierra Madre Occidental (SMO) in the late Oligocene-middle Miocene (25-15 million years ago, mya) had dramatic

impacts on the biota of northwestern Mexico (Van Devender, 2002).

Tropical forests, once found from coast-coast across the continent,

were restricted to the coastal lowlands. As the region gradually dried

out in the Miocene, tropical deciduousforest isolated in northwestern

Mexicowastransformedfirstintothornscrubandthenintodesertscrub,

as the Sonoran Desert formed about 8 million years ago (Axelrod,

1979). In the Pliocene (2.4-5 mya), there was a reversal to more

tropical climates, when thornscrub presumably expanded back into

desert areas.

	 The climatic changes during the Pleistocene with 15-20 glacial/

interglacial climatic cycles had dramatic impacts on the Sonoran

Desert as pinyon-juniper woodlands moved to low elevations in

southwestern Arizona and southeastern California (Van Devender,

1990). Ice age climates differed from modern ones in much cooler

summers, fewer winter freezes, reduced summer rains from tropical

oceans, andincreasedwinterprecipitation fromPacific frontalstorms.

For 80-90% of the last two million years, the Sonoran Desert was

greatly reduced in area, but thornscrub may have expanded as TDF

dried out to the south.

Thornscrub

	 There are two types of thornscrub in Sonora (Martin and others

1998). Coastal thornscrub is on the flat coastal plain of the Gulf of

California from Empalme-Guaymas southward (Friedman 1996).

Foothills thornscrub is on inland rocky slopes. In southern Sonora,

FTS is below TDF in elevation. In central Sonora, it is the transitional vegetation between the Plains of Sonora subdivision Sonoran

desertscrub on the west and oak woodland in Sky Island mountain

ranges and the Sierra Madre Occidental to the east. In the north,

FTS is replaced by desert grassland as winters become colder and

periodic fires become ecological processes. The northern limits of

FTS in Sonora are at about 30°11’N east of Sinoquipe in the Río

Sonora Valley and 30°26’N at PresaAngostura on the Río Bavispe at

the southern end of the Sierra El Tigre. FTS does not reach Arizona,

but the distributions of quite a few FTS species cross the border into

southern Arizona in desert grassland or oak woodland.

	 We conclude that foothills thornscrub is a distinct vegetation type

that evolvedinnorthwesternMexicoandisbestrepresentedinSonora,

where itiswidespread,important, andunder-appreciated. Ourfindings

suggest that the floristic composition of FTS variesin different areas,

but similarities in community structure (height, strata, and density)

and dominants, habitats, and climate make it readily recognizable.

Acknowledgments

	 Father William Trauba, Thomas F. Daniel, Patty West, and Kim

Baker helped with the fieldwork in the Yécora area. We thank Don

Pedro Avilez, Don Francisco Encinas, Doña Rosa, and Yaneth of

Curea for guidance and hospitality.We thank Sky IslandAlliance and

the Universidad de la Sierra for support of the MABA Expedition to

the Sierra de la Madera. We thank John A. Anderson for his careful

review of the paper.

References

Axelrod, Daniel I. 1979. Age and origin of the Sonoran Desert. California

Academy of Sciences. Occasional Papers, 132: 1-74.

Friedman, Samuel. L. 1996. Vegetation and flora of the coastal plains of the

Río Mayo region, southern Sonora, Mexico. M.S. Thesis. Tempe: Arizona

State University.

Hale, Stephen F.; and May, Clarence. J. 1983. Status Report for Rana tarahumarae Boulenger. Arizona Natural Heritage Program, Tucson. Report

to Office of Endangered Species, U.S. Fish and Wildlife Service, Albuquerque, NM. 99 p.

Lumholtz, Carl S. 1902. Unknown Mexico: A record of five years’ exploration among the tribes of the western Sierra Madre; in the Tierra Caliente

of Tepic and Jalisco; and among the Tarascos of Michoacán. New York:

Charles Scribner’s Sons.

Martin, Paul S.;Yetman, DavidA.; Fishbein, Mark;[and others]. (eds.). 1998.

Gentry’s Río Mayo Plants. The tropical deciduous forest and environs of

northwest Mexico. Tucson: University of Arizona Press. 558 p.

Rzedowski,J. 1978. Vegetación de México. México: Editorial Limusa. 432 p.

Van Devender, Thomas R. 1990. Late Quaternary vegetation and climate of

the Sonoran Desert, United States and Mexico. Pp. 134-165 in Betancourt,

Julio L.; Van Devender, Thomas R.; and Martin, Paul S. eds. Packrat

Middens. The Last 40,000 Years of Biotic Change. Tucson: University of

Arizona Press. 467 p.

Van Devender, Thomas R. 2002. Environmental history of the Sonoran

Desert. Pp. 3-24 in Fleming, Theodore H.; and Valiente-Banuet, Alfonso.

eds. Evolution, Ecology and Conservation of Columnar Cacti and their

Mutualists. Tucson: University of Arizona Press. 371 p.

Valenzuela-Yánez, Melissa. 2012. Flora vascular del Ejido Tonibabi; Moctezuma, Sonora. Memoria Professional para Licenciado en Biología,

Universidad de la Sierra, Moctezuma, Sonora. 62 p.

White, Stephen S. 1948. The vegetation of flora of the region of the Río de

Bavispe in northeastern Sonora, Mexico. Lloydia, 11: 229‐302.

Yanes-Arvayo, Gertrudis; Montañez-Armenta, María de la Paz; Silva-Kurumiya, Hugo; Gil-Montaño, Enrique;Van Devender,ThomasR.; andReinaGuererro,Ana L. 2011.Catálogo de Plantas de Sierra La Madera.Arbóreas,

Arbustivas y Herbáceas. Universidad de la Sierra, Hermosillo, Sonora.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 243

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Biogeographic Perspective of Speciation Among Desert 

Tortoises in the Genus Gopherus: A Preliminary Evaluation

Taylor Edwards

University of Arizona Genetics Core, University of Arizona, Tucson, Arizona

Mercy Vaughn

Paso Robles, California

Cristina Meléndez Torres

Comisión de Ecología y Desarrollo Sustentable del Estado de Sonora, Sonora, Mexico

Alice E. Karl

Alice E. Karl and Associates, Davis, California

Philip C. Rosen

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Kristin H. Berry

U.S. Geological Survey, Western Ecological Research Center, Riverside, California

Robert W. Murph

Royal Ontario Museum, Toronto, Canada

Abstract—The enduring processes of time, climate, and adaptation have sculpted the distribution of organisms 

we observe in the Sonoran Desert. One such organism is Morafka’s desert tortoise, Gopherus morafkai. We 

apply a genomic approach to identify the evolutionary processes driving diversity in this species and present 

preliminary findings and emerging hypotheses. The Sonoran Desert form of the tortoise exhibits a continuum 

of genetic similarity spanning 850 km of Sonoran desertscrub extending from Empalme, Sonora, to Kingman, Arizona. However, at the ecotone between desertscrub and foothills thornscrub we identify a distinct, 

Sinaloan lineage and this occurrence suggests a more complex evolutionary history for G. morafkai. By using 

multiple loci from throughout the tortoise’s genome, we aim to determine if divergence between these lineages 

occurred in allopatry, and further to investigate for signatures of past or current genetic introgression. This 

international, collaborative project will assist state and federal agencies in developing management strategies 

that best preserve the evolutionary potential of Morafka’s desert tortoise. Ultimately, an understanding of 

the evolutionary history of desert tortoises will not only clarify the forces that have driven the divergence in 

this group, but also contribute to our knowledge of the biogeographic history of the Southwestern deserts 

and how diversity is maintained within them.

Introduction

	 The diverse assemblages of flora and fauna in the Sonoran Desert

have been sculpted by the enduring processes of time, climate, and

adaptation. The Sonoran Desert developed gradually over the last 8

million yearsthrough multiple glacial and interglacial climatic cycles

that resulted in repeatedly expanding and shrinking woodlands,

desertscrub, and thornscrub communities (Van Devender 1990). The

modern characteristic assemblage that includes columnar cactus and

palo verde trees that define Arizona Upland Sonoran desertscrub

likely did not (re)emerge until roughly 9,000 years ago (Van Devender 1990). We are interested in how this fluctuating vegetation

community has shaped the evolution of one of the Sonoran Desert’s

most charismatic reptiles, the Morafka’s desert tortoise, Gopherus 

morafkai. In particular, we explore how genetic variation within this

species fits the biogeographic history of the landscape.

	 The genus Gopherus contains of two species of desert tortoise, one

native to the Mojave and Colorado deserts(Agassiz’s desert tortoise;

G. agassizii) and another in the Sonoran Desert region (Morafka’s

desert tortoise, G. morafkai; Murphy and others 2011). Gopherus 

agassizii occurs almost entirely north and west of the Colorado River

andG. morafkai inhabitsregionssouth and east oftheColoradoRiver,

extending southward into tropical deciduous forests of Southern

Sonora, Northern Sinaloa, and extreme Southwestern Chihuahua.

The Mojave and Sonoran lineages of Gopherus differ in morphology,

seasonal activity,reproductive ecology,habitatselection, andgenetics
244 USDA Forest Service Proceedings RMRS-P-67. 2013

Edwards and others Biogeographic Perspective of Speciation Among Desert Tortoises in the Genus Gopherus: . . .

(e.g., Lamb and others 1989; McLuckie and others. 1999; Berry and

others 2002).

	 The U.S. Fish and Wildlife Service federally listed the Mojave

population of the desert tortoise (G. agassizii) as threatened (USFWS 1990). The Sonoran population (G. morafkai) is not federally

listed in the United States, but became a candidate for federal listing

in 2010 and is considered a Species of Special Concern in Arizona

(USFWS2010).MexicanpopulationsofGopherusreceiveprotection

asthreatenedspecies(CategoryA“Amenazada” inNOM-059;D.O.F.

2002). To assesstrendsin Morafka’s desert tortoise, it is necessary to

consider populations in Mexico that occupy about two-thirds of the

geographic range. Unfortunately, little data exist on the distribution,

ecology, and taxonomic status of the tortoise in Mexico. Driven by

this lack of knowledge, in 2005 a team of Mexican, Canadian, and

American collaborators began amultifaceted study of desert tortoises

throughout their range in Mexico focusing on the tortoises’ health,

genetics, general biology, and ecology to investigate their status and

assess conservation needs. Here we present preliminary findings and

our emerging hypotheses.

	 Across its large geographic range in Mexico, Morafka’s desert

tortoise occupies a variety of habitatsincluding desertscrub, foothills

thornscrub, andthenorthernpartofsemiaridSinaloantropicaldeciduous forest (TDF) of southern Sonora and northern Sinaloa (Fritts and

Jennings 1994). Behavioral patterns and ecological challenges for

tortoisesin these different environments can vary greatly, but there is

currently very little published on the behavior or ecology of tortoises

inthornscrubandTDFcommunities.Genetically,mitochondrialDNA

(mtDNA)sampled from tortoisesin the southern portion of the range

is deeply divergent from haplotypes in the central part of the species’range, suggesting the potential for a unique lineage (Lamb and

others 1989; Berry and others 2002). Thus, one challenge currently

facing conservation initiatives for G. morafkai is to determine the

taxonomic status and distribution in the southern portion of itsrange.

The “Sinaloan” type tortoise may be a cryptic species (Murphy and

others 2011).

	 Taxonomic assessments of tortoisesin Mexico can be bolstered by

identifying geographic barriers to dispersal that contribute to genetic

isolation, analogous to the Colorado River, which separates G. agassizii and G. morafkai. In contrast, the identification of genetic clines

infers that taxonomic recognition of multiple species is unwarranted

because such designations would serve to recognize the extremes

only. It is equally critical to understand the rates and patterns of

genetic differentiation expected between two populations. Previous

work has established that in both Mojave and Sonoran environments,

desert tortoises exhibit a population genetic structure of isolation by

distance (IBD: Edwards and others 2004; Murphy and others 2007;

Hagerty and others 2011). IBD occurs when the geographic distance

between populations is the major limitation to gene flow. Panmixia

is expected at all sampling localities yet allele frequencies at sites

separated by great geographic distances may differ significantly. In

contrast, when alleles that distinguish species boundaries are under

strong selection, the rate of genetic introgression should be near zero

in one or both directions (Payseur 2010).

Methods

	 Between 2005 and 2011 we collected blood samplesfrom over 130

wild-caught G. morafkai throughout the species’range in Mexico and

in three different vegetation communities: TDF, foothillsthornscrub,

and Sonoran desertscrub. Targeted sampling involved the ecotone

between foothills thornscrub and Sonoran desertscrub (fig. 1). We

collected whole blood via subcarapacial or brachial venipuncture.We

also used salvaged red blood cellsstored at the University ofArizona

Genetics Core. Blood collection followed Animal Use Protocols approved by the Royal Ontario Museum Animal Care Committee, and

by theUniversity ofArizona (IACUC09-138).All genetic procedures

and analyses were conducted in the University of Arizona Genetics

Core following procedures described in Edwards and others (2004)

and Murphy and others (2007). We genotyped individuals for 16

microsatellite loci (Edwards and others 2004; Murphy and others.

2007) and sequenced approximately 1,200 base pair portions of

mtDNA(Murphy and others 2007).In addition,we utilized additional

samples from throughout the range of desert tortoises in the United

States(Edwards and others 2004; Murphy and others 2007; Edwards

and others 2010).

	 We used PAUP* 4.0b10 (Swofford 2002) to reconstruct the maternal (mtDNA) phylogeny using a 50% majority rule consensustree

based on unweighted maximum parsimony analysis. In addition to

the samples collected in this study, we also included data from G. 

morafkai collected inArizona,G. agassizii collected inCalifornia and

Utah (Murphy et al. 2007), and we utilized other Gopherus spp. (G. 

berlandieri, G. flavomarginatus and G. polyphemus) as outgroups.

Figure 1—Sample locations in Mexico for this study are represented by 

circles; solid circles represent sites where the sampled mitochondrial 

haplogroup is strictly “Sonoran,” gray-shaded circles represent sites where 

the sampled mitochondrial haplogroup is strictly “Sinaloan” and open 

circles represent sites where a mix of Sonoran and Sinaloan haplotypes 

are observed. The double-dashed vertical line running down the middle 

of the image roughly defines the boundary of Sonoran desertscrub.
USDA Forest Service Proceedings RMRS-P-67. 2013 245

Biogeographic Perspective of Speciation Among Desert Tortoises in the Genus Gopherus: . . . Edwards and others

We performed a bootstrap analysiswith 1,000 replicates of 10 random

addition sequences per replicate to determine the support for the inferred relationships. For our microsatellite analyses we incorporated

samples previously collected in Arizona and estimated the degree

of population differentiation within and among populations using

AMOVA (analysis of molecular variance) in ARLEQUIN 3.5.1.2

(Excoffier and others 2005). We calculated genetic distances based

on pairwise ΦST (FST sensu Weir and Cockerham 1984) and defined

populations by their proximity to major cities; Kingman, Phoenix,

and Tucson, Arizona, USA, and Hermosillo, Ciudad Obregón, and

Álamos, Sonora, Mexico.

Results

	 Analyses of mtDNA sequences resolved three distinct maternal

lineages corresponding to the Mojave, Sonoran, and Sinaloan

populations. Sonoran and Sinaloan tortoises differentiated along the

ecotone between foothills thornscrub and Sonoran desertscrub. Distinct Sinaloan haplotypes occurred only in foothills thornscrub and

TDF environments while Sonoran haplotypes were mostly restricted

to desertscrub environments (fig. 1). Several sample sites along the

ecotone of foothillsthornscrub and desertscrub exhibited individuals

representing both Sonoran and Sinaloan maternal lineages (fig. 1).

Modal Sonoran haplotype SON01 (Murphy et al. 2007) was observed

at all sites exhibiting Sonoran lineages.

	 In the analyses of bi-parentally inherited microsatellite loci we

observed a continuum of genetic similarity spanning 850 km of Sonoran desertscrub, ranging from Hermosillo, Sonora, northwards to

Kingman,Arizona (table1).This correspondedwiththedistributionof

Sonoran mtDNAhaplotype SON01. Tortoises within this continuum

had a population structure consistent with IBD (Mantel test:r

2 = 0.75,

p<0.01). However, strength of the correlation began to break down

when assessed acrossthe ecotonal boundary between desertscrub and

thornscrub-TDF (Mantel test: r2 = 0.38, p<0.01). Fixed differences

in microsatellite alleles occurred in foothills thornscrub and TDF in

southern Sonora and clearly distinguish a unique Sinaloan genotype.

Discussion

	 These analyses suggest that tortoises living in foothills thornscrub

and TDF in southern Sonora and Sinaloa have a level of differentiation equal to that between G. agassizii and G. morafkai fromArizona

(Murphy and others 2011). All three lineages—Mojave, Sonoran,

and Sinaloan—likely diverged from a common ancestor 5–6 million

years ago (fig. 2; Lamb and others 1989; Avise and others 1992;

McLuckie and others 1999). A cursory analysis of the three mtDNA

lineages suggests they are on independent evolutionary trajectories

and potentially constitute separate species. However, spatial overlap

of several haplotypes occurs at the eastern and southern boundaries

Table 1—Comparison of pairwise genetic distance, ΦST (FST sensu Weir and Cockerham 1984), 

among sites spanning the geographic range of desert tortoises in the Sonoran Desert. 

Data generated from 16 microsatellite loci. Sites are categorized by their proximity to major 

cities; Kingman, Phoenix, and Tucson, Arizona, USA, and Hermosillo, Ciudad Obregón, 

and Álamos, Sonora, Mexico. Dotted line indicates transition from localities representing 

desertscrub (above) and thornscrub and tropical deciduous forest (below). This analysis 

was performed excluding samples from the 4 sites with mixed mitochondrial lineages.

Kingman Phoenix Picacho Tucson Hermosillo Obregón 

Kingman 

Phoenix 0.007 

Picacho 0.043 0.024 

Tucson 0.067 0.042 0.008

Hermosillo 0.090 0.055 0.051 0.041

Obregón 0.251 0.201 0.185 0.181 0.145

Álamos 0.231 0.191 0.173 0.172 0.153 0.072

Figure 2—A 50% majority rule consensus tree based on maximum 

parsimony for 1,200 base pairs of sequence from the ND4 region 

of the of mitochondrial genome of desert tortoises (Gopherus 

spp.); SON = G. morafkai, SON mex = mtDNA haplotypes 

observed only in Sonora, Mexico, MOJ = G. agassizii, MEX = 

Sinaloan populations of Gopherus morafkai and outgroups G. 

berl (G. berlandieri), G. flav (G. flavomarginatus), and G. poly 

(G. polyphemus). Numbers above branches are bootstrap values.
246 USDA Forest Service Proceedings RMRS-P-67. 2013

Edwards and others Biogeographic Perspective of Speciation Among Desert Tortoises in the Genus Gopherus: . . .

of Sonoran desertscrub and this suggests a complex history in this

vegetative transition zone (fig. 1).The transitionsfrom desertscrub to

thornscrub toTDF are generally considered gradual, clinal transitions

between somewhat artificial categories. Even though about 85% of

foothills thornscrub is in Sonora, phytogeographers have characterized it as “Sinaloan” as a result of floristic affinities to TDF (Brown

1982). This ecotone is likely to have expanded and retreated many

times during the relatively recent transformation into the Sonoran

Desert (Van Devender 2000) and this dynamic system has undoubtedly influenced genetic differentiation among deserttortoise lineages.

	 Whereas the divergence between G. agassizii and G. morafkai is

a classic example of allopatric speciation resulting from geographic

isolation by the Colorado River (Lamb and others 1989; Avise and

others 1992; McLuckie and others 1999; Murphy and others 2011),

divergenceofSonoranandSinaloanG. morafkaisuggeststhat adifferentmechanismisin play. Sonoran and Sinaloan tortoises differentiate

along an ecotone of foothills thornscrub and Sonoran desertscrub

communities; ecotones can play critical roles in evolution because

hybridization can occur in these settings (Barton and Hewitt 1985).

Interestingly, our analyses do not detect clear signals of hybridization between Sinaloan and Sonoran type tortoises despite the spatial

proximity oflineagesin this extensive vegetative transition zone (fig. 1).

This is in contrast to G. agassizii and G. morafkai where hybridization has been observed at locations of secondary contact (Edwards

and others 2010). Unlike the clear allopatric pattern of divergence

between G. agassizii and G. morafkai, Sonoran-Sinaloan lineages of

G. morafkai may have diverged under a parapatric model where the

current contact zone in Mexico may be a result of recent secondary

contact after isolation prior to the formation of the Sonoran Desert.

Chance interbreeding (during times of sympatry) may have driven

or be driving the development of reproductive isolating mechanisms.

Alternatively, the sharp break in the cline may be a result of insufficient time for genes to flow far beyond the boundary. Our future

efforts will attempt to discern whether genetic differentiation is a

result of past periods of isolation, or if differentiation has evolved in

the absence of physical barriers to gene flow.

	 Taxonomic clarification is of upmost importance for management

of G. morafkai. Our continuing effort to resolve the taxonomy of the

species in the context of its geographic distribution will provide a

basisfor prioritizing resources and conservationmeasureswhere they

are most needed and will ensure that data are available to maintain

genetic diversity acrossthe species’range. Future efforts will resolve

the taxonomic uncertainty of G. morafkai in Mexico by testing the

null hypothesis of conspecificity and determining the mechanism(s)

of divergence between the Sonoran and Sinaloan lineages. Clarification of these evolutionary processes will directly inform efforts to

preserve the evolutionary potential of the species. Ultimately, an

understanding of the evolutionary history of desert tortoises will not

only clarify the forces that have driven differentiation in this group,

but also contribute to our knowledge of the biogeographic history of

the Southwest deserts and how diversity is maintained within them.

Acknowledgments

	 We obtained Mexican samplesthrough a multinational, collaborative effort including CEDES and F. R. Méndez de la Cruz, Instituto

de Biología, Universidad Nacional Autónoma de México (UNAM).

Special thanks go to the dedicated field crew that volunteered to make

this work possible. Permits for Mexican samples were facilitated by

Secretaría de MedioAmbiente y Recursos Naturales(SEMARNAT).

TheTucsonHerpetological Society,DesertTortoiseCouncil,Arizona

Game and Fish Department, University of Florida, University ofArizona GeneticsCore, U.S. Geological Survey,Royal Ontario Museum

Foundation, Natural Sciences and Engineering Research Council of

Canada Discovery Grant 3148 and private donors provided funding and support for sample collection in Mexico. Any use of trade,

product, or firm names is for descriptive purposes only and does not

imply endorsement by the U.S. Government.

References

Avise,J. C., B. W. Bowen, T. Lamb,A. B. Meylan, and E. Bermingham. 1992.

Mitochondrial DNA evolution at a turtle’s pace: evidence for low genetic

variability and reducedmicroevolutionary rate in theTestudines.Molecular

Biology and Evolution 9:457−473.

Barton, N. H., and G. M. Hewitt. 1989. Adaptation, speciation, and hybrid

zones. Nature 341:497−503.

Berry, K.H., D. J. Morafka, and R. W. Murphy. 2002. Defining the desert

tortoise(s): our first priority for a coherent conservation strategy.Chelonian

Conservation and Biology, 4:249−262.

Brown, D. E. (ed.). 1982. Biotic communities of the American Southwest–

United States and Mexico. Desert Plants 4:1−342.

D.O.F. 2002. NOM-059-ECOL-2001. Protección ambiental- especies nativas

de México de flora y fauna silvestres - Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio - Lista de especies en riesgo.

Diario Oficial de la Federación, 6 March 2002.

Edwards, T., C. J. Jarchow, C. A. Jones, and K. E. Bonine. 2010. Tracing

genetic lineages of captive desert tortoises in Arizona. Journal of Wildlife

Management 74:801−807.

Edwards, T., C. R. Schwalbe, D. E. Swann and C. S. Goldberg. 2004. Implications of anthropogenic landscape change on inter-populationmovements of

thedeserttortoise (Gopherus agassizii).ConservationGenetics5:485−499.

Excoffier,L.G.Laval, and S. Schneider. 2005.Arlequin ver. 3.0:An integrated

software package for population genetics data analysis. Evolutionary Bioinformatics Online 1:47−50.

Fritts, T. H., and R. D. Jennings. 1994. Distribution, habitat use, and status

of the desert tortoise in Mexico. Pages 49−56 in Bury, R.B., and D.J.

Germano, eds. Biology of North American Tortoises. Fish and Wildlife

Research 13. Washington, DC: U.S. Department of the Interior, National

Biological Survey, 204pp.

Hagerty B. E., K. E. Nussear, T. C. Esque, and C. R. Tracy. 2011. Making

molehills out of mountain landscape genetics of the Mojave desert tortoise.

Landscape Ecology 2: 267–280.

Lamb, T, J. C. Avise, and J. W. Gibbons. 1989. Phylogeographic patterns

in mitochondrial DNA of the desert tortoise (Xerobates agassizi), and

evolutionary relationships among the North American gopher tortoises.

Evolution 43:76−87.

McLuckie,A. M.,T. Lamb,C.R. Schwalbe, andR. D. McCord. 1999. Genetic

and morphometric assessment of an unusual tortoise (Gopherus agassizii)

population in the Black Mountains of Arizona. Journal of Herpetology

33: 36−44.

Murphy, R.W., K. H. Berry, T. Edwards andA. M. McLuckie. 2007.Agenetic

assessment ofthe recovery unitsfortheMojave Population ofthe desert tortoise,Gopherus agassizii.ChelonianConservationandBiology6:229−251.

Murphy, R. W., K. H. Berry, T. Edwards, [and others]. 2011. The dazed and

confusedidentityofAgassiz’slandtortoise,Gopherusagassizii (Testudines:

Testudinidae) with the description of a new species, and its consequences

for conservation. ZooKeys 113:39−71.

Payseur,B.A.2010.Usingdifferentialintrogression inhybridzonestoidentify

genomic regions involved in speciation. Molecular Ecology 10:806−820.

Swofford, D. L. 2002. Phylogenetic Analysis Using Parsimony. Sunderland,

MA: Sinauer Associates Inc, software.

U.S. Fish and Wildlife Service [USFWS]. 1990. Endangered and threatened

wildlife andplants;determinationofthreatenedstatusoftheMojavepopulation of the desert tortoise. Federal Register 55:12178−12197.
USDA Forest Service Proceedings RMRS-P-67. 2013 247

Biogeographic Perspective of Speciation Among Desert Tortoises in the Genus Gopherus: . . . Edwards and others

U.S. Fish and Wildlife Service [USFWS]. 2010. Endangered and threatened 

wildlife and plants; 12-month finding on a petition to list the Sonoran 

population of the desert tortoise as endangered or threatened; proposed rule. 

Federal Register 75(293): 78094−78146. December 14, 2010.

Van Devender, T.R. 1990. Late Quaternary vegetation and climate of the 

Sonoran Desert, United States and Mexico. Pp. 134–65 in Betancourt, 

J.L., T. R. Van Devender, and P. S. Martin, eds. Packrat Middens: The last 

40,000 years of biotic change. Tucson, AZ: University of Arizona Press.

Van Devender, T. R. 2000. The deep history of the Sonoran Desert, pp. 61 −69 

in Phillips, S. J. and P. W. Comus, eds. A natural history of the Sonoran 

Desert. University of California Press, Berkeley, CA.

Weir, B. S., and C. C. Cockerham. 1984. Estimating F-statistics for the analysis 

of population structure. Evolution 38:1358–1370. 

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
248 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Richness of Mammals on the San Bernardino Ranch in the 

Municipality of Agua Prieta, Sonora, Mexico

Mario Erandi Bonillas-Monge and Carlos Manuel Valdez-Coronel

	 Departamento de Investigaciones Científicas y Tecnológicas de la Universidad de Sonora, Sonora, Mexico

Abstract—Historically, the San Bernardino Ranch has performed, as economical activities, livestock and farming, which has contributed to the deterioration of regional ecosystems. The ranch is ecologically important 

due to the diverse types of habitats of conservation interest such as the semi-desert grassland, the riparian 

vegetation, and a large ciénega, in which restoration efforts are being applied through water harvesting by 

gabions. Mammals are essential in the ecosystem. Knowing about the actual species that live on the different sites of the ranch tells us about the healthy state of the environments. We used the direct and indirect 

sampling methods to identify 26 species of terrestrial mammals on the Ranch. Using this information, we 

compared the richness and diversity of the ranch species with the species living in the San Bernardino 

National Wildlife Refuge located next to the border, which was a helpful comparison since the Refuge has 

more years of restoration experience than does the San Bernardino Ranch.

Introduction

Due to the biogeographic location, Sonora is in a transition zone 

between the Nearctic and Neotropical region, and is considered an 

arid region because its large surface is covered by desert shrubs; 

however, there are deciduous forests in the southern part of the state 

and pine-oak woodlands in the Sierra Madre Occidental, providing 

a diversity of ecosystems (Molina-Freaner and Van Devender 2009). 

This diversity is a reflection of the mammals in the state including 

the presence of 126 identified species and 35 as possible occurrences 

(Castillo and others 2010). Mammals are a fundamental part in an 

ecosystem covering a huge variety of niches. Some species, like 

carnivors, are indicators of the healthy state of the ecosystem and are 

key in the maintenance of an ecological balance; they also provide 

data for diagnosing conservation 

Several areas of Sonora are considered of special interest because 

their resources are important to the population, as is the case of 

the San Bernardino Ranch, located at the Northeastern border with 

the United States inside the priority land region (RTP) number 45 

denominated by San Luis-Janos. This area contains the headwaters 

of the Rio Yaqui River and is important for the diverse ecosystems 

of grasslands and mountains (Arriaga and others 2000) consisting 

of a series of tributary rivers and streams (Hudson and others 2005) 

with vegetation of pine-oak woodlands, low open forest, chaparral, 

thornless shrub, grasslands, mesquites and chollal (Rzedowski 1978).

According to the classification of provinces by Caire (1978), there 

are five mammal provinces in Sonora with the Mapími province covering the San Bernardino area having the highest diversity of rodents 

in Sonora. There was a big cienega about 7000 years ago where the 

San Bernardino Ranch is presently located. But with the arrival of the 

Europeans, the Cienega became a field supporting agricultural crops 

and livestock grazing, causing the deterioration of the habitat. This 

area now is part of southeast Arizona and northeast Sonora (Minckley 

and others 2006).

According to Castillo and others (2010), rivers in the region function like tropical fauna corridors to the north, while the foothills 

of Sierra Madre Occidental function as corridors as well, allowing 

interchange of species between the north moist temperate and south 

tropical regions. Such is the case of Neotropical species of great interest like jaguar (Panthera onca), ocelot (Leopardus pardalis) and 

jaguarundi (Puma yagouaroundi), which historically had their north 

limit in southern Arizona (Grigione and others 2007). Herein is the 

importance of conserving this area.

The main concerns for conservation on the San Bernardino Ranch 

is the Agua Prieta-Janos road, which is the international border with 

United States of America, with a history of residual degradation 

(CEDES 2011). The first efforts for conserving the general area were 

undertaken in 1982, when the U.S. Fish and Wildlife Service declared 

the area in Arizona as San Bernardino National Wildlife Refuge for 

the protection of water resources and to provide a habitat for native 

species. This area has a register of 315 bird species, 55 amphibians 

and reptiles, and 66 species of mammals and more than 490 types of 

plants according to the U.S. Fish and Wildlife Service (http://www.

fws.gov/refuges/profiles/index.cfm?id=22523).

In Sonora, the conservation efforts in the area are carried out by 

the Association “Cuenca Los Ojos A C,” who work to restore the 

eroded soil, recharge groundwater aquifers, and promote original 

vegetation growth. However, these efforts are carried out only half 

of the time as those in the San Bernardino NWR so we do not know 

whether the health of the ecosystem is similar on both sides of the 

border. A listing of the small non-flying mammal species of the San 

Bernardino Ranch will help us to know more about the health of the 

ecosystem. Obtaining such a listing was the purpose of this study. 

One of the principal motives that led us to undertake this work was 

the demand for biological inventories of the fauna resources of the 

state and particularly about mammals. Also, whether similar mammal 
USDA Forest Service Proceedings RMRS-P-67. 2013 249

Richness of Mammals on the San Bernardino Ranch . . . Bonillas-Monge and Valdez-Coronel

species occur on the San Bernardino Ranch and the San Bernardino 

NWR may help to determine better ways of environmental management. Therefore, determining the mammal species of this Ranch will 

provide interesting information about the restoration efforts that are 

taking place nowadays, and will help us know how these efforts are 

improving the state of the ecosystem in this important area for wildlife.

Methodology

The study was made on the San Bernardino Ranch, located in the 

northeast corner of the Sonora state in Mexico, 30 km from Agua 

Prieta within the coordinates of 31°19’ 0.20” N y 109° 15’ 59.91” 

W. This study was conducted as a part of our professional practices 

during the summer of 2011 from June 27 to July. We later came back 

in winter 2012 from March 15-18 to expand our database.

We used two types of methods: direct sampling and non-direct sampling. With the direct method, we captured small- and medium-sized 

mammals and we were able to obtain information on their weight, 

sex, reproductive state, body measurements, and other characteristics 

that helped us understand the biology of the studied species. With 

the non-direct method we collected information about the species 

without direct measurements by identifying mammals by associated 

tracks like pad prints, scats, body parts, and nests.

Direct Sampling Methods

To know the specific richness of small (non-flying) mammals of the 

San Bernardino Ranch, we established three study sites: one located 

near a dam (Site 1), another localized in Silver Creek (Site 2) and 

a third in the Cienega (Site 3). The criteria for selecting these sites 

were the closeness to water and being able to place the traps on a 

gradient from more to less humidity. The coordinates were taken at 

the beginning and ending of the each studied transect on each site.

The method of capture and recapture was used, which can be used 

for measuring the population structure (Krebs 1985). To capture small 

mammals, 10 Sherman traps were placed on four lines on each site 

and were separated by 10 m from one another, totaling a total of 40 

traps for each site. Every trap was baited with a mixture of peanut 

butter and seeds, like oats, on 3 nights at each site. Every night the 

traps were baited and activated. In the morning (5 AM), the traps 

were checked for captures. Each collected individual was marked 

with purple ink and measured from total body, tail, ear, rear foot, 

and weight. Identification was determined through field guides. The 

individual was then release on the site of the capture.

For medium mammals, eight Tomahawk traps were placed around 

each site. These traps were baited with sardine, tuna, egg, and fresh 

vegetables. For each individual captured, the guides of Whitaker 

(1996) and Reid (2006) were consulted for their identification. All 

captured animals were released back into their area of capture after 

identification.

Non-Direct Sampling

Non-direct sampling was used to know the presence of some medium 

and large mammals. For this purpose we made walks on the main 

paths of the Ranch looking for tracks, footprints, scats, remains, and 

other marks. When these indicators were found, we used a scale and 

a camera with GPS to take pictures and obtain coordinates of each 

associated trace. The field guides of Aranda (2000) and Elbronch 

(2003) were used for identification of the traces. There were also 

five interviews with people of the Ranch (supported by illustrations) 

between the ages of 16 and 61 years to provide an idea of the mammals that we might find.

Results and Discussion

The total registration of mammals obtained in the study was 26 

species in the two sampling seasons of the year. This information was 

distributed in 4 orders, 11 families and 17 genera. The order with higher 

number of species was Rodentia with 14 (53.84% of total results) and 

seconded by the Carnivora order with 7 species representing 26.92%, 

a listing of which is presented in table 1. The 26 species register for 

the San Bernardino Ranch represent 20.63% of the total species of 

Sonora. This highlights the importance of conserving the mammals 

on the ranch.

Through the direct methods for small (non-flying) mammals, the total 

presence registered was 89 individuals, while there were 8 skunks 

(Mephitis mephitis) captured in the Tomahawk traps as summarized 

in table 2. The results of the non-direct methods of recoding mediumlarge mammals were the 43 proved observations shown in table 3, 

giving us a total of 140 proved data for the San Bernardino Ranch, 

which are included in the MABA data base.

Within the observed species, there was Desert shrew (Notiosorex 

crawfordi), the only species in the category of special protection (Pr) 

according to NOM-059-SEMARNAT-2010. We found in the Red List 

of IUCN that all registered species are under the category of least 

concern, but the populations of Mountain Lion (Puma concolor) and 

hog-nose skunk are under consideration for listing.

Through interviewing the people of the Ranch, we identified species such as Ringtale (Bassariscus astutus), Western spotted skunk 

(Spilogale gracilis), Cotton-tailed Rabbit (Sylvilagus audubonii), 

Pocket gopher (Thomomys bottae) and Badger (Taxidea taxus), but 

they were not included in the listing because of the lack of confirmation of their presence. We collected three skulls of dead animals that 

were taken to the Mammals Collection in the laboratory of terrestrial 

resources of the University of Sonora; further results are presented 

by the study sites in the following paragraphs.

Site 1

On this site we found the greatest amount of available water. This 

site is dominated by bulrush (Scirpus americanus) and was the site 

with more sampling success in the summer. Sigmodon arizonae was 

the species with more observations; this can be due to this species 

being associated with dense grasses to feed and make their burrows 

(Wilson and Ruff 1999). This was the only site where we found the 

desert shrew (N. crawfortii) in an area dominated by scrub. The site 

was relocated in the winter due to the increased amount of water. In 

this season we had more observations including two species, Neotoma 

albigula and N. Mexicana, which we did not have in the first phase 

of the study.

Site 2

We only had 2 individuals of Chaetodipus penicillatus and three 

skunks (M. mephitis) in the Tomahawk traps in the summer; this small 

number of captures might be because of the conditions of the site. 

Most of the area had bare ground, which does not provide adequate 

shelter for rodents. This number changed drastically in the winter, 

with observations of 28 individuals and 7 different species, resulting 

in the most captured individuals and with a major number of species 

at this site. This finding makes us think that the relation between the 
250 USDA Forest Service Proceedings RMRS-P-67. 2013

Bonillas-Monge and Valdez-Coronel Richness of Mammals on the San Bernardino Ranch . . .

Table 1—List of the mammals found in the San Bernardino Ranch.

Small non-flying mammals IUCNa Medium and large mammals IUCNa

population population

RODENTIA LAGOMORPHA 

Familia Cricetidae Familia Leporidae

Sigmodon arizonae Unknown Lepus californicus Stable

Sigmodon hispidus Increasing CARNIVORA

Reithrodontomys megalotisb Stable Familia Canidae

 Stable Canis latrans Increasing

Reithrodontomys montanusc Stable Urocyon cinereoargenteusb Stable

Peromyscus maniculatus Stable Familia Felidae

Peromyscus leucopus Stable Lynx rufusb Stable

Peromyscus boyliic Stable Puma concolorb Decreasing

Peromyscus eremicus Stable Familia Mephitidae

Baiomys taylori Stable Conepatus leuconotusb Decreasing

Familia Muridae Mephitis mephitis Stable

Neotoma albigulac Stable Familia Procyonidae

Neotoma mexicanac Stable Procyon lotor Increasing

Familia Heteromyidae ARTIODACTYLA

Chaetodipus penicillatus Stable Familia Tyassuidae

Chaetodipus intermedius Stable Pecari tajacu Stable

SORICOMORPHA Familia Cervidae

Familia Sorcidae Odocoileus hemionusb Stable

Notiosorex crawfordib Stable Odocoileus virginianus Stable

a

Red list of threatened species by the International Union for Conservation of Nature.

b

Species found only in Summer.

c

Species found only in Winter.

Table 2—Total records of small non-flying mammals.

Records of species

Species Summer Winter

Sigmodon arizonae 10 3

Sigmodon hispidus 3 5

Reithrodontomys megalotis 1 0

Reithrodontomys fulvencens 3 0

Reithrodontomys montanus 0 1

Peromyscus maniculatus 5 9

Peromyscus leucops 1 19

Peromyscus boylii 0 2

Peromyscus eremicus 0 13

Notiosorex crawfordi 1 0

Chaetodispus penisillatus 2 2

Chaetodispus intermedius 2 2

Bayomis tayloris 1 1

Neotoma albigula 0 2

Neotoma mexicana 0 1

Total records 89

Table 3—Total records of medium and large mammals.

Record of species

Species Summer Winter

Puma concolor 7 0

Procyon lotor 7 1

Conepatus leuconotus 1 0

Mephitis mephitis 5 5

Pecari tajacu 3 2

Odocoileus hemionus 2 0

Urocyon cinereoargenteus 1 0

Lepus californicus 2 0

Canis latrans 2 2

Lynx rufus 11 0

Total records 51

presence of water and the number of individuals was due to the dates 

of water flow in Silver Creek and a high density of vegetation giving 

a good habitat for rodents.

Site 3

On this site we expected to find the species of S. ochrognathus

that has been recorded in the San Bernardino NWR but not found 

on this side of the border. However, we found the only individual of 

the species Baiomys taylori. The number of species and individuals 

increased in the winter and it was the only place where we found an 

individual of the species Reithrodontomys montanus.

Associated Traces

The bobcat (Lynx rufus), followed by the raccoon (Procyon lotor), 

were the species with more traces that had many footprints over the 

paths, mainly on the shores of the rivers and also it was possible to 

observe some latrines on fallen trees and over gabions with water, 

which can indicate the adoption for these places with water (Urban 

1970). We also observed raccoon scats, a variety of insects, little 

mammals, and some molluscs.

We did not find many skunk scats (M. mephitis), probably due to the 

aleatory selection of defecation or to hidden places like rock crevices; 

however, the footprints were common on the paths (Elbroch 2003). 

The fewest tracks we observed were of gray fox (Urocyon cinereo-
USDA Forest Service Proceedings RMRS-P-67. 2013 251

Richness of Mammals on the San Bernardino Ranch . . . Bonillas-Monge and Valdez-Coronel

argenteus). This may have been due to the avoidance of competing 

with another carnivor like coyote that has similar preys; on occasion, 

coyotes can depredate the gray fox (Merlin and Siminski 2000). We 

found a few traces of mountain lion (Puma concolor), which is normal due to being a top predator and less common than the rest of the 

animals because of their solitary and territorial nature. However, their 

presence tells us about the good conditions of the place correlated to 

the abundance of the preys (Pacheco and others 2004).

Conclusions

The San Bernardino Ranch is a place that is undergoing restoration, and, therefore, it is possible to observe favorable changes in the 

fauna composition. These changes are reflected in the individual and 

traces found near water, which is fundamental to the establishment 

of species richness. Comparing our list of species and the list of the 

San Bernardino NWR, we can see that many species remain to be 

documented, due to the fact of the differences between the time of 

restoration on the side of the United States and Mexico. However, 

we recommend that this study be continued to constantly sample 

mammals in the San Bernardino Ranch for several years in a major 

amount of points to obtain reliable results.

The San Bernardino Ranch is a great site for future research efforts 

centered on monitoring the fauna with trap cameras, determining 

the population structure of small mammals, and studying the eating 

habitats of carnivors. This type of research can show us a broader 

picture about the healthy state of the ecosystem in the Ranch. The 

purpose of the work presented in this paper was to obtain presence 

data. However, the necessary preliminary data were also taken for 

future research on the population structures of small (non-flying) 

mammals.

References

Aranda, J.M. 2000. Huellas y otros rastros de los mamíferos grandes y medianos 

de México. CONABIO, Instituto de Ecología, A.C. Xalapa, Veracruz. 212 pp.

Arriaga, L., J.M. Espinoza, C. Aguilar, E. Martínez, L. Gómez y E. Loa (coordinadores). 2000. Regiones terrestres prioritarias de México. Comisión 

Nacional para el Conocimiento y uso de la Biodiversidad, México.

Bojórquez-Tapia, L. A., P. Balvanera, and D.A. Cuaron. 1994. Biological 

inventories and computer data bases: their role in environmental assessments. Environmental Management, 18:775-785.

Burt, W.H. 1943. Territoriality and home range concepts as applied to mammals. Journal of Mammalogy 24: 346-352. 

Caire, William. 1978. The distribution and zoogeography of the mammals of 

Sonora, Mexico. The university of New México. Albuquerque, New México.

Castillo-Gámez, R.A., J.P. Gallo-Reynoso, J. Egido-Villarreal, and W. Caire. 

2009. Los mamíferos. En: F.E. Molina-Freaner y T.R. Van Devender, eds. 

Diversidad biológica de Sonora. UNAM, México, pp. 421-436.

Ceballos G. and G. Oliva 2005. “Los mamíferos silvestres de México” Comisión 

Nacional Para el Conocimiento y Uso de la Biodiversidad.

Ceballos, Gerardo., Arroyo Cabrales, Joaquín., Medellin, Rodrigo A., Domínguez Castellanos, Yolanda. 2005. Lista actualizada de los mamíferos de 

México. Revista Mexicana de Mastozoología. 9:21-71. 2005.

CEDES. 2011. Estudio previo justificativo para proponer la región de San 

Bernadino, municipio de Agua Prieta, Sonora como reserva estatal.

Elbroch, Mark. 2003. Mammal tracks and sign: A guide to North American 

species. StackpoleBooks: Mechanicsburg, PA. 

Grajales-Tam, K.M., R. Rodríguez-Estrella y J. Cancino-Hernández. 2003. 

Dieta estacional del Coyote Canis latrans durante el periodo 1996-1997 

en el Desierto de Vizcaíno, Baja California Sur, México Acta Zoologica 

Mexicana (N.S.) 89: 17-28.

Grigione, M., A. Scoville, G. Soville, and K. Crooks. 2007. Neotropical 

cats in southeast Arizona and surrounding areas: Past and present status 

of jaguars, ocelots and Jaguarundis. Mastozoología Neotropical, 14(2): 

189-199, Mendoza, 2007.

Hudson, P.F., D.A. Hendrickson, A.C. Benke, A. Varela-Romero, A. RodilesHernández y W.L. Minckley. 2005. Rivers of Mexico. En: Benke, A.C. and 

C.E. Cushing, (eds.), Rivers of North America. Elsevier Academic Press, 

Toronto, Canadá.

Krebs, C.J. 1985. Ecología. Estudio de distribución y la abundancia. 2a Ed. 

Harla. México.

Leopold, A.S., 1977. Fauna silvestre de México. Instituto Mexicano de Recursos 

Naturales Renovables. Pax-México, México. 608 pp.

Merlin P. and Siminski P. 2000. Coyote and fox. In: S.J. Phillips and P. 

Wentworth-Comus, eds. A natural history of the Sonoran Desert. The 

University of California Press.

Minckley T.A. and A. Brunelle. 2006. Paleohydrology and growth of a desert 

ciénega. Journal of Arid Environments 69 (2007): 420-431.

Molina-Freaner, Francisco and Thomas R. Van Devender. 2010. Diversidad 

Biologica de Sonora. Mora-Cantúa Editores, S. A. de C. V. México, Distrito Federal.

Molina-Freaner, F.E. and T.R. Van Devender, eds. 2010. Diversidad biológica 

de Sonora. UNAM, México,

Pacheco, L.F., A. Lucero, and M. Villca. 2004. Dieta del Puma (Puma concolor) 

en el parque Nacional Sajama, Bolivia y su conflicto con la ganadería. 

Ecologia en Bolivia. 29: 75-83.

Reid, Fiona and A. Peterson 2006. Field guide to mammals of North America. 

Fourth Edition. New York: national Audubon Society.

Rzedowski, J. 1978. Vegetacion de Mexico. Editorial Limusa, S. A. Mexico D.F.

Urban, D. 1970. Racoon populations, movement patterns, and predation on a 

managed waterflow marsh. Journal of Wildlife Management 34: 372-382.

Whitaker, John O., Jr. 1996. National Audubon Society Field Guide to North 

American Mammals. New York: Alfred A. Knopf. 

Wilson Don E. and Sue Ruff. 1999. The Smithsonian Book of North American 

Mammals. Smithsonian Institution Press.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
252 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Preliminary List of Flying Mammals in the Ajos-Bavispe 

National Forest Reserve and Wildlife Refuge, Sonora

Rosa Elena Jiménez

CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

Christ D. Weise

U.S. Fish and Wildlife Service, Kofa National Wildlife Refuge, Yuma, Arizona

Mario Cirett-Galán, Guadalupe Flores, and Manuel Munguia

CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

E. Isaías Ochoa

Naturalia, A.C., Hermosillo, Sonora

Abstract—Information on bat communities, including their composition, abundance, distribution and ecology, 

can support management programs in protected areas, and also provide information and initiatives for the 

designation of new protected areas. In 2010 and 2011, monitoring was conducted in the Ajos Mountains, 

a sky island, that is part of the Ajos Bavispe Reserve. During this period of time, we recorded a total of eight 

species (Myotis californicus, Myotis auriculus, Myotis volans, Myotis thysanodes, Lasiurus cinereus, Lasiurus 

blossevillii, Eptesicus fuscus y Choeronycteris mexicana), in riparian and oak-pine forests. This preliminary 

species list will assist in assessing ecosystem health in this part of the reserve and provides an example of 

the monitoring we are hoping to extend to the other Sky Islands within the Ajos-Bavispe Natural Protected 

Area: La Madera, El Tigre, Buenos Aires and La Purica. The intention of this paper is to use the information 

we have gathered on abundance and distribution of bat species as a tool to assess the conservation status 

and ecosystem health of these Sky Islands. Bats are a good indicator taxon because they are abundant, 

diverse, and relatively easy and cost effective to monitor. As a result the status of bat communities (composition and abundance) will support conservation and management decisions in the short and long term in 

this important protected area.

Introduction

Research on the abundance, diversity, distribution, ecology, and 

species composition is an essential part of the development and implementation of management programs in natural protected areas (Kalko 

1993). Therefore, it is necessary to develop projects and monitoring 

protocols for natural protected areas to complement and guarantee the 

conservation of the ecosystems with proper management of resources 

over time.

Castner and others (1996) conducted an inventory of bats in wet 

Beaver Creek Wilderness Area (Coconino National Forest) to establish 

a bat management project which implies the importance of and results 

in the preservation of bats. The National Forest Reserve and Refuge 

Fauna Silvestre Ajos- Bavispe (RFN y RFS Ajos-Bavispe), are the 

oldest reserves of Mexico. They were designated in the first decree in 

1936 and included barely 21,494 hectares of national land formed by 

the mountains of “Los Ajos”, “Buenos Aires” and “Purica”. By 1939, 

the Presidential Decree was ratified adding a total of 184 000 hectares 

including the sierras “El Tigre” and “Pilares of Teras”, “Pilares of 

Nacozari”, “San Diego”, and “la Madera”. RFN y RFS Ajos-Bavispe 

is located in the physiographic region of the Sierra Madre Occidental 

(CONANP 2001). These ranges are part of the region called islands 

of the sky (The Sky Islands) and are known to be complex systems 

with uniquely high biodiversity (McLaughlin 1995; Warshall 1995). 

Information on flora and fauna of Ajos Bavispe, a Protected Natural 

Area (ANP) has been collected in recent years; however, information on volant mammals (bats) in the ANP is not known. We believe 

that bats can be used as indicators to assess the conservation status 

of different ecosystems present within the reserve, because of their 

diversity, and the relative ease and cost effectiveness of monitoring 

bats.

The order Chiroptera is ecologically diverse and has a global. 

distribution The development of flight and the sophisticated sensory 

adaptations (echolocation) of these flying mammals has allowed bats 

to inhabit extremely diverse habitats and has provided them with the 

ability to exploit a wide variety of resources for food (Sampaio and 

others 2003, Ávila Flores and Medellín 2004). 

This study is intended to provide an inventory of bats for the RFN 

y RFS Ajos-Bavispe and thus identify sites with greatest diversity and 

abundance. This information will be used for planning and decision 
USDA Forest Service Proceedings RMRS-P-67. 2013 253

Preliminary List of Flying Mammals in the Ajos-Bavispe National Forest Reserve . . . Jiménez and others

making of refuge management when prioritizing conservation actions 

and habitat protection.

Methodology

Sampling was conducted in section 4 of the RFN y RFS Ajos-Bavispe 

situated in the Sierra Los Ajos in Gallery forests and pine-oak forest. 

In addition abandoned buildings, caves and mine adits were sampled. 

All sample sites were near perennial water sources. The sample sites 

are: Cañon de Evans, La Sal and El Guerigo (fig. 1).

The monitoring took place in October 2010 and May, July and 

August 2011, by surveying for bats with mistnets (Ávila and Medellín, 2004; Rabe and Rosenstock 2005). Three to four mistnets of 3, 

6, 9 and 12 m (Martinez 2007; Sánchez and Medellín 2007) were 

deployed at sunset. Sampling was concluded when bat activity was 

no longer detectable or when weather conditions (i.e. low temperatures, high winds or rain) necessitated closing the nets. The mist nets 

were installed close to or over bodies of water, and in flyways used 

by bats (Lopez and Garcia 2006). For the monitoring in abandoned 

buildings, caves or mines hand nets were used to capture and identify 

bats (Lopez and Garcia 2006).

Captured bats were identified and sex, age (adult or juvenile), 

reproductive status (pregnant, nursing, post-lactante, scrotal, nonbreeding, not scrotal), weight, forearm length and ear length (Ávila 

and Medellín 2004; Lopez and Vaughan 2004; Russell and others 2005; 

López and García 2006; Sánchez and Medellín 2007;) were recorded. 

For species identification a key to the bats of Mexico (Medellín and 

others 2008) was used and when necessary we consulted experts to 

assist in species determination. For taking measurements we used 

dial calipers (DialMax) and a ruler, mass was measured with a pesola 

scale (100 g) and photos were taken of all captured bat species.

Results and Discussion 

During our surveys in 2010-2011 we recorded a total of eight bat 

species (Choeronycteris mexicana, Myotis auriculus, Myotis californicus, Myotis thysanodes, Myotis volans, Eptesicus fuscus, Lasiurus 

blossevilli and Lasiurus cinereus) in our study sites. These species 

represent two families of bats Phyllostomidae (leaf-nosed bats) and 

Vespertilionidae (Vesper or evening bats) (table 1). Our sample 

recorded 34% of the 23 possible species that may occur in this area 

according to distribution maps in Medellín and others (2008).

Choeronyteris mexicana was found in an abandoned cabin in the 

months July and August and we estimated the number of individuals 

by having three observers count individuals at the same time. This 

method yielded a result of 90 to 100 estimated individuals. The discovery of this colony for the ANP is important evidence that the Ajos 

Mountains are being used as a roost by this important pollinating bat 

species which is listed as threatened in the NOM-059 (SEMARNAT 

2010). We plan to monitor this roost at Güerigo regularly in order 

to obtain additional information on size and times of roost use. This 

information will assist in determining the actions necessary for protection of this colony by the ANP. 

Figure1—This map depicts the monitoring locations for bats in the Los Ajos Mountains.
254 USDA Forest Service Proceedings RMRS-P-67. 2013

Jiménez and others Preliminary List of Flying Mammals in the Ajos-Bavispe National Forest Reserve . . .

r

This species uses a wide range of habitats including desert scrub, 

deciduous tropical forests and pine-oak forests (Arroyo - Cabrales 

and others 1987). Our study found C. mexicana associated with oak 

forest and forest galleries represented mainly by Quercus oblongifolia, Quercus eggersii, Juniperus deppeana and forest of galleries 

represented chiefly by alder (Platanus wrightii), alamos (Populus 

fremontii), and güerigos (Populus brandengeei). 

Arroyo-Cabrales and others (1987) writes that for C. mexicana migration to Arizona and New Mexico may take place in June, however, 

we recorded the presence of this colony in July and August 2011; it is 

clear that more population and ecological data of this colony is needed 

to understand this species spatial and temporal use of the habitat in the 

Sky Island region. A better understanding of its dynamics is needed 

to make decisions for species conservation. Myotis auriculus, Myotis 

californicus, Myotis thysanodes, Myotis volans, Eptesicus fuscus, 

Lasiurus blossevilli, Lasiurus cinereus, were found in Oak-Juniper 

Forests and forest riparian forests represented mainly by Quercus arizonica, Juniperus deppeana, Acer grandidentatum, Platanus wrightii, 

with perennial water sources. Even though these species may occur 

in various types of habitat, the habitat they use is usually in good 

condition (Shump and Shump 1982, Warner Richard 1982, Warner 

and Czaplewsky 1984; Simpson 1993; Avila and Fenton 2005). As 

preliminary result we can assume that the Canyon de Evans and La 

Sal are in good condition and we recommended the use these sites 

as permanent monitoring stations to obtain abundance, diversity, and 

community structure data for the bat species present in the reserve.

No settlements or anthropogenic activities (i.e. livestock, agriculture) 

are within these sampled sites. They are nationally owned lands which 

have resulted in a good quality of habitat, as mentioned above these 

three sites have perennial water sources. For the Cañon of Evans and 

la Sal, we recorded the presence of Ambystoma rosaceum which are 

indicators of good water quality (Rorabaugh 2008) and complement 

the ecological importance of these sites in the Sierra Ajos.

A greater sampling effort is needed at the study sites, especially 

in the early spring. Bat surveys at additional sites within the Ajos 

mountains are necessary for a comprehensive assessment of the area. 

Also it is important to include Sierra del Tigre, Madera and the Purica 

which are part of the ANP in future sampling efforts.

Acknowledgments

Thanks to Erin Fernandez, Jim Rorabaugh, Scott Richardson, Cat 

Crawford, Gerardo Carreon for their teachings in field and sharing 

their knowledge. Nalleli Lara, Alejandro González, Heli Colonel for 

their support in field.

References

Avila-Flores, R.; M. B. Fenton. 2005. Use of spatial features by foraging 

insectivorous bats in a large urban landscape. Journal of Mammalogy. 86 

(6): 1193-1204.

Arroyo Cabrales, J.; R. R. Hollander; J. Knox Jones.1987. Choeronycteris 

Mexicana. Mammalian Species. 291: 1-5.

Ávila-Flores, R.; R. A. Medellín. 2004. Ecological, taxonomic, and physiological correlates of caves use by Mexican bats. Journal Of Mammalogy, 

85(4). Pp.675-687

Castner, S. V.; T. K. Snow; A. T. Holycross; N. Branch. 1996. Bat inventory 

of the wet beaver creek wilderness area (Coconino County, Arizona). Tech. 

Rep. 106. Arizona Game and Fish Departament, Nongame and Endangered 

Program, Phoenix, AZ.

Comisión Nacional de Áreas Naturales Protegidas. 2001. Estudio previo 

justificativo, área de protección de flora y fauna Mavavi. SEMARNAT. 

México, D.F. 

Kalko, E. K.V. 1993. Acustic survey of neotropical bats. Advantanges and 

disadvantanges. International workshop on neotropical mammals. The 

Nature Conservancy. Panama

Kalko, E. R. 1997: Diversity in tropical bats. In: Urich, H., ed. Tropical 

biodiversity and systematics. Proceeding of the International Symposium 

on Biodiversity and sistematics in tropical ecosystems, Bonn, 1994. Zoologisches Forschungsinstitut And Museum Alexander Koenig: 13-43.

López González, C.; D. F. García Mendoza. 2006. Murciélagos de la sierra 

Tarahumara, Chihuahua, México. Acta Zoológica Mexicana. 22 (2): 109-135. 

López, J. E.; C. Vaughan. 2004. Observations on the role of frugivorous bats 

as seed dispersers in Costa Rican secondary humid forests. Acta Chiropterologica. 6(1): 111-119

Martínez Gallardo, R. 2007. Inventario de los murciélagos de los parques 

nacionales sierra de san pedro mártir y constitución de 1857, baja california, 

México. universidad autónoma de Baja California. Informe Final SNIBCONABIO Proyecto No. BK009 México D. F.

Medellín, R. A.; H. Arita; O. T. Sánchez. 2008. Identificación de los murciélagos de campo de México Clave de campo. Segunda edición. Instituto de 

Ecología, UNAM. México 

Mclaughlin, S. P. 1995. An Overview Of The Flora Of The Sky Islands, 

southeastern Arizona: Diversity, affinities, and insularity. In: Debano, L.F. 

and others, eds. Biodiversity and management of the Madrean Archipelago. 

Gen. Tech. Rep. RM-GTR 264. Fort Collins, CO: U.S. Department Of 

Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 60-70

O’Farrell, M. J.; E. H. Studier. 1980. Myotis thysanodes. Mammalian Species. 137: 1-5.

Rabe, M. J.; Rosenstock S. S. 2005. Influence of water size and type on bat 

captures in the lower Sonoran Desert. Western North American Naturalist.65(1): 87-90.

Rorabaugh, J. C. 2008. An introduction to the herpetofauna of mainland Sonora, 

México, with comments on conservation and management. Journal of the 

Arizona-Nevada Academy of Science. 40(1): 20-65.

Table 1—Displays the documented species in RFN and RFS Ajos-Bavispe.

Family	 Scientific name	 Common name	 Number of individuals

Phyllostomidae Choeronycteris mexicana Murciélago nariz de cochi 90-100

Vespertilionidae Myotis auriculus Miotis Suroeste 7

Myotis californicus Murciélago de California 5

Myotis thysanodes Miotis Bordado 6

Myotis volans Murciélago de patas largas 2

Eptesicus fuscus Murciélago grande café 7

Lasiurus blossevilli Murciélago rojo occidental 2

Lasiurus cinereus Murciélago canoso 4
USDA Forest Service Proceedings RMRS-P-67. 2013 255

Preliminary List of Flying Mammals in the Ajos-Bavispe National Forest Reserve . . . Jiménez and others

Russell A. L; R. A Medellín; G .F. Mccracken.2005. Genetic variation and 

migration in the mexican free-tailed bat (Tadarida Brasiliensis Mexicana). 

Molecular Ecology. 14: 2207-2222.

Sampaio E. M.; E. K. V. Kalko; E. Bernard; B. Rodriguez- Herrera; C., Jr. 

Handley. 2003. A biodiversity assessment of bat (Chiroptera) in a tropical 

lowland rain forest of central Amazonia, including methodological and 

conservation considerations. Estudies on Neotropical Fauna and Environment. 38(1):17-31.

Sánchez, R.; R. A. Medellín. 2007. Food habits of the threatened bat Leptonycteris Nivalis (Chiroptera: Phyllostomidae) in a mating roost in Mexico. 

Journal Of Natural History. 41(25):1753 – 1764.

Simpson, M. R. (1993). Myotis californicus. Mammalian Species. 428: 1-4.

SEMARNAT. 2010. NORMA Oficial Mexicana NOM-059-ECOL-2010, 

Protección Ambiental-Especies Nativas De México De Flora Y Fauna 

Silvestres-Categorías De Riesgo Y Especificaciones Para Su Inclusión, 

Exclusión O Cambio-Lista De Especies En Riesgo. Diario Oficial 30 De 

Diciembre De 2010. México D.F.

Snow, T.; B. Taubert.; M. Tuttle. D. Pictorial key to the bats of Arizona. Arizona 

Game And Fish Department; Bat Conservation International.

Warshall, P. 1995. The Madrean Sky Island Archipelago: A planetary overview. 

In: DeBano, L. F. and others, eds. Biodiversity and management of the 

Madrean Archipelago. Gen. Tech. Rep. RM-GTR 264. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest 

and Range Experiment Station: 7-18

Warner, R, M. (1982). Myotis auriculus. Mammalian Species.191:1-3. 

Warner, R. M.; N. J. Czaplewsky. 1984. Myotis volans. Mammalian Species. 

224:1-4.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
256 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Decade of Wildlife Tracking in the Sky Islands

Jessica A. Lamberton-Moreno

Wildlife Linkages Program, Sky Island Alliance, Tucson, Arizona

Sergio Avila-Villegas

Northern Mexico Program, Sky Island Alliance, Tucson, Arizona

Abstract— In 2001 Sky Island Alliance developed a citizen science program that uses track and sign identification 

and count surveys to monitor potential wildlife corridors throughout southeastern Arizona and southwestern 

New Mexico. The goal of the Wildlife Linkages Program is to protect and advocate for an interconnected 

landscape where wildlife, based on their ecological needs, can move easily between core habitats, the Sky 

Island mountain ranges. Currently, we train and engage volunteers in the monitoring of fifty 1.5-mile long 

transects within seven priority linkage areas; the majority of these study areas are located on public lands. To 

date we have conducted over 1,000 track count surveys and documented over 4,100 records for more than 

40 different animal species in the region. Sky Island Alliance has successfully applied the resulting species 

presence data to land-use policy and permanent land conservation, incorporating wildlife data and corridor 

priorities into the Sonoran Desert Conservation Plan, the Santa Cruz County Comprehensive Plan, the Pima 

County Wildlife Connectivity Assessment, and the Arizona Wildlife Linkages Assessment. 

Introduction

	 The ability for long-ranging species to disperse between mountain ranges and intervening valleys is paramount to species survival

(Opdam 1990; Hass 2000; Bennet 2003; Arizona Wildlife Linkages

Workgroup 2006). Preserving, restoring, and increasing connected

ecosystems within the Sky Island region, has ever-greater urgency.

Habitat fragmentation and destruction of open space caused by irresponsible off-road vehicle use and the development of new roads

and highways, transmission lines, border infrastructure, and other

effects of rapid human expansion are the most serious short-term

threats to Sky Island species and their habitats (Avila and others in 

press). Division of otherwise continuous habitat will prevent natural

movements of some species whose distribution in the U.S is already

limited, confining them to isolation and potential extinction. In an

elevational gradient, limiting wildlife migration movements from

lowerdesertregionstohigher elevations couldbedevastating(Misztal

and others 2012). This would also have far-reaching effects on other,

interconnected species, such as prey.

	 In the arid southwest, where extreme summer temperatures and

long distances between watersources already affectspeciessurvival,

climate change is adding another layer of difficulty. The ability of

species to adapt and respond to a changing climate will depend on

their ability to move freely across the landscape (Misztal and others

2012).An intact healthy landscape maintainsthe ability of wildlife to

movebetweencoreareaswherespecies—bothplantandanimal—have

sufficient resources to survive, reproduce, and otherwise facilitate

ecological processes (Hass 2000; Soulé and Terborgh 1999).

	 Tracking is an excellent tool for wildlife corridor monitoring over

time. The skill of wildlife tracking is hundreds of thousands of years

old and seen by some as the origin of science (Long and others 2008;

CyberTracker Conservation 2012). Although somewhat forgotten

except as a primitive skill and recreational practice, tracking as a

technique continues to have implications for modern science and

for wildlife conservation (Haynes and others 2005; CyberTracker

Conservation 2012).

	 Trackingisanon-invasivemonitoringtechniquethat,withdeliberate

and careful study design, can provide presence, relative abundance,

occurrence, distribution, and population density information (Long

and others 2008). Wildlife tracking is also an effective tool to engage

and educate the public, and attracts a wide range of people from

different backgrounds as supporters and advocates for wildlife and

their habitats. Track count events have become extremely valuable in

bringingtogether activists,hunters,biologists, agencyrepresentatives,

military personnel, planners, landowners, academics and volunteers

in a shared interest (Linton 2003; Lamberton,Avila and Morris 2011;

Sky Island Alliance 2012). Track and sign identification has some

limitations in terms of its subjectivity, variable observer skills, and

difficulty in distinguishing individual animals, determining gender,

or estimating populations.

	 Tracking remains a cost-effective, easy to learn, and engaging

monitoringmethodbywhichwe candocumenttrendsinsigndetection

over time and wildlife presence in specific areas, particularly with

protocolstoprovidedata consistencybasedona scientific studydesign

(Haynes and others 2005; Beier and Cunningham 1996; Smallwood

and Fitzhue 1995; Lamberton, Avila and Morris 2011). In addition,

track and sign surveys can confirm the presence ofrare speciesin new

areas(Long and others 2008; Squires and others 2004). For example,

Sky IslandAlliance’s first documentation of jaguar in the SierraAzul

range of Sonora, Mexico, came from the tracks discovered in 2009,

one year before the animal triggered a nearby remote camera.
USDA Forest Service Proceedings RMRS-P-67. 2013 257

Decade of Wildlife Tracking in the Sky Islands Lamberton-Moreno, and Avila-Villegas

	 In 2001, Sky Island Alliance (SIA) developed a citizen science

program that uses track and sign identification and surveys to monitor potential wildlife corridors throughout southeastern Arizona and

southwesternNewMexico.ThegoaloftheWildlifeLinkagesProgram

is to protect and advocate for an interconnected landscape where

wildlife, based on their ecological needs, can move easily between

core habitats, the Sky Island mountain ranges. We use wildlife tracking to gather data on wildlife presence in key wildlife linkage, and

engage citizen scientists and advocates. Over the last decade we have

monitored fifty 1.5-mile long transects within seven priority linkage

areas in southern Arizona and New Mexico; the majority of these

located on public lands.This article focuses on ourlongestmonitoring

effort located in the Las Ciénegas Wildlife Linkage, using Davidson

Canyon transects as case study examples.

Study Area

	 The Sky Island region is a 70,000-square-mile world biodiversity

hotspot (Mittermeier and others 2005), a biological bridge where the

temperateRockyMountainsandColoradoPlateaumeetthesubtropics,

the Sierra Madre Occidental, and the Sonoran and the Chihuahuan

Deserts.

	 Within this biological region, the Las Ciénegas Wildlife Linkage,

which connectsthe Rincon Mountainsto the Empire,Whetstone, and

Santa Rita Mountains, has been identified as a high-priority wildlife

linkage (Arizona Wildlife Linkages Workgroup 2006; Beier, Majka,

and Bayless 2006). Sky IslandAlliance’s work in this corridor began

in 2001 when we conducted a 4-month assessment on 10 transects

located throughout the area. The results were included in the “Missing Link Report” (also referred to as the Ciénega Creek Corridor),

presentedtotheBureauofLandManagement(SonoranInstitute2003).

Subsequently, Pima County recognized the ecological importance of

this area and designated the linkage as a Critical Landscape Connection in the Pima County Sonoran Desert Conservation Plan (2004).

	 SIA continued this study and has monitored 10 transects consistently between 2002 and 2011 in the Las Ciénegas Wildlife Linkage

(fig.1). Transects occur both north and south of Interstate-10; four

transects occur within the Las Ciénegas National Conservation Area

under Bureau of Land Management jurisdiction.

	 Davidson Canyon is an ephemeral stream that runs south to north,

provides wildlife habitat and water recharge in the Ciénega-Rincon

watershed.Itsrelationship to the perennialCiénegaCreek and endangeredspeciessuchastheChiricahualeopardfrog(Rana chiricahuensis)

andgila topminnow(Poeciliopsis occidentalis occidentalis) hasledto

concerns over proposed mining projects in the Santa Rita mountains

(Coronado National Forest 2011; Sky IslandAlliance 2011).The high

and expansive bridge thatspansthe canyon at Interstate-10 enhances

the importance of Davidson Canyon as a wildlife corridor, which is

one of the few drainages in the Las Ciénegas Wildlife Linkage that

facilitates safe passage of wildlife across Interstate-10. Davidson

Canyon also is a direct connector for the Santa Rita Mountains across

Scenic Highway 83, making it an integral piece of the Santa RitaRincon-Whetstone mountain linkage complex.

	 There are several recent confirmed sightings of jaguars and ocelots

in southeast Arizona (Avila and others In press). It is possible that

these species are moving north from the Patagonia Mountains to the

Santa Rita Mountains, and across Highway 83 utilizing Davison

Canyon and the Las Ciénegas Wildlife Linkage to reach large habitat

blocks in locations to the east, such as the Whetstone and Dragoon

Mountains.Conversely, black bear periodically use the same pathway

moving south from theRincon Mountainsto the SantaRitas(Atwood

and others 2011), thereby bolstering the genetic health of a newly

discovered Sonoran population, a population ironically made distinct

by itsisolation from northernArizona mountain ranges(Atwood and

others 2011).

Methods 

	 The Wildlife Linkages Program track identification and data

collection protocol is adapted from Keeping Track, Inc. (Hass and

others 2000; Przybyl 2003) to consider regional requirements, like

dry tracking conditions in sand and dirt rather than snow. Transects

were established in areas most likely to provide evidence of wildlife

activity, primarily in sandy washes, dirt roadways, or riparian edges,

where there issuitable substrate fortracking.We established strategic

1.5-mile longby60-feetwide transectsinpotentiallinkages,surveyed

every 6 weeks by teams of trained volunteers. In aggregate, teams

conduct nine surveys per year: two surveys occurring each season,

with three surveys occurring in summer to capture data during presummer and monsoon conditions.

	 Volunteers go through a five-day training course in track and sign

identification and documentation in the field. Teams of two to four

trained trackers are then permanently assigned to one transect, and

are provided with datasheets, GPS units, cameras, track identification cards and tracking rulers. A minimum of two trained trackers is

required for each survey. A data point is recorded when the tracking

team come to consensus about the identification of a track or sign.

	 Tracking teams survey in early morning, when temperatures are

cool, the angle of light refraction is optimal for track visibility and

photo-documentation, and nocturnal species tracks are freshest.

Tracking surveys conducted later in the day have a lower chance

of detecting nocturnal or crepuscular species and a higher chance

of detecting diurnal species (Hanson and Hanson, personal communication). In early morning, there is also a higher probability of

seeing wildlife tracks that increased human and cattle activity might

eliminate during the day. No surveys are conducted during rainfall,

due to low visibility of tracks and field safety.

	 Sky Island Alliance’s Wildlife Linkages Program focuses on regional animal focal species categorized as a keystone, umbrella, or

indicator species, habitat specialists, or have an unknown or shifting

population or distribution. Threatened or endangered species often

fit many of these categories. Focal species are plants or animals

whose survival needs are also requirements of a healthy ecosystem,

therefore making them indicators of ecosystem health (Miller and

others 1999). Eight focal species were identified for study: jaguar

(Panthera onca), ocelot (Leopardus pardalis), bobcat (Lynx rufus),

mountain lion (Puma concolor), Mexican gray wolf (Canis lupus 

baileyi), black bear (Ursus americanus), white-nosed coati (Nasua 

narica), and western ornate box turtle (Terrapene ornata).

	 Each occurrence of focalspecies’track or sign is documented on a

data collection form, and assigned a data point and photo number. A

data point is a sign or set of signs made by a single animal at a single

time. Tracks and other sign identified belonging to focal species are

photographed with a reference ruler, measured, and information is

collected on direction of travel and location. Any non-focal species

indentified on a survey are recordedwith an abundance rating, defined

as A (one to five occurrences of sign); B (six to 10 occurrences of

sign); and C (>10 occurrences of sign). Additional comments and

observations are included; the resulting field notes provide excellent

anecdotal information. In addition to track and sign counts we gather

baseline information to document changes in transect condition, including evidence of off-road vehicle use, erosion, presence of stock

animals, and new barriers to wildlife movement.
258 USDA Forest Service Proceedings RMRS-P-67. 2013

Lamberton-Moreno, and Avila-Villegas Decade of Wildlife Tracking in the Sky Islands

Figure 1—Study area within the Las Ceiengas Wildlife Linakge and locations of the ten tracking transects (2002-2011) (map by Nick Deyo).
USDA Forest Service Proceedings RMRS-P-67. 2013 259

Decade of Wildlife Tracking in the Sky Islands Lamberton-Moreno, and Avila-Villegas

Results

	 SIAhas conducted over 1,000 track-countsurveys and documented

over4,100recordsformorethan40differentanimalspeciesthroughout

the region since 2001. In the Las CiénegasWildlife Linkage, we have

documented 31 species with track and sign surveysthroughout the 10

transect study sites between 2002 and 2011 (table 1). The majority

are medium to large mammals, including both habitat specialists and

generalists. We were successful in identifying tracks or sign for the

majority ofmediumto largemammals known to be presentin the area,

including all fourspecies of native skunks. We identified lagomorphs

to genus, including the eastern and desert cottontailrabbit(Sylvilagus 

spp.), and the antelope and black-tailed jackrabbit (Lepus spp.).

	 We compared transect results using the proportion of surveys with

track and sign detection of similar large and medium sized mammal

species (fig. 2). We categorized these by obligate carnivores: mountain lion and bobcat; omnivores: black bear and white-nosed coati;

and herbivores: mule deer (Odocoileus heminonus), white-tail deer

(O. virginianus) and collared peccary (Tayassu tajacu). Detection of

ungulate track and sign in these transects appeared to be lower than

in the more northern transects in the study area, possibly due to these

species preferring open habitat found away from Cienega Creek.

	 The two transects that make up the 3-mile stretch of Davison

Canyon where it flows underneath Interstate 10 have been monitored

simultaneously between 2002 and 2006. We documented 24 species

here during the 4-year period (table 1). We compared North and

South Davidson Canyon transects for four of these species: black

bear, bobcat, white-nosed coati, and mountain lion (fig. 3). There

is a marked difference between the two transects for both mountain

lion and white-nosed coati, although we must acknowledge that this

could be a factor of observer bias or occasional track detectability

changes caused by substrate or weather.

	 One noteworthy resultfromthisstudy wasthat black beartrack and

signincreaseddramaticallyduringthemonthsofApril andSeptemberOctober across all transects. Although the sample size was relatively

small, all black bear tracks documented in North and South Davison

Canyon transects during this period showed upstream travel (north to

south).Thisis consistent withAtwood’s finding that individualsfrom

a northern population periodically disperse to a genetically distinct

population of black bears found south of Interstate 10 (Atwood and

others 2001).

Applying Tracking to Conservation

	 Protected wildlife linkages are the future of sustainable wildlife

conservation and rural landscape protection. Over 250 dedicated

citizens have participated in our program collecting information and

advocating for wildlifemovement in the Sky Island region.One ofthe

most impressive successes out of the decade of wildlife monitoring

at Sky Island Alliance is the shift in perception about wildlife corridors and landscape level planning by collaborating agencies and

stakeholders.

	 Wehavesuccessfullyappliedtheresultingspeciespresenceinformationtoland-usepolicyandpermanentlandconservation,incorporating

wildlife data and linkage priorities into the Pima County Sonoran

Desert Conservation Plan (2004) and the Santa Cruz County Comprehensive Plan (2005).These results also informed the 2006Arizona

Wildlife Linkages Assessment and the 2012 Pima County Wildlife

ConnectivityAssessment, two comprehensive mapping projects that

identify at-riskwildlife corridors andcontribute recommendationsfor

transportation planners and engineers, created through the Arizona

WildlifeLinkagesWorkgroup(ArizonaWildlifeLinkagesWorkgroup

2006; Arizona Game and Fish Department 2012).

	 Wildlife monitoring data and volunteer advocacy have been instrumental in documenting Wilderness quality lands and providing

momentumtoeffortstopermanentlyprotect corehabitat asdesignated

Wilderness. In addition, with wildlife tracking information we have

provided strong, accurate responses to transportation infrastructure,

mining and energy development projects that threaten to impact key

wildlife blocks and adversely affect associated wildlife linkages.

For example, we provided strong evidence to defeat a proposed

amendment to the 2040 Regional Transportation Plan that called for

widening Scenic Highway 83 from two to four lanes. This project

would have significantly damaged the integrity of the Las Ciénegas

Wildlife Linkage without extensive wildlife crossing structures in

place to mitigate the loss of connectivity and allow animals to cross

the highway safely.

	 Three wildlife crossing structures, including the second wildlife

overpassto be constructed inArizona, were recently approved as part

of a needed expansion of StateRoute (SR) 77 in Oro Valley. Ourtrack

and sign monitoring data, theArizona Wildlife LinkagesAssessment

(2006) and detailed corridor modeling (Beier, Garding and Majka

2006) in the Catalina-Tortolita mountain wildlife linkage supported

this decision. Similarly, in December 2011, 10 years after tracking

volunteers first put feet to the ground, a new project on SR86 near

Kitt Peak proposed by the Tohono O’Odham Nation and informed

by the Arizona Wildlife Linkages Assessment gained final approval

Table 1—Species documented by tracks and sign on Las Cienegas 

Linkage transects (n = 10) between 2002-2011.

Common name Scientific name

Badger Taxidea taxus

Black beara Ursus americanus

Bobcata Lynx rufus

Collared peccarya Tayassu tajacu

Cottontail rabbita Sylvilagus spp.

Coyotea Canis latrans

Domestic cata Felis catus

Domestic cattlea Bos taurus

Domestic doga Canis lupus familiaris

Domestic horsea Equus caballus

Gambel’s quaila Callipepla gambelii

Gila monstera Heloderma suspectum

Gray foxa Urocyon cinereoargenteus

Great blue heron Ardea herodias

Hooded skunka Mephitis macroura

Jackrabbita Lepus spp.

Kangaroo rat Dipodomys spp.

Mountain liona Puma concolor

Mourning dove Zenaida macroura

Mule deera Odocoileus hemionus

Ornate box turtle Terrapene ornata

Raccoona Procyon lotor

Ringtaila Bassariscus astutus

Roadrunnera Geococcyx californianus

Sonoran desert tortoise Gopherus morafkai

Striped skunka Mephitis mephitis

Western hognose skunka Conepatus mesoleucus

Western spotted skunka Spilogale gracilis

White-nosed coatia Nasua narica

White-tailed deera Odocoileus virginianus

Wild turkey Meleagris gallopavo 

a Species documented in Davidson Canyon.
260 USDA Forest Service Proceedings RMRS-P-67. 2013

Lamberton-Moreno, and Avila-Villegas Decade of Wildlife Tracking in the Sky Islands

Figure 2—A comparison of the frequency of track and sign detection in 10 study sites in the Las Ceiengas Wildlife Linakge for 

select carnivore, omnivore, and herbivore species (2002-2011). The location of Interstate-10 is indicated on each graph, and 

transects are arranged left to right as south to north to provide a spatial illustration. The southernmost transects (Gardner Canyon 

A, Ciénega Creek A, North Canyon, and Forty-nine East) occur within the Las Ciénegas National Conservation area.
USDA Forest Service Proceedings RMRS-P-67. 2013 261

Decade of Wildlife Tracking in the Sky Islands Lamberton-Moreno, and Avila-Villegas

for PimaCountyRegionalTransportationAuthority.This project will

include two wildlife underpasses and a vegetated wildlife overpass.

The SR77 wildlife-crossing project, scheduled to break ground in

2013, is serving as a model for similar projects in Pima County. As

population and transportation needs increase, these critical wildlife

connections will become more necessary. There is continued value

forpublic engagement,increasedpublicunderstandingofthe intrinsic

value of wildlife and their habitats, and the information gathered that

track and sign monitoring provides.

Conclusions

	 Long term tracking results have provided useful information about

the presence and movement of wildlife in the Las Ciénegas Wildlife

Linkage and Davidson Canyon. Track and sign monitoring provides

an excellent opportunity for cost-effective, publicly engaging, data

collection that has strong implications for conservation. It serves as

both an advocacy tool as well as a scientific method of determining

wildlife presence. We recommend using track and sign identification

for landscape scale wildlife monitoring efforts as a non-invasive

alternative or to provide supplemental data where technicians or

volunteers are conducting field investigations and the project may

benefit from these additional observations.

Acknowledgments

	 This program and its success would not be possible without the

dedication and skill of our volunteer trackers. We would like to thank

our tracking instructors over the years: Steve Bless,Janay Brun,Jack

Childs, Jonathan and Roseann Hanson, Christine Hass, Lisa Haynes,

AletrisNeils,HarleyShaw,SheridanStone, andCynthiaWolf.Kristen

Campbell,JillKelleman,MelissaLamberton,LouiseMisztal,Christine

St. Onge, and Alex Smith assisted with data synthesis and editing.

Dr. David Bygott and Dr. Jeanette Hanby provided useful reviews.

Our methods are based on the Keeping Track, Inc.© project and

data management protocol. We appreciate Susan Morse for her assistance in the establishment of the monitoring and training protocol,

and Janice Przybyl and Roseann Hanson who were instrumental in

the long-term development of the Wildlife Linkage Program. Many

thanks to Karen Simms and the Bureau of Land Management for

working with Sky Island Alliance over the last decade to monitor

transects onLasCiénegas NationalConservationArea.We also thank

our funders: Kresge Foundation, MET Foundation, the Nina Mason

PulliamCharitableTrust,SummerleeFoundation,TransWildAlliance

and anonymous donors.

References 

ArizonaGame andFishDepartment.2012.ThePimaCountyWildlifeConnectivityAssessment:ReportonStakeholderInput.Available:http://www.azgfd.

gov/w_c/documents/PimaCountyWildlifeConnectivityAssessment.pdf.

Arizona Wildlife Linkages Workgroup. 2006. Arizona’s Wildlife Linkages

Assessment.Available:http://www.azdot.gov/inside_adot/OES/AZ_WildLife_Linkages/assessment.asp.

Atwood, T. C.,J. K.Young,J. P. Beckmann, S.W. Breck,J. Fike, O. E. Rhodes

Jr., and K. D. Bristow. 2011. Modeling connectivity of black bears in a

desert sky island archipelago. Biological Conservation 144.

Avila, S. 2009. The Cuatro Gatos Project: wildlife research and conservation

in northwestern Mexico and implications of the border fence. Wild Felid

Monitor. Winter 2009. Vol. 2(1).

Avila, S., Quigley, M., Hare, T., Carter, J. and M. Emerson. In press. Sky

Island Alliance: Bi-National Efforts to Protect and Restore Native Wildlife and Habitats both Sides of the Border. In: The Border Wall between

Mexico and the United States. Venues, Mechanisms and Stakeholders for

a Constructive Dialogue, Cordova, A. y C. de la Parra (coords). Colegio

de la Frontera Norte.

Beier,P., andS.C.Cunningham.1996.Poweroftracksurveystodetect changes

in cougar populations. Wildlife Society bulletin 24:540-546.

Beier, P., D. Majka, and T. Bayless. 2006.Arizona Missing Linkages: RinconSanta Rita-Whetstone Linkage Design. Report to Arizona Game and Fish

Department. School of Forestry, Northern Arizona University.

Figure 3—Number of track and sign counts documented on South and North Davidson Canyon transects over a period of 4 

years (2006-2009)
262 USDA Forest Service Proceedings RMRS-P-67. 2013

Lamberton-Moreno, and Avila-Villegas Decade of Wildlife Tracking in the Sky Islands

Beier, P., E. Garding, and D. Majka. 2006.Arizona Missing Linkages: Tucson

– Tortolita – Santa Catalina Mountains Linkage Design. Report toArizona

GameandFishDepartment.SchoolofForestry,NorthernArizonaUniversity.

Bennet, A. F. 2003. Linkages in the Landscape: The Role of Corridors and

Connectivity in Wildlife Conservation. IUCN, Gland, Switizerland and

Cambridge, UK. Xiv + 254 pp.

Coronado National Forest. 2011. Draft Environmental Impact Statement for

the Rosemont Copper Project: A Proposed Mining Operation. Coronado

National Forest, Pima County, Arizona. Vol. 1. Available: http://www.

rosemonteis.us/draft-eis.

CyberTracker Conservation. 2012. The art of tracking. Available: http://

cybertracker.org/tracking/the-art-of-tracking.

Hanson,J., andR. Hanson. 2011.Tracking etiquette. Presentation atIntroduction to Wildlife Tracking workshop; July 14; Tucson, AZ.

Hass,C.C.2000.Landscapefragmentationandconnectivityforcarnivoresinthe

UpperSanPedroBasin.FortHuachucaWildlifeOffice,FortHuachuca,AZ.

Hass, C.C., S. C. Morse, and H. G. Shaw. 2000. Keeping Track Project and

Data Management Protocol. Keeping Track, Inc. Vermont, USA.

Haynes, L., Z. Hackl, and M. Culver. 2005. Wild Cats of the Sky Islands: A

SummaryofMonitoringEffortsUsingNoninvasiveTechniques.InConnecting mountain islands and desert seas: biodiversity and management of the

MadreanArchipelago II, compiled by Gottfried, Gerald J.; Gebow, Brooke

S.; Eskew, Lane G.; and Edminster, Carleton B. Proceedings RMRS-P-36.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 631 p.

Lamberton, J. A., S. Avila and K.Morris. 2011. Where jaguar and black

bear meet: tracking the pathway of temperate and tropical species in an

international Sky Island corridor. In Proceedings of the 2011 International

Conference on Ecology and Transportation, edited by Paul J. Wagner,

Debra Nelson, and Eugene Murray. Washington State Department of

Transportation, Seattle, WA.

Linton, T. 2003. Fort Huachuca Surveys Big Cats. U.S. Army Environmental

Command, Fall 2003 Environmental Update.

Long, R. A., P.MacKay, J. C. Ray, W. J. Zielinski, ed. 2008. Noninvasive

Survey Methods for Carnivores. Island Press. Washington, DC.

Miller, B., R. Reading, J. Strittholt, [and others]. 1999. Using focal species

in the design of nature reserve networks. Wild Earth Winter 1998/1999.

Misztal,L.M,A. Score, andL.Hansen. 2012.Between aRock and aHot Place:

Climate Change Adaptation and Resource Management for the Sky Island

Region Workshop Summary Report. Sky Island Alliance and Ecoadapt.

Available:http://www.skyislandalliance.org/adaptationworkshop2011.htm

Mittermeier R.A., P. Robles Gil, M. Hoffman, [and others]. 2005. Hotspots

Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial

Ecoregions. Conservation International, Washington, DC, 392 pp.

Opdam, P. 1990. Dispersal of fragmented populations: the key to survival.

pp. 3-17 In Species Dispersal in Agricultural Habitats, edited by R.G.H.

Bunce and D.C. Howard. Belhaven Press: London.

Pima County Sonoran Desert Conservation Plan. 2004. Biological Corridors

and Critical Habitat. Available: http://www.pima.gov/sdcp.

Przybyl, J.. 2003. Management Model and Training Program for Sky Island

Alliance’s Volunteer-based Wildlife Monitoring Program. Master’s thesis.

Prescott College, AZ.

SantaCruzCountyComprehensivePlan.2005.Available:http://www.co.santacruz.az.us/com_development/pdf/complete-plan-2005.pdf

Sky Island Alliance. 2011. Comments re: Draft Environmental Impact Statement for the Rosemont Copper Project, A Proposed Mining Operation,

Coronado National Forest, Pima County, AZ. Submitted to the U.S. Forest

Service January 18, 2012. Available: https://cara.ecosystem-management.

org/Public/ReadingRoom?Project=24544

Sky Island Alliance. 2012. Wildlife Linkages 10 Year Report: Celebrating a

decade of citizen science and Sky Island corridor conservation. Sky Island

Alliance, Tucson, AZ.

Smallwood, K. S., and E. L. Fitzhugh. 1995. A track count for estimating

mountain lion Felis concolor californica population trend. Biological

Conservation 71:251-259.

Sonoran Institute. 2003.MissingLinkAssessmentforProtectionMeasuresfor

Certain Lands North of the Sonoita ValleyAcquisition Planning District in

Pima County,Arizona. Final Report for Congress, prepared for the Bureau

of Land Management, U.S. Department of the Interior.

Soulé, M. E., and J. Terborgh, ed. 1999. Continental conservation: scientific

foundations of regional reserve networks. Island Press, Washington, DC.

Squires, J. R., K.S. McKelvey, and L.F. Ruggiero. 2004. A snow tracking

protocol used to delineate local lynx, Lynx Canadensis, distributions.

Canadian Field-Naturalist 118:583-589.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 263

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Black Bear Population and Connectivity in the Sky Islands 

of Mexico and the United States

N. E. Lara-Díaz and C. A. López-González

Universidad Autónoma de Querétaro, Qro., Mexico

H. Coronel-Arellano and A. González-Bernal

Naturalia, A. C. Sonora, Mexico

Abstract—The Sky Island region is a mountainous region surrounded by grasslands, deserts and intermountain 

valleys, located between Mexico and the United States. However, different land management and human 

impact can have an effect on its wildlife populations. Currently, the border wall poses an immediate threat 

to the survival of black bears (Ursus americanus), considered an endangered species in Mexico. Our aim 

was to determine the conservation status of black bears in the Sierra San Luis as affected by the border 

fence. We determined population size through camera traps and radio-telemetry, and modeled population 

occupancy using PRESENCE. We documented a bear population with more than 500 individuals. Surveys 

along the border failed to detect bears crossing it, but we identified linkages between the two countries, 

which shall be important for future landscape planning. Increased vehicular traffic, migration, and drug traffic 

have a negative effect on bear populations, exacerbated by an increase in anthropogenic activities resulting 

from the construction and maintenance of the border wall. We recommend modifications to the structure of 

the border wall, and to increase wildlife monitoring by the United States authorities in order to reduce the 

potential impacts that this structure has on black bears and other wildlife populations.

Introduction

The black bear is the largest carnivore and the only Ursid species 

present in Mexico (Hall 1981; Doan-Crider and Hellgren 1996). 

Historically, its distribution included the States of Sonora, Chihuahua, 

Coahuila, Tamaulipas, Durango, Zacatecas, Sinaloa, San Luis Potosí, 

Jalisco, Nayarit, and Aguascalientes (Leopold 1959; Baker and Greer 

1962; Hall 1981; Delfín-Alfonso and others 2011). However, during 

the 20th century, black bears were hunted, captured, and poisoned 

as a means of protection, because they were perceived as a threat to 

livestock and crops; the bears were harvested commercially for their 

skins and fat (Baker 1956; Leopold 1959; Medellín and others 2005). 

These actions caused to loss of 80% of their range (SEMARNAP-INE 

1999).

	 Black bears currently are classified as endangered in most of its

distribution in Mexico (SEMARNAP-INE 1999; SEMARNAT 2010). 

Although harvesting and hunting is prohibited (CITES 2009) and 

there are programs for black bear conservation (PROCER; CONANP 

20011), their populations are at risk because of habitat loss, fragmentation (Leopold 1959; Pelton 1982; Robins and others 2004), and other 

human activities (Costello and others 2001).

The populations in the Sierra Madre Oriental has increased due 

to protection through the implementation of Management Units for 

Wildlife Conservation (UMAs) and Natural Protected Areas. These 

populations are more abundant compared with populations in the 

Sierra Madre Occidental (Doan-Crider 1995a; López-González and 

Lara-Díaz 2010).

In the Northern part of the Sierra Madre Occidental (SMO), bears 

remain in a naturally fragmented habitat, the Sky Islands, (Varas 

2010; Delfín-Alfonso and others, in press) in Mountain ranges as 

San Luis, Los Ajos, El Tigre, Sierra Azul, La Elenita, Buenos Aires, 

and El Gato (Gallo and Garza 2002). However, information about 

their population status is poor. Sierra de San Luis may be the “source 

population” of individuals to the rest of the Sky Islands in Mexico 

and one of the primary connectivity areas with the populations in 

United States, therefore, contributing to the recovery of black bears 

in Mexico (Lara-Díaz 2010; López González and Lara-Díaz 2010; 

Varas 2010). Additionally, in the SMO and Sky Islands, there are 

immediate threats that can bring negative effects from short- to longterm for black bear populations (List 2007) mainly because of the 

construction and operations of the border fence between Mexico and 

the Unites States (Public Law 109-367 2006), and the expansion in 

two additional lanes of Federal Highway No. 2 (Agua Prieta-Janos) 

located within 5 km south of the border.

Black bears must maintain their mobility and their ability to grow 

and maintain viable populations if they are to survive in a highly 

fragmented landscape (Meffe and Carroll 1997; Crooks 2002; Duffy 

2002; List 2007). Given the uncertain status of black bears in the Sky 

Islands, however, it becomes necessary to establish conservation 

and habitat management actions, through a solid base of biological 

knowledge of the species, in order to give viability to their populations in the long-term (Rogers 1993; Robins and others 2004). Our 
264 USDA Forest Service Proceedings RMRS-P-67. 2013

Lara-Díaz and others Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States

objective was to generate information on the current status of the 

black bear population in Sierra de San Luis and identify key areas of 

connectivity between the two countries.

Study Area

	 The Sierra de San Luis (SSL; fig. 1) is located at the NE of Sonora

(31° 11’ N, 108° 56’ W) within the Sky Islands region, which consists 

of mountains that are isolated from one another by large extensions 

of desert and grasslands (Onorato et al. 2003; The Wildlands Project 

2000). The area is also under the influence of the SMO, the Rocky

Mountains, and the Chihuahuan and Sonoran deserts (Ponce-Guevara 

and others 2005).

The SSL in characterized by a dry temperature with summer rains. 

The average temperature in the summer is between 18 oC and 7 oC in 

winter (INEGI 1973). The average annual precipitation ranges from 

450 mm (~1500 m) to 700 mm (~2500 m; Íñiguez et al. 2005). July 

receives the highest rainfall (86 to 106 mm) and the lowest precipitation occurs in May (2.7 to 3.2 mm; INEGI, 1973). The elevation of 

the locations ranges from 1,500 to 1800 m (Rodríguez-Martínez and 

others 2008). The hydrogeology is characterized by the presence of 

theCajonBonitoRiver and several artificialwater bodies and puddles

(Rodríguez-Martínez and others 2008).

The dominant vegetation consists of grasslands and scrub. The representative species are the palmilla (Nolina microcarpa), Engelmann ́s 

nopal (Opuntia pheacantha), maguey (Agave palmeri) and sotol 

(Dasylirion wheeleri). There are also areas with gallery forests 

dominated by aspen (Populus sp.) and in some areas juniper-oak or 

huizachal-oak associations can be found (Rodríguez-Martínez and 

others 2008).

Methods

Population Size

To estimate the black bear population size in SSL, we placed camera traps (WildView5®) in two sites from July to August 2010—23 

camera traps in El Pinito Ranch and 29 camera tramps in Los Ojos 

Ranch. The camera traps were placed on roads identified as fauna

passages and were separated about 1 km. They were placed with S-N 

orientation. As bait, we placed in front of each camera a combination 

ofsardinesin tomato sauce, amixture of oat-corn, and finally a vanilla

extract sprayed with an atomizer into the above mixture. The camera 

traps were programmed to record the date and time (military format) 

and to take three successive photographic events every minute if they 

were activated.

	 Black bears in the photographic records were identified at the

individual level through a comparative analysis of opinions where 

five observers discriminated individuals between the images obtained

independently. Unusable photographs to distinguish individuals were 

eliminated, producing a consensus by which was established the 

average number of individuals; this approach has been used successfully in other studies (Kelly and others 2008). With these results, we 

constructed a capture-recapture history for each individual generating 

Figure 1—Study area and records of Black Bear in Sierra de San Luis, Sonora, México.
USDA Forest Service Proceedings RMRS-P-67. 2013 265

Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States Lara-Díaz and others

a database of presence (1) or absence (0) according to the sampling 

days.

The abundance was estimated with the MARK 6.0 program (White 

2008), using the Capture tool, considering the assumptions of a closed 

population through the appropriate model of capture probability. We 

calculated the effective sample area generating a buffer surrounding 

each camera, using the average home range for black bear in SSL 

(32.33 km2

; López-González and Lara-Díaz 2010), in the ArcMap 

ver. 9.3 program (ESRI 2008), with the “buffer” tool, and avoiding 

overlapping areas. The final area was calculated using the extension

Xtools Pro (ver. 5.3) for ArcMap. For black bear density, the abundance parameter was divided by the effective sampling area, being 

reported as number of individuals per km2

. Finally, the density data 

(individuals/km2

) was extrapolated to the SSL area.

Occupancy and Connectivity

Black bear records obtained during the study (including photographs, 

feces, capture, telemetry, traces) were used to generate the occupancy 

probability of the species in the study area, using the PRESENCE 

ver. 3.0 program (Hines 2006) and following the protocol established 

by Donovan and Hines (2007), which considers a single species, in 

a season of sampling, with covariates associated to the sites.

We considered an area of 26,307 km2 (fig. 2) for calculating the

occupancy probability. With the help to GIS, we divided the area into 

a grid with sampling units of 1 km each. Each sampling unit visited, 

where we placed a camera trap or searched for traces, was associated 

to black bear presence or absence. Sampling units that were not visited 

during the monitoring events were listed with a dash (-) because it was 

unknown if the species was present or not. Thus, the detection history 

of the species was generated. Additionally, we generated a matrix of 

covariates associated with each sample unit: elevation (USGS 1993), 

Normalized Difference Vegetation Index (NDVI; Earth Observatory 

2010), roughness (USGS 2001) and slope (USGS 2001), all of them 

important variables for black bear presence (Moreno 2008).

To associate the covariates of elevation, roughness, and slope to 

sample units, we used the ArcView 3.2 program (ESRI 1999) where 

each covariate is represented by a raster layer, and they were clipped 

to the study area through the Spatial Analyst extension (ver. 1.1). The 

information of each layer was added to the grid of the study area through 

the Get Grid Value extension 2 (Davies 2000). Moreover, the NDVI 

values for each of the sample units was generated by a LANDSAT 

satellite image using Image Analysis extension (Pinedo and Pinedo 

2006) for Arcview 3.2 program (ESRI 1999). The covariates were 

normalized to be introduced to the program (Mackenzie and others 

2003).

Based on the black bear detection matrix and the covariates matrix, 

we generated probability models with the PRESENCE program (Hines 

2006). Each model considered the occupancy probability and detection 

probability associated with one or more covariates, also considering 

Figure 2—Occupancy probability of black bear (Ursus americanus) in the border region of Sonora and Chihuahua, México.
266 USDA Forest Service Proceedings RMRS-P-67. 2013

Lara-Díaz and others Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States

a constant covariate and a time-dependent covariate (Donovan and 

Hines 2007). Thus, we generated 1,056 models, given by the possible 

combinations among covariates. The best model to explain the occupancy and detection probabilities was selected through the lower 

Akaike Information Criteria (AIC ) (King and others 2007).

Once we obtained the best model and given the occupancy probabilities per sample unit of 1 km2

 (obtained by the program), these 

probabilities were associated with a space-map to get a representative 

image of the occupancy probability distribution of black bears in the 

study area. This was achieved by using the ArcMap 9.3 program (ESRI. 

2008) considering areas with the highest occupancy probability and 

greater coverage, which are the more likely for having connectivity 

between areas.

Results

Population Size

We obtained a total 30 different individuals in El Pinito Ranch and 

18 different individuals in Los Ojos Ranch with a sampling effort 

of 4,413 camera-days. The abundance was estimated at 34 and 24 

individuals, respectively. El Pinito Ranch had a higher density (0.322 

individual/km2

) compared to Los Ojos Ranch (0.161 individual/km2; 

table 1).

The SSL has an approximate area of 2,300 km2 including 50 km 

south of the international border. Extrapolating the estimated average 

density obtained through our study, we believe there may be a black 

bear population of about 550 individuals; however, this population 

estimate may be reduced depending on management and predator 

control being carried out by different ranches in the area.

Occupancy and Connectivity

In the period from June 23 to November 3, 2010, we obtained a 

total of 209 records that documented the black bear presence. These 

records included 149 feces, six sets of traces, 41 camera-trap stations, 

four captures and 10 telemetry locations (fig. 1).

The best occupancy probability model (AIC = 390.37) was the 

one associated with the occupancy probability (psi) to NDVI, and the 

detection probability (p) to time. According to this model, the most 

likely area where black bears can be found is SSL, whose connectivity 

to the United States occurs in two main points from Puerto de San 

Luis (Mexico) to Animas Mountains (United States) and from the 

west of the SSL to Peloncillo Mountains (United States), eventually 

connecting to the Chiricahua Mountains (fig. 2). The region of the

Sierra de Enmedio, in the Chihuahua State, hardly connects to the 

north with appropriate areas for the black bear, making it likely that 

individuals migrate north along the Sierra de San Luis. The other 

connectivity passage to the United States for black bear populations 

in the Sky Islands is the north part of the Sierra de Ajos (Mexico) 

through the Sierra San José (Mexico) and connecting with Mule 

Mountains (United States; fig. 2).

Discussion and Conclusions

The black bear population in Sierra de San Luis seems to be the 

largest in Sonora and Chihuahua. The abundance and density data 

obtained, as well as indirect records, have been higher compared with 

mountain ranges as Ajos, El Tigre and La Madera in México, where 

the densities have been estimated to be less than 0.05 individuals/

km2

 (Lara-Díaz 2010).

The monitoring carried out during the study period shows a potential increase in black bear population in SSL relative to previous 

years (Sierra Corona and others 2005; Rodríguez and others 2008). 

This increase allows this population to be a source of individuals to 

other populations in the region. This result may be the outcome of 

multiple management strategies in the region, with areas dedicated 

to conservation with no hunting and a limited predator control. However, we must consider that some ranches still continue to implement 

measures against predators, and, therefore, the habitat for the black 

bears may be less suitable. Consequently the population size in SSL 

may be smaller.

In terms of connectivity, black bears should not have restrictions 

to continue crossing the border on the existing vehicle barriers. We 

believe that asking for modificationsto the construction structure and

the placing of it would be unrealistic. However, we can recommend 

to remove or modify the barbed wire that makes up the lower portion 

of the barrier and to raise the final strand; these simple actions can

facilitate transition of young or juvenile black bears. Adults seem to 

have no problem crossing because they can easily climb the vehicle 

barriers.

	 It is important to reduce people and vehicular traffic in the areas

identified as priority for possible connectivity of black bear populations from the two countries. If possible, increasing remote monitoring 

techniques (cameras and unmanned aerial vehicles) by the border 

agents would help the bears to continue crossing in these priority 

areas. Likewise, implementation of crossing points for wildlife with 

the expansion of Federal Highway No. 2 should reduce the number 

of collisions and, thus, would increase the probability that a black 

bear makes it to the border and across it. If black bear populations 

are to survive in the Sky Islands of the Northwest of Mexico, it is 

essential to maintain the connectivity sites from SSL and Sierra de 

Ajos to the United States.

Although the black bear population in SSL is reproducing and has 

recently increased, individuals of this population are not sufficient

to reevaluate the endangered category in the region. Black bears 

are still widely pursued under the belief that they seriously affect 

production activities in the area. Finally, continuous monitoring of 

black bears populations in the Sky Islands is essential to assess the 

species status in the medium and long-term, and it is also essential 

to develop strategies for conservation and appropriate management.

Table 1—Abundance and density of black bear (Ursus americanus) at two study sites in the Sierra de 

San Luis, Sonora, México.

Abundance Interval Effective Sampling Density

Site	 (individuals)	 S. E.	 (95% confidence)	 area (km2

) (Ind./km2) S. E.t

El Pinito 34 3.98 31 a 50 individuals 105.600 0.322 0.387

Los Ojos 24 4.51 20 a 40 individuals 149.340 0.161 0.369
USDA Forest Service Proceedings RMRS-P-67. 2013 267

Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States Lara-Díaz and others

Acknowledgments

We are in debt to the Instituto Nacional de Ecología and to the Program for the Improvement of Teachers (PROMEP) who provided the 

funds to buy the required materials for the research projects - “Diagnóstico poblacional del oso negro (Ursus americanus) en las serranías 

de los estados de Sonora y Chihuahua y sus posibles afectaciones 

por el muro fronterizo” and “FNB-2008-04, Respuesta poblacional, 

espacio-temporal y fisiológica del oso negro (Ursus americanus) a 

un gradiente latitudinal como prueba de una hipótesis demográfica,”

respectively. We are also in debt to the Fundación Cuenca los Ojos, 

owners and staff of the monitored properties for the support and the 

facilities provided. We also need to thank the U.S. Border Patrol in 

Douglas, Arizona. We thank Eugenia Espinosa, Diana Zamora and 

Daniel Ávila for their support in the monitoring, especially to Sandra 

Lanham for support during aerial telemetry. Finally, thanks to Dr. 

Christian A. Delfín Alfonso and Dr. A. Vladimir Cachón Guillen for 

their valuable reviews this manuscript.

References

Baker, Rollin H. 1956. Mammals of Coahuila. Publications of the Museum 

of Natural History, University of Kansas 9:127-335.

Baker, Rollin H.; J.K. Greer. 1962. Mammals of the Mexican state of Durango. 

Michigan State University Museum Publ., Biol. Ser. 2.

CITES (Convención sobre el Comercio Internacional de Especies Amenazadas 

de Fauna y Flora Silvestres). 2009. Apéndices. Online: http://www.cites.

org/esp/index.shtml. 

CONANP (Comisión Nacional de Áreas Naturales Protegidas). 2011. Programa 

de conservación de especies riesgo. PROCER. 2007-20012. Online: http://

www.conanp.gob.mx/contenido/pdf/PROCER-TdeR-INT.pdf

Costello, Christine, M.; D.E. Jones; G. Hammond; [and others]. 2001. A study 

of black bear ecology in New Mexico with models for population dynamics 

and habitat suitability. Federal Aid in Wildlife Restoration Project W-131-R. 

Santa Fe, New Mexico, USA.

Crooks, Kevin R. 2002. Relative sensitivities of mammalian carnivores to 

habitat fragmentation. Conservation Biology. 16:488–502.

Davies, John. 2000. Get Grid VAlue Extension 2. ArcView GIS. Online: 

www.ersi.com.

Delfín-Alfonso, Christian. A.; C. A. López-González; N. E. Lara-Díaz. 2011. El 

oso negro americano en el Noroeste de México: Recuperación de Registros 

de ocurrencia. Acta Zoológica Mexicana (n.s.). 27:777-801.

Delfín-Alfonso, Christian A.; C. A. López-González; M. Equihua-Zamora. (In 

press). Potential distribution of American black bears in northwest Mexico 

and implications for their conservation. Ursus 23 (1).000-000

Doan-Crider, Diana. L. 1995. Population characteristics and home range 

dynamics of a black bear population in northern Coahuila, México. M.S. 

thesis, Texas A&M University, Texas, USA.

Donovan, T. M.; J. Hines. 2007. Exercises in occupancy modeling and estimation. Online: http://www.uvm.edu/envnr/vtcfwru/spreadsheets/occupancy/

occupancy.htm

Doan-Crider, Diana. L.; E. C. Hellgren. 1996. Population characteristics and 

winter ecology of black bears in Coahuila, Mexico. Journal of Wildlife 

Management. 60:398-407.

Duffy, J. Emmett. 2002. Biodiversity and ecosystem function: the consumer 

connection. Oikos. 99:201-219.

Earth Observatory. 2010. EOS Project Science Office. NASAGoddard Space

Flight Center. Online: http://earthobservatory.nasa.gov/Features/MeasuringVegetation/measuring_vegetation_1.php.

ESRI (Environmental Systems Research Institute) .1999. ArcView. Version 3.2. 

Environmental Systems Research Institute, Inc. Redlands, California, USA.

ESRI (Environmental Systems Research Institute). 2008. ArcGIS. Arc View 

GIS, Ver. 9.3. Windows. USA.

Gallo, Juan P.; F. Garza. 2002. Estado Actual del Oso Negro (Ursus americanus machetes) en el Estado de Sonora, México. VI Congreso Nacional 

de Mastozoología, Departamento de Ecología y Comportamiento Animal, 

Veracruz, Veracruz.

Hall, Eugene R. 1981. The mammals of North America. Vol. 2. John Wiley 

& Sons, New York. 1175 pp.

Hines, James E. 2006. PRESENCE 2-Software to estimate patch occupancy 

and related parameters. USGS-PWRC. Online: http://www.mbr-pwrc.usgs.

gov/software/presence.html.

INEGI (Instituto Nacional de Estadística y Geografía). 1973. Cartas topográficas Estado de Sonora, Escala 1:250,000.INEGI.Aguascalientes, México.

Íñiguez, José M.; J. L. Ganey; P. J. Daugherty; J. D. Bailey. 2005. Using cluster 

analysis and a clasification and regression tree model to developed cover

types in the Sky Islands of southeastern Arizona. In: Gottfried, G. J., B. S. 

Gebow, L.G. Eskew, C. B. Edminster (comp.). 2005. Connecting Mountain 

Islands and Deserts Seas: biodiversity and Management of the Madrean 

Archipelago II. Proc. RMRS-P-36. Fort Collins, CO: U. S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station: 195-200

Kelly, Marcella J.; A. J. Noss; M. S. Di Bitetti; [and others]. 2008. Estimating puma densities from camera trapping across three study sites: Bolivia, 

Argentina, and Belize. Journal of Mammalogy. 89:408–418.

King, Carolyn M.; R. M. McDonald; R. D. Martin; [and others]. 2007. 

Continuous monitoring of predator control operations at landscape scale. 

Ecological Management and Restoration. 8:133-139.

Lara-Díaz, Nalleli E. 2010. La Comunidad de mamíferos previa a la reintroducción de Canis lupus baileyi en sonora, México. Tesis Maestría. Facultad 

de Ciencias Naturales. Universidad Autónoma de Querétaro. Querétaro, 

México. 123 pp.

Leopold, Aldo S. 1959. Wildlife of Mexico. University of California Press, 

Berkeley. 608 pp.

List, Rurik. 2007. Los impactos del muro fronterizo sobre los mamíferos silvestres. In: Córdova, A. y C. A. de la Parra (coords). Una barrera a nuestro 

ambiente compartido. El muro fronterizo entre México y Estados Unidos. 

Secretaría del Medio Ambiente y Recursos Naturales. Instituto Nacional 

de Ecología. El Colegio de la Frontera Norte. Consorcio de Investigación 

y Política Ambiental del Suroeste. México. 214 pp.

López-González, Carlos A.; N. E. Lara-Díaz. 2010. Diagnóstico poblacional 

del oso negro (Ursus americanus) en las serranías de los estados de Sonora 

y Chihuahua y sus posibles afectaciones por el muro fronterizo. Informe 

Final. Instituto Nacional de Ecología-Universidad Autónoma de Querétaro. 

México. 94 pp.

MacKenzie Darryl I.; J. D. Nichols; J. E. Hines; M. G. Knutson; A. D. Franklin. 

2003. Estimating site occupancy, colonization and local extinction when a 

species is detected imperfectly. Ecology. 84:2200–2207.

Medellin, Rodrigo A.; C. Manterola; M. Valdéz; D. G. Hewitt; D. L. Doan 

Crider; T. E. Fulbright. 2005. History, ecology, and conservation of the 

pronghorn antelope, bighorn sheep, and black bear in Mexico. In: Biodiversity, Ecosystems, and Conservation in Northern Mexico, J. E. Cartron, 

G. Ceballos, and R. S. Felger (eds). New York: Oxford University Press: 

387-404.

Meffe, Gary K.; C. R. Carroll. 1997. Principles of Conservation Biology, 2nd 

Edition. Sinauer Associates, Sunderland, MA.

Moreno, Claudia. 2008. Ecología Conductual del Oso Negro (Ursus americanus) en la Sierra Madre Occidental. Tesis de Maestría. Instituto de Ecología, 

A. C. Xalapa, Veracruz. 85 pp.

Onorato, Dave P.; E.C. Hellgren; F.S. Mitchell; J.R. Skiles, Jr. 2003. Home 

range and habitat use of American black bears on a desert montane island 

in Texas. Ursus. 14(2): 120-129

Pelton, M.R. 1982. Black bear. In: J.A. Chapman; G.A. Feldhamer (eds). Wild 

mammals of North America: biology, management, and economics. John 

Hopkins University Press. Baltimore and London: 504-514.

Pinedo, A. Carmelo; A. A. Pinedo. 2006. SIG: Sistemas de Información 

Geográfica para el monitoreo y evaluación de los recursos forestales.

Facultad de Zootecnia, UACH.

Ponce-Guevara, Eduardo, K. Pelz-Serrano; C. A. López-González. 2005. 

Coyote Abundance in Relation to Habitat Characteristics in Sierra San 

Luis, Sonora, México. In: Gottfried, G. J., B. S. Gebow, L.G. Eskew, C. 

B. Edminster (comp.). 2005. Connecting Mountain Islands and Deserts 

Seas: biodiversity and Management of the Madrean Archipelago II. Proc. 
268 USDA Forest Service Proceedings RMRS-P-67. 2013

Lara-Díaz and others Black Bear Population and Connectivity in the Sky Islands of Mexico and the United States

RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station:337-340.

Public Law 109-367-OCT.26. 2006. Secure Fence Act of 2006. To establish 

operational control over to international land and maritime norders of 

the United States. Congresional record Vol. 152. Weekly compilation of 

presidential documents, Vol. 42.

Robbins, C. T.; C. C. Schwartz; L. A. Felicetti. 2004. Nutritional ecology of 

ursids: a review of newer methods and management implications. Ursus. 

15:161–171. 

Rodríguez-Martínez, Adriana; C. Moreno-Arzate, R. González-Sierra; C. A. 

López-González. 2008. Uso de hábitat, hábitos alimentarios y estructura 

poblacional del oso negro (Ursus americanus) en la Sierra Madre Occidental 

de México. In: Avances en el Estudio de los Mamíferos de México II. Lorenzo, C. y Espinoza, E. (eds.) Asociación Mexicana de Mastozoología A. C.

Rogers, Lynn L. 1993. The role of habitat quality in the natural regulation of 

black bear populations. Proc. 4th Western Black Bear Workshop: 95-102; 

Yosemite National Park, CA. Tech. Rep.NPS/NRWR/NRTR-93/12.

SEMARNAP-INE. 1999. Proyecto para la conservación y manejo de oso 

negro (Ursus americanus) en México. 112 pp.

SEMARNAT. 2010. Norma OficialMexicana NOM-059-SEMARNAT-2010,

Protección ambiental-Especies nativas de México de flora y fauna silvestres-Categorías de riesgo especificaciones para su inclusión, exclusión o

cambio-lista de especies en riesgo. 

Sierra-Corona, Rodrigo; I. A. Sáyago Vázquez; M. C. Silva Hurtado; C. A. 

López González. 2005. Black bear abundance, habitat use, and food habits 

in the Sierra San Luis, Sonora, México . In: Gottfried, G. J., B. S. Gebow, 

L.G. Eskew, C. B. Edminster (comp.). 2005. Connecting Mountain Islands 

and Deserts Seas: biodiversity and Management of the Madrean Archipelago 

II. Proc. RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station.

The Wildlands Project. 2000. Sky Islands wildlands network, conservation 

plan: An extraordinary plan for a place beyond compare. Tucson, AZ: The 

Wildlands Project. 

USGS (U. S. Geological Survey). 1993. Digital elevations models, data user 

guide 5: Reston, Virginia. U.S. Geological Survey. 48 pp.

USGS (US Geological Survey). 2001. HYDRO 1k, elevation derivative 

database. Sioux Falls, SD: U.S. Geological Survey. Online: http://www.

edcdaac.usgs.gov/gtopo30/hydro.

Varas, A. Cora. 2010. Conservation genetics of black bears in Arizona and 

Northern México. Ph.D. Thesis. Tucson: University of Arizona.

White C. Gary. 2008. Mark and Recapture Parameter Estimation. Version 6.0. 

Colorado State University. Online: http://welcome.warnercnr.colostate.

edu/~gwhite/mark/mark.htm. 

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 269

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Inventory of Terrestrial Mammals in the Rincon Mountains 

Using Camera Traps

Don E. Swann and Nic Perkins

Saguaro National Park, Tucson, Arizona

Abstract— The Sky Island region of the southwestern United States and northwestern Mexico is well-known 

for its diversity of mammals, including endemic species and species representing several different biogeographic provinces. Camera trap studies have provided important insight into mammalian distribution and 

diversity in the Sky Islands in recent years, but few studies have attempted systematic inventories of one 

or more mountain ranges with a repeatable, randomized study design. We surveyed medium and large 

terrestrial mammals of the Rincon Mountains within Saguaro National Park, and compared the results with 

previous surveys of the Rincons. We sampled in random locations in four elevational strata from May 2011 

through April 2012. We detected 23 native species of mammals and estimated species richness to be 24.8 

species. We failed to detect four native species documented by other methods during 1999-2012, as well 

as five species (bighorn sheep, grizzly bear, jaguar, gray wolf, and North American porcupine) documented 

during 1900-1999 that may be extirpated from the Rincons. Advances in camera trap technology, as well 

an expanding use of this technology by educators and the public, suggest this method has the potential to 

be a cost-effective and reliable method for both inventory and long-term monitoring of terrestrial mammals 

of Sky Island region. 

Introduction

The Sky Island region of the southwestern United States and 

northwestern Mexico is known for exceptional biological diversity, 

including species from the Sierra Madre, Rocky Mountain, Sonoran 

Desert, and Chihuahuan Desert biogeographic provinces (Swann 

and others 2005) and as many as 15 mammals on the edges of their 

biogeographic range (Warshall 1995). Southeastern Arizona, in particular, has been characterized as possessing the greatest mammalian 

diversity in North America north of Mexico (Turner and others 1995) 

including species with more northern affinities such as black bears 

(Ursus americanus) and tropical species such as ocelots (Leopardus 

pardalis), jaguars (Panthera onca), coatis (Nasua narica), and others. 

Several high profile species have been extirpated, including grizzly 

bear (Ursus arctos) and Mexican gray wolf (Canis lupus), while 

others such as jaguars, ocelot, bighorn sheep (Ovis canadensis), and 

pronghorn (Antilocapra americana) are considered highly vulnerable.

Species inventories, or verified lists of species that occur in a 

particular area, have value for many reasons (Sidner and Stone 

2005) including studies of biogeography (Newmark 1995; Lomolino 

and others 1989), determining environmental change (McDonald 

and Brown 1992), and determining status of sensitive species. In 

addition, repeated inventories may provide data for monitoring 

long-term changes in plant and animal communities if conducted 

using a randomized, repeatable study design that allows estimates 

to be made of measures such as native species richness (the number 

of native species that occur in an area). Loss of biodiversity is one 

of the world’s greatest environmental concerns (Wilson 1988) and 

detecting changes in diversity can be accomplished through repeated 

inventories conducted in a framework for drawing inference about 

variation in species richness over time (Nichols and others 2011).

Because of the significance of the Sky Island region, taxonomic 

inventories of its individual mountain ranges have been conducted 

for over a century (Swarth 1904) and remain a principal focus of 

biologists to this day. Recent inventories of reptiles (Flesch and others 2010), vascular plants (Van Devander and Reina 2005; Bowers 

and McLaughlin 1996), and other taxonomic groups exist for many 

prominent Sky Islands but inventories of mammals have generally 

lagged behind (Koprowski and others 2005). Although older surveys 

exist for selected ranges (Cahalane 1939; Hoffmeister and Goodpasture 

1954; Lange 1960) mammals as a group are generally less well-studied 

and a recent paper (Koprowski and others 2005) identified a “dearth 

of data” on the mammals of the region. In addition, although medium 

and large mammals have a higher profile with the public than smaller 

mammals and bats, past inventories (Hoffmeister and Goodpasture 

1954) have tended to focus on the latter groups because more of 

those species can be easily captured using a single method such as 

live-trapping and mist-netting. Inventory methods for larger mammals 

include collection of specimens by trapping and shooting, but these 

techniques are generally discouraged today. Other methods include 

use of sightings, tracks, and other sign, but these approaches have the 

disadvantage of not always being verifiable by other researchers. In 

addition, the probability of detection varies widely among species, 

habitats, and time. Many mammals are very elusive, rare, or both and 

may be present at a site but not detected during even a multi-year 

study (Tobler and others 2008).
270 USDA Forest Service Proceedings RMRS-P-67. 2013

Swann and Perkins Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps

Infrared-triggered cameras, also called camera-traps, represent an 

important newer technology in mammal studies and are now used 

widely to study animal behavior (Bridges and Noss 2011), estimate 

abundance (Karanth and others 2011), and inventory mammals and 

monitor communities (Tobler and others 2008; O’Brien and others 

2011). Camera-traps have a great advantage over observational studies in that they provide a record of each species, similar to a voucher 

specimen, which can be viewed by independent observers. They 

are relatively inexpensive to operate and can be left alone to gather 

data for long periods. Most importantly, camera-traps provide data 

that can be used to estimate the detection probability of individual 

species in different habitats, camera settings, time periods, and other 

factors that are known to influence detectability (O’Connell and others 

2006; O’Brien and others 2011). In recent years, camera-traps have 

made a significant contribution to our knowledge of jaguars, ocelots, 

and other tropical cats of the Sky Island region (McCain and Childs 

2008) while providing important information for conservation and 

management of mammals in small reserves (Crooks and others 2008; 

Swann and others 2010). However, most of these studies were in a 

relatively small area over a short time-frame.

We conducted an inventory of medium and large mammals in both 

districts of Saguaro National Park from May 2011 through June 2012. 

Our goals were to use a randomized study design to collect photos of 

all mammals that could be detected by our cameras, and to use these 

data to estimate species richness, as well as detectability and percent 

occupancy for common species. In addition to providing baseline 

monitoring data, we wanted to compare our results with a similar 

study during 1999-2005 (Swann and Powell 2006) and historic data 

(Davis and Sidner 1992; Swann 2011) to determine if species richness or distribution has changed significantly during the past decade 

and past century. Finally, we hoped to use photographs to interpret 

wildlife in the park through social media outlets while also providing opportunities for service learning and citizen science. Here, we 

present partial data (May 2011-March 2012) for the park’s Rincon 

Mountain District.

Study Site

Saguaro National Park is located in Pima County, Arizona, and 

consists of two disjunct districts separated by the city of Tucson 

(fig. 1). The park includes 37,005 ha, of which 28,694 ha (78%) are 

designated Wilderness. The Rincon Mountain District covers a large 

portion of the Rincon Mountains, a northern Sky Island range adjacent 

and biologically connected to the Santa Catalina Mountains. The 

portion of the Rincon Mountains within the park ranges in elevation 

from 814-2,614 m. Biotic communities include Sonoran desertscrub, 

semi-desert grassland, oak savannah, oak pine-oak forest and woodland, and mixed conifer forest. The Rincon Mountains also contain 

riparian forest and riparian woodland in canyon bottoms, scattered 

wet and dry meadows at higher elevations, and perennial springs and 

spring-fed rock pools (tinajas).

Methods 

We used a stratified random design with four elevational strata: 

814-914 m (hereafter “low elevation”), 914-1,524m (“low-medium”), 

1,524-2,134 m (“medium-high”), and 2,134-2,641 m (“high elevation”) 

(fig. 2). Previous studies (Davis and Sidner 1992; Swann and Powell 

2006) suggested that mammal distribution in the Rincon Mountains 

varies among these strata, which may be associated with differences 

in soil type, vegetation, and climate. We divided the park into 1-km 

plots, deleting any that were >50% outside the park boundary and 

then randomly selected 15 plots (from a total of 60 plots). If a selected 

plot included two strata, it was assigned to the strata with the greatest 

amount of area. We established a minimum distance of 200 m between 

points.

Within in each plot we placed four Cuddeback Capture cameratraps (Non Typical Inc., Green Bay, Wisconsin). We set cameras at 

four random points generated for each plot but occasionally used an 

alternative random point if we determined that a point was not safe 

Figure 1—Location map for the Rincon Mountains, east of Tucson, Arizona.
USDA Forest Service Proceedings RMRS-P-67. 2013 271

Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps Swann and Perkins

for technicians to place a camera (e.g., on a cliff). We navigated to 

each point using a Garmin GPSmap 76 GPS unit (Garmin International, Olathe, Kansas) and recorded the actual camera location. We 

set cameras as close to the random point as possible (within 20 m), 

choosing the closest location that had a sufficiently open field of view 

to provide the potential to detect an animal if it was present. We made 

an effort to reduce technician bias in setting cameras through training, 

but we suspect that some bias on this small scale (e.g., setting cameras 

where a scat was present) crept in. We set cameras 30-60 cm above 

ground to detect both larger as well as relatively small mammals, 

although ground height typically varied within the detection zone. 

We targeted all terrestrial mammals that were squirrel-sized or larger 

(>80 g), except for nocturnal rodents and pocket gophers due to the 

difficulty in identifying them to species level. We did not provide bait 

or any attractants.

We left cameras in place for 6 weeks. We typically kept two plots 

active in each of the four strata (total of 32 cameras), although this 

routine was altered by an extreme fire danger closure of Saguaro 

National Park in June 2011, a major event (the BioBlitz) in October 

2011, and by occasional weather or logistical concerns. We checked 

plots at least once (but typically twice) during each 6-week interval to 

change batteries and verify that cameras were working properly. On 

each visit we replaced the camera memory card and also downloaded 

photos and viewed them in the field using a MP3 player.

We downloaded memory cards back in the office and placed photos 

in digital computer folders arranged by location and named by camera 

placement date and a unique camera ID. We examined each photo 

and identified all mammals to species. We independently confirmed 

all identifications using local experts and sent questionable identifications to taxonomic specialists. We entered all photo data in Excel 

spreadsheets using a separate spreadsheet for every camera location. 

We independently checked all data entry. To organize and analyze 

photo data, we created a file geodatabase with shapefiles for study 

plots, random points, active camera locations, and other information 

associated with each camera location. For statistical analysis, we 

imported a master Excel file that contained all photo records into a 

GIS (Arcmap) project and added attributes such as geographic coordinates and plot name, and then summarized the records by species to 

identify misspellings and other errors. We used ArcGIS-based models 

created for the project to build tables summarizing photo records of 

species by district and strata.

Figure 2—Study design for camera trap placement in Rincon Mountain District of Saguaro National Park. Four camera traps were placed randomly 

in each randomly located plot for 6 weeks, with all four strata being sampled from May 2011-March 2012. 
272 USDA Forest Service Proceedings RMRS-P-67. 2013

Swann and Perkins Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps

To calculate species richness of native mammals, we converted 

these tables to binary data, and used the species richness program 

EstimateS (http://viceroy.eeb.uconn.edu/EstimateS) that computes 

a variety of biodiversity functions, estimators, and indices for biotic 

sampling data (Colwell and others 2012). We used the Jack1 estimator in EstimateS; Jackknife estimators perform well when there is 

heterogeneity in capture probabilities among species as would be 

expected with camera-traps and past studies have indicated that Jack1 

provides the most reliable estimates for large number of camera 

nights (Tobler and others 2008). For purposes of species richness 

estimation, we assumed that Abert squirrels (Sciurus aberti), which 

were introduced into the Santa Catalina Mountains in the 1940s and 

first observed in the Rincons in 1959 (Davis and Sidner 1992), are 

now “native” and that all cottontails found above 2000 m are eastern 

cottontails (Sylvilagus floridanus), although a specimen is needed to 

confirm identification of this species.

Results

During May 2011 to March 2012, we sampled for approximately 

7,837 camera nights (tables 1 and 2). The cameras functioned during 

most but not all camera-checks and so the number of actual operational 

camera nights is fewer and unknown. We sampled at 173 points: 47 

in the low elevation strata, 47 low-medium, 40 in medium-high, 

and 39 in high in elevation. We obtained 1,943 photos containing 

medium-to-large mammals that could be identified to species, as well 

as many photos of nocturnal rodents, birds, reptiles, humans, and other 

non-target species. We detected a total of 23 native species during the 

study period, including 17 in low elevation, 18 in low-medium, 12 in 

medium-high, and 10 in high elevation (table 1). We estimated the 

native species richness of the Rincon Mountains to be 24.8 species 

(SD = 1.2).

The most commonly photographed species were desert cottontail 

(Sylvilagus audubonii), white-tailed deer (Odocoileus virginianus), 

and antelope jackrabbit (Lepus alleni) (fig. 3). Five species were 

photographed less than 6 times during the study period: mule deer 

(Odocoileus hemionus), Abert’s squirrel (Sciurus aberti), round-tailed 

ground squirrel (Spermophilus tereticaudus), Western spotted skunk 

(Spilogale gracilis), and American badger (Taxidea taxus). We did 

not detect any species that had not been previously documented for 

the Rincon Mountains, although we did photograph one species 

(American badger) that had not been previously documented by 

camera-traps. We failed to detect two species (Virginia opossum 

[Didelphis virginiana] and Northern raccoon [Procyon lotor] that were 

detected in the Rincon Mountains by non-randomly placed cameras 

during the same study period (Saguaro National Park, unpublished 

data). We also failed to detect one species (cliff chipmunk [Tamias 

dorsalis]) that was confirmed by numerous reliable observations 

and one species (Arizona gray squirrel [Sciurus arizonae]) that was 

detected during the inventories in 1999-2005. We did not detect five 

species, including grizzly bear, jaguar, gray wolf, bighorn sheep, and 

North American porcupine (Erethizon dorsalis) that were confirmed 

either by specimen or very reliable historical record to be present in 

the Rincon Mountains at some time during the period between 1900 

and 1999 (Swann 2011). We did not detect free-ranging, non-native 

domestic dogs (Canis familiaris), domestic cat (Felis catus), or cattle 

(Bos taurus), which are known to occur in the Rincon Mountains at 

times (Swann 2011) but probably not in established populations.

Discussion

Our study demonstrates some advantages and limitations of using 

infrared-triggered camera-traps to inventory and monitor medium 

and large mammals, where we define the term inventory as the development of a species list for an area and monitoring as a method to 

determine changes in a measure such as species richness over time. 

Camera-traps have some major advantages over traditional inventory 

and monitoring methods for larger mammals. These methods include 

kill-trapping and hunting which provide reliable identification but 

raise ethical issues and have the potential to impact populations; livetrapping, which provides reliable identification but is expensive and 

has safety issues for animals and researchers; visual identification of 

scat and sign, which is non-invasive but the results cannot be verified 

to prevent false identification; genetic identification of scat and hair, 

which is growing in its potential as a non-invasive, reliable method, 

but can be expensive and incomplete due to DNA degradation of scat 

in a field environment (Piggot and Taylor 2003); observation records, 

which are inexpensive to gather but produce results that cannot be 

independently verified (Giordano and others 2011); and collection 

of voucher specimens of roadkills and other dead animals, as well as 

collection of vouchers photographs by citizen scientists, which are 

inexpensive and verifiable but require a very long effort in time due 

to species avoidance of humans and raise complicated study design 

issues.

One advantage of camera-traps for inventory and monitoring is 

that the results can be independently verified. The results of some 

classic papers in conservation biology (Newmark 1995) have been 

questioned due to their use of unverified observational records (Parks 

and Harcourt 2002). In addition, camera-traps can be easily deployed 

in a repeatable study design. Our study demonstrates that cameras 

placed at random locations, as opposed to only at water holes, game 

trails, and other locations known to attract certain species, will detect 

large numbers of species present in an area even if no bait or other 

attractants are used. Baiting camera-traps for a study of multiple 

Table 1—Summary data on effort and results of camera trap study in the Rincon Mountains, May 2011-March 2012. 

 Area of No. of No. of Species

Elevation sampling camera camera No. of No. of richness

Strata range unit points nights photos species estimate

(m) (ha) (SD)

1 814-914 5,066 47 2,015 1,216 17 19.9 (1.9)

2 914-1,524 27,714 47 1,876 302 18 18.9 (0.9)

3 1,524-2,134 27,040 40 1,900 218 12 14.7 (1.9)

4 2,134-2,641 7,469 39 2,046 207 10 11.8 (1.2)

Total 67,289 173 7,837 1,943 57 24.8 (1.2)
USDA Forest Service Proceedings RMRS-P-67. 2013 273

Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps Swann and Perkins

species raises the potential for biasing results in studies of mammal 

communities (Kays and others 2011). We obtained a large number 

of photos of animals without using bait and without targeting water 

holes, game trails, or other macro-habitat features.

A major issue in inventories as opposed to monitoring (see below) 

that cannot be resolved by camera-traps is that of incomplete detectability or the failure to detect species when they are indeed present. 

Camera-traps are like live-traps and other methods in that they vary 

in their ability to detect species, especially when they are set in a 

standard manner at random points. We expect that some individual 

animals may notice and avoid cameras, some may be attracted to them, 

and some may not be detected by cameras due to their small size or 
274 USDA Forest Service Proceedings RMRS-P-67. 2013

Swann and Perkins Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps

quick movements. Despite a large effort, we did not detect several 

species that were photographed in the Rincon Mountins during the 

past decade (Swann 2011). We suspect that these species were present in the park for at least part of our study but went undetected due 

to their relative rarity, small size (cliff chipmunk), or specific habitat 

needs (e.g., Arizona gray squirrels who occur mostly in trees). We 

believe we would detect more species with increased sampling effort, 

although this effort may need to be very large. Therefore, for a mammal inventory of an area to be as complete as possible, we recommend 

that some cameras be set in targeted habitats. For example, setting 

cameras at the base of oak trees to detect Arizona gray squirrels, a 

rare habitat specialist, may be the most efficient way to detect this 

species in the Rincon Mountains.

For monitoring, the problem of incomplete detectability is common to all wildlife monitoring studies and can be overcome through 

a design that quantifies the probability of detection and estimates the 

measure of interest. Species richness, particularly of native species, 

is often used to determine changes in biological diversity, and can 

be estimated from camera trap studies using a number of models 

and approaches (Tobler and others 2008; O’Brien and others 2011), 

including a fairly simple approach using program EstimateS, as we 

have done in this paper. Our estimated native species richness of 24.8 

+1.2 for the Rincon Mountains seems to be reasonable, given that 27 

native species have been detected by all methods since 1999.

As one the most northern and western of the Sky Islands, the Rincon 

Mountains might be expected to have a reduced number of mammals 

of tropical origin compared to more southern and eastern ranges such 

as the Chiricahua Mountains, Huachuca Mountains, and others. Compared to other Sky Islands, the Rincon Mountains have fewer reptile 

and amphibian species, and lack at least seven montane species found 

even as close as the nearby Santa Rita Mountains (Swann and others 

2005). Medium and large mammals should be able to move among 

Sky Islands more easily than reptiles and some other taxa, and indeed 

a number of recent genetic studies have shown that larger mammals 

such as bears can be closely related among nearby sky islands (Varas 

and others 2005). We photographed opossums at non-random sites, 

a more southern species that appears to be expanding its range to the 

north (Babb and others 2004), but did not detect ocelots or jaguars. 

There are apparently no confirmed records of ocelots from the Rincons 

but at least four jaguars were confirmed (killed by hunters) in the 

Rincon Mountains during 1900-1932 (Swann 2011). Several other 

terrestrial mammals that occur in the area, such as kit fox (Vulpes 

macrotis), have never been documented in the Rincon Mountains.

At least five species that we did not detect are known to have occurred in the Rincon Mountains during the past century (Swann 2011). 

In addition to jaguar, these include grizzly bear (last record a hunter 

kill in the 1920s); Mexican gray wolf (last records from the 1950s); 

bighorn sheep (last records from the 1950s); and North American 

porcupine (last record a reliable observation in the 1990s). Of these 

five species, grizzly bear and Mexican gray wolf are extirpated from 

the region. Jaguars and bighorn sheep are probably extirpated from 

the Rincon Mountains, but both are wide-ranging species that could 

be expected to occur occasionally. Porcupines appear to be declining throughout Arizona, possibly due to the increase in predators, 

especially mountain lions (Brown and Babb 2009).

Management Implications

Despite the great interest in mammals of the Sky Islands, the need 

for regional monitoring and improvements in our ability to detect 

rare species such as jaguars, many Sky Islands still do not have 

basic inventories and the region does not have a current standardized monitoring plan. Camera-traps are an emerging technology that 

may provide opportunities to address these issues. Comprehensive 

monitoring need not be an expensive investment, due to volunteer 

efforts by groups such as the Sky Island Alliance (SIA), and because 

monitoring intervals can be widely separated in time, but it would 

require a coordinated effort. A great advantage of using camera-traps 

over other monitoring methods is that they provide interesting photos 

and videos that not only possess important scientific value but can be 

enjoyed by the public through a wide variety of social media. Photos 

and videos from our study have been made available through Facebook and a number of websites including Flickr, National Geographic 

Society, and Friends of Saguaro National Park.

Figure 3—Number of photos for each of the 23 native terrestrial mammal species (excluding nocturnal rodents and pocket gophers) photographed by 

camera traps in the Rincon Mountains during May 2011-March 2012. 
USDA Forest Service Proceedings RMRS-P-67. 2013 275

Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps Swann and Perkins

The project described in this paper was initiated and inspired by the 

2011 BioBlitz, a major biodiversity event that took place at Saguaro 

National Park in October 2011 and was supported by the National 

Geographic Society, Friends of Saguaro National Park, and many 

other organizations. The BioBlitz matched nearly 200 scientists and 

experienced naturalists in the field with more than 2,500 Tucson school 

children and adult volunteers to sample the park’s biodiversity. As part 

of the BioBlitz, we developed a program with SIA, the National Park 

Foundation, AmeriSchools College Preparatory Academy, and others 

that enabled middle and high-school students to track wildlife with 

expert SIA trackers and download wildlife photos from camera traps 

that had been previously set by high school volunteers. The program 

provided a hands-on wildlife experience, with students succeeding in 

capturing a number of species on film including mountain lions 

(fig. 4) and coatis. The growing use of camera-traps by both biologists and educators has great potential to engage the public directly 

in the conservation of Sky Island mammals while also providing data 

useful for long-term conservation.

Acknowledgments

This project could not have been accomplished without the support of the Friends of Saguaro National Park. We are grateful to the 

many interns and volunteers who regularly assisted in setting cameras 

particularly Mary Beth Benton, Brittany Bankovich, Dan Bell, Emma 

Fajardo, Katie Keck, Johnny Ortiz, Kris Ratzlaff, Rafael Rojas, 

Adam Springer, and Albi von Dach. We appreciate reviews by John 

Koprowski and Natasha Kline that greatly improved the paper.

References

Babb, R. D.; D. E. Brown; and J. L. Childs. 2004. On the status of the opossum 

(Didelphis virginiana) in Arizona. Journal of the Arizona-Nevada Academy 

of Science 36:120–126.

Bowers; J. E.; S. P. McLaughlin. 1996. Flora of the Huachuca Mountains, a 

botanically rich and historically significant Sky Island in Cochise County, 

Arizona. Journal of the Arizona-Nevada Academy of Science 29:66-107. 

Bridges, A. S.; A. J. Noss. Behavior and activity patterns. Pp 57-70 in

O’Connell, A. F.; J. D. Nichols; and K. U. Karanth, eds. Camera traps in 

wildlife ecology. Springer Publishing, Tokyo. 271 p. 

Brown, D. E.; R. D. Babb. 2009. Status of the Porcupine (Erithizon dorsatum) 

in Arizona, 2000–2007. Journal of Arizona-Nevada Academy of Science 

41:36-41. 

Cahalane, V. H. 1939. Mammals of the Chiricahua Mountains, Cochise County, 

Arizona. Journal of Mammalogy 20: 418-440. 

Colwell, R. K.; A. Chao; N. J. Gotelli; S.-Y. Lin; C. X. Mao; R. L. Chazdon; 

and J. T. Longino. 2012. Models and estimators linking individual-based and 

sample-based rarefaction, extrapolation, and comparison of assemblages. 

Journal of Plant Ecology 5:3-21.

Crooks, K. R.; M. Grigione; A. Scoville; and G. Scoville. 2008. Exploratory 

use of track and camera surveys of mammalian carnivores in the Peloncillo 

and Chiricahua mountains of southeastern Arizona. The Southwestern 

Naturalist 53: 510-517.

Figure 4—Mountain lion (Felis concolor) photographed by a camera trap in the Rincon Mountains for the BioBlitz educational 

program in October, 2011. 
276 USDA Forest Service Proceedings RMRS-P-67. 2013

Swann and Perkins Inventory of Terrestrial Mammals in the Rincon Mountains Using Camera Traps

Davis, R.; R. Sidner. 1992. Mammals of woodland and forest habitats in the 

Rincon Mountains of Saguaro National Monument, Arizona. Technical 

Report NPS/WRUA/NRTR-92/06 (CPSU/UA no. 47). National Park 

Service, Cooperative Park Studies Unit, School of Natural Resources, 

University of Arizona, Tucson. 

Flesch, A. D.; D. E. Swann; D. Turner; and B. F. Powell. 2010. Amphibians 

and reptiles of the Rincon Mountains, a Sky Island in southern Arizona. 

Southwestern Naturalist 55:240-253. 

Giordano, A. J.; R. Carrera; and W. Ballard. 2011. Assessing the credibility 

of Jaguarundi (Puma yagouaroundi) observations using diagnostic criteria 

and witness qualification. Human dimensions of wildlife 16:360-367. 

Hoffmeister, D. F.; W. W. Goodpaster. 1954. The mammals of the Huachuca 

Mountains, southeastern Arizona. Illinois Biological Monographs. 24:1-

152. Urbana, IL: University of Illinois Press.

Karanth, K. U.; J. D. Nichols; and N. S. Kumar. 2011. Estimating tiger abundance from camera trap data: field surveys and analytical issues. Pp 97-118 

in O’Connell, A. F.; J. D. Nichols; and K. U. Karanth, eds. Camera traps 

in wildlife ecology. Springer Publishing, Tokyo. 271 p. 

Kays, R.; S. Tilak; B. Kranstauber; P. A. Jansen; C. Carbone; M. Rowcliffe; 

T. Fountain; J. Eggert; and Z. He. 2011. Camera traps as sensor networks 

for monitoring animal communities. International Journal of Research and 

Reviews in Wireless Sensor Networks 2:19-29. 

Koprowski, J. L.; A. J. Edelman; B. S. Pasch; and D. C. Buecher. 2005. A dearth 

of data on the mammals of the Madrean Archipelago: what we think we 

know and what we actually do know. Pp in 412-415 in Gottfried, G. J.; S. B. 

Gebow; L. G. Eskew; and C. B. Edminster, eds. Connecting mountain islands 

and desert seas: biodiversity and management of the Madrean Archipelago 

II. Proceedings RMRS-P-36. Fort Collins, Colorado: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station, 631 p. 

Lange, K. I. 1960. Mammals of the Santa Catalina Mountains, Arizona. 

American Midland Naturalist 64: 436-458. 

Lomolino, M. V.; J. H. Brown; and R. Davis. 1989. Island biogeography of 

montane forest mammals in the American Southwest. Ecology. 70: 180-194.

McCain, E. B.; J. L. Childs. 2008. Evidence of resident jaguars (Panthera 

onca) in the southwestern United States and the implications for 

conservation. Journal of Mammalogy 89:1-10. 

McDonald, K. A.; J. H. Brown. 1992. Using montane mammals to model 

extinctions due to global change. Conservation Biology. 6: 409-415.

Newmark, W. D. 1995. Extinction of mammal populations in western American 

national parks. Conservation Biology 9:512–526.

Nichols, J. D.; K. U. Karanth; and A. F. O’Connell. 2011. Science, conservation, and camera traps. Pp 45-56 in O’Connell, A. F.; J. D. Nichols; and 

K. U. Karanth, eds. Camera traps in animal ecology. Springer Publishing, 

Tokyo. 271 pp. 

O’Brien, T. G.; M. F. Kinnaird; and H. T. Wibisono. 2011. Pp 233-252 in

O’Connell, A. F.; J. D. Nichols; and K.U. Karanth, eds. Camera traps in 

animal ecology. Springer Publishing, Tokyo. 271 pp. 

O’Connell, A. F.; L. L. Bailey. 2011. Inference for occupancy and occupancy 

dynamics. Pp 191-206 in O’Connell, A. F.; J. D. Nichols; and K. U. Karanth, 

eds. Camera traps in wildlife ecology. Springer Publishing, Tokyo. 271 pp. 

Parks, S. A.; A. H. Harcourt. 2002. Reserve size, local human density, and 

mammalian extinctions in U.S. protected areas. Conservation Biology 

16:800-808. 

Piggot, M. P.; A. C. Taylor. 2003. Remote collection of animal DNA and its 

application in conservation management and understanding the population 

biology of rare and cryptic species. Wildlife research 30:1-13. 

Sidner, R.; H. S. Stone. 2005. First records of two species of mammals in the 

Huachuca Mountains: results of ecological stewardship at Fort Huachuca. 

Pages 131–134 in Gottfried, G. J.; S. B. Gebow; L. G. Eskew; and C. B. 

Edminster, eds. Connecting mountain islands and desert seas: biodiversity 

and management of the Madrean Archipelago II. Proceedings RMRS-P-36. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station, 631 p. 

Swann, D. E. 2011. Mammals of Rincon Mountain District, Saguaro National Park. National Park Service Natural Resource Report NPS/SODN/

NRR—2011/437. National Park Service, Fort Collins, Colorado. 

Swann, D. E.; T. M. Mau-Crimmins and E. W. Stitt. 2005. In search of the 

Madrean Line: biogeography of herpetofauna of the Sky Islands. In Gottfried, G. J.; S. B. Gebow; L. G. Eskew; and C. B. Edminster, eds. Connecting mountain islands and desert seas: biodiversity and management 

of the Madrean Archipelago II. Proceedings RMRS-P-36. Fort Collins, 

Colorado: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station, 631 p. 

Swann D. E; and B. F. Powell. 2006. Mammal inventory of Saguaro National 

Park, Rincon Mountain District. Pp 69-97 in Powell, B. F.; W. L. Halvorson; 

and C. A. Schmidt, eds. Vascular Plant and Vertebrate Inventory of Saguaro 

National Park, Rincon Mountain District. Open-File Report 2006-1075, 

U. S. Geological Survey, Southwest Biological Science Center, Sonoran 

Desert Research Station, University of Arizona, Tucson, Arizona. 156 p.

Swann, D. E.; M. Bucci; A. J. Kuenzi; B. N. Alberti; and C. R. Schwalbe. 

2010. Challenges to natural resources monitoring in a small border park: 

terrestrial mammals at Coronado National Memorial, Cochise County, 

Arizona. Pp 225-239 in Halvorson, W. L.; C. R. Schwalbe; and C. R. van 

Riper, III, eds. Southwestern Desert Resources. University of Arizona 

Press, Tucson. 360 p.

Swarth, H. S. 1904. Birds of Huachuca Mountains. Pacific Coast Avifauna 

4:1-70. 

Tobler, M. W.; S. E. Carillo-Percastegui; R. L. Pitman; R. Mares; and G. Powell. 

2008. An evaluation of camera traps for inventorying large- and mediumsized terrestrial rainforest mammals. Animal Conservation 11:169-178. 

Turner, D. S.; S. Brandes; M. Fishbein; and P. W. Hirt. 1995. Preserve Design for maintaining biodiversity in the Sky Island region. Pp 524-530 

in: DeBano, L. F.; P. F. Ffolliott; A. Ortega-Rubio; G. J. Gottfried; R. H. 

Hamre; and C. B. Edminster (tech. cords.). Biodiversity and Management 

of the Madrean Archipelago: the sky islands of southwestern United States 

and northwestern Mexico. Gen. Tech. Report RM-GTR-264. Ft. Collins, 

Colorado: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Forest and Range Experiment Station. 

Van Devender, T. R. and A. L. Reina G. 2005. Floristic comparison of an 

Arizona Sky Island and the Sierra Madre Occidental in Eastern Sonora: 

the Huachuca Mountains and the Yecora area. Pp 154-157 in Gottfried, 

G. J.; S. B. Gebow; L. G. Eskew; and C. B. Edminster, eds. Connecting 

mountain islands and desert seas: biodiversity and management of the 

Madrean Archipelago II. Proceedings RMRS-P-36. Fort Collins, Colorado: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain Research 

Station, 631 p. 

Varas, C.; C. A. Lopez-Gonzalex; P. R. Krausman; and J. R. Morgart. 2010. 

Conservation genetics of black bears in the Sky Islands of Arizona and 

Northern Mexico. Pp. 275-282 in Halvorson, W. L.; C. R. Schwalbe; and 

C. R. van Riper, III, eds. Southwestern Desert Resources. University of 

Arizona Press, Tucson. 156 p.

Warshall, P. 1995. The Madrean Sky Island Archipelago: a planetary overview. 

Pp 6-18 in: De Bano, L. F.; P. F. Ffolliott; A. Ortega-Rubio; G. J. Gottfried; 

R. H. Hamre; and C. B. Edminster (tech. cords.). Biodiversity and Management of the Madrean Archipelago: the sky islands of southwestern United 

States and northwestern Mexico. Gen. Tech. Report RM-GTR-264. Ft. 

Collins, Colorado: U.S. Department of Agriculture, Forest Service, Rocky 

Mountatin Forest and Range Experiment Station. 

Wilson, D. E.; S. Ruff. 2000. The Smithsonian Book of North American 

Mammals. Smithsonian Books: Washington, DC. 750 p. 

Wilson, E. O.; F. M. Peter, eds. 1988. Biodiversity. National Academies Press, 

Washington, DC.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 277

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Habitat Type and Permanence Determine Local Aquatic 

Invertebrate Community Structure in the Madrean Sky 

Islands

Michael T. Bogan

Oregon State University, Corvallis, Oregon

Oscar Gutierrez-Ruacho and J. Andrés Alvarado-Castro,

CESUES, Hermosillo, Sonora, Mexico

David A. Lytle

Oregon State University, Corvallis, Oregon

Abstract—Aquatic environments in the Madrean Sky Islands (MSI) consist of a matrix of perennial and 

intermittent stream segments, seasonal ponds, and human-built cattle trough habitats that support a diverse suite of aquatic macroinvertebrates. Although environmental conditions and aquatic communities 

are generally distinct in lotic and lentic habitats, MSI streams are characterized by isolated perennial pools 

for much of the year, and thus seasonally occur as lentic environments. In this study, we compared habitat 

characteristics and Coleoptera and Hemiptera assemblages of stream pools with those of true lentic habitats (seasonal ponds and cattle troughs) across the MSI. We identified 150 species across the 38 sites, and 

despite superficial similarities in habitat characteristics, seasonal ponds and stream pools in the MSI support 

distinct aquatic insect communities. Stream-exclusive species included many long-lived species with poor 

dispersal abilities, while pond-exclusive species tended to have rapid development times and strong dispersal 

abilities. We suggest that, in addition to perennial streams, seasonal aquatic habitats should also be a focus 

of conservation planning in the MSI. 

Introduction

	 Abiotic factors canbehighlyinfluentialindetermininglocal aquatic

invertebrate community structure, including presence or absence of

flowing water (lotic vs. lentic) and hydroperiod (how long water lasts

in a given habitat). Lotic and lentic environments generally support

distinct aquatic insect communities (Huryn and others 2008). The

physical hydraulics of flow, oxygen availability, substrate, and temperature are among the primary physical drivers of this community

distinctness between lotic and lentic habitats (Vannote and Sweeney

1980; Resh and others 2008). Additionally, seasonal habitat drying

can act as a strong filter on aquatic insect communitiesin both streams

and ponds(Williams 2005).Seasonalstreams and ponds(habitatsthat

dry for extended periods oftime each year) often contain species with

life-history adaptations to drought (Wiggins and others 1980).

	 The Madrean Sky Island region (MSI) in the southwestern United

States and northwesternMexico contains a diversity oflotic and lentic

habitats, including streams with perennial and intermittent surface

water, seasonal ponds, and human-built cattle troughs. Perennial

streams are found in the higher elevations (1200-2200 m) of most

MSI mountain ranges. While wet winters and monsoon rains can

result in high-flow periods where stream pools are scoured and connected to one another, many MSI streams have low to zero flow for

much of the year, and the remaining isolated perennial pool habitats

approximate lentic environments. Seasonal ponds form in adjacent

valley bottoms; these ponds fill after summer monsoon rains or

prolonged winter storms and usually contain water for fewer than 4

months. Cattle troughs also provide small lentic habitats, which often

contain water year-round. All three of these environments provide

lentic habitat, but whether or not they were utilized by the same taxa

was previously unknown.

	 In this study, we surveyed stream pool, seasonal pond, and cattle

trough aquatic insect assemblages across the MSI. We focused our

analyses on beetles (Coleoptera) and true bugs (Hemiptera) because

species-level identification is possible and they comprise over 80% of

aquatic insect individuals in MSI stream pools during the dominant

low-flow period (Bogan and Lytle 2007). Our objectives were to

document the distribution of these taxa across the MSI and to examine assemblage differences among stream pool, seasonal pond, and

cattle trough habitats. While MSI stream pools approximate lentic

environments for much of the year, we hypothesized that stream

pools, seasonal ponds, and cattle troughs would contain distinct

beetle and true bug assemblages due to environmental differences

(e.g. hydroperiod, water temperature) among these habitat types.
278 USDA Forest Service Proceedings RMRS-P-67. 2013

Bogan Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure . . .

Methods

Habitat Types

	 Perennial stream pools occur as small, fragmented habitats within

MSI ranges(generally between 1200 and 2200 m), and contain water

year-round.Seasonalponds(occurringbelow1200m)thatwesurveyed

in 2004 were dry in April and June, wetted in August, and dry again

in November. Thus, these ponds contained water for no more than 4

consecutive months in 2004. Cattle troughs were found in the lower

reaches of canyons(usually below 1500m) and in valleys below 1200

m. These were small (1 to10 m2) concrete and metal tanks that were

fed by spring or well water. Cattle troughs generally contain water

year-round, but can dry occasionally if the pumps, hoses, or pipes

that feed them break. The history of individual troughs (e.g. when

they were constructed) was not available for this study. None of the

streams, seasonal ponds, or cattle troughs surveyed contained exotic

vertebrate predators (e.g. large-mouthed bass, bullfrogs) when they

were surveyed.

Collection Methods

	 Aquatic beetle and true bug assemblages were sampled from pools

in 25 streams, from 7 seasonal ponds, and from 5 cattle troughs

between May and July 2004. Streams were the focus of a separate

study (Bogan 2005), which is why more streams were surveyed than

ponds ortroughs.Streampoolswere sampled by vigorously sweeping

a D-net (1 mm mesh) above all pool substrata and on the surface of

the water for 10 s m-2 of pool area, an effort that detected approximately 95% of beetle and true bug taxa in a given pool (see Bogan

and Lytle 2007, 2011). Ponds, and occasionally troughs, were too

large to effectively sample the entire habitat as was done for stream

pools, so we focused our efforts on near-shore habitats.At each pond

and trough sampling site, a 1-mm mesh D-net was used to vigorously

sweep substrate and vegetation from the shore to 2 m offshore along

the entire habitat perimeter. All collected insects were placed in 95%

ethanol, andlateridentifiedatOregonStateUniversity.At eachsample

site, temperature, pH, and conductivity were recorded at a depth of

10 cm using hand-held meters. Since sampling did not always occur

at the same time of day, instantaneous water temperature values may

Table 1—Location of study sites in the Madrean Sky Islands region (SON = Sonora, Mexico; 

AZ = Arizona; NM = New Mexico).

Habitat type Site State Latittude Longitude

Stream pools Agua Caliente- Aconchi SON 29.846 110.2849

Agua Caliente- Cajon SON 31.2824 108.9963

Cañon Alacrán SON 28.01 111.2

Ash Creek AZ 32.50679 110.23639

Bear Canyon AZ 31.41056 110.2826

Cajon Bonito SON 31.27803 109.00212

Chulo Canyon AZ 31.48545 109.98251

Dixie Canyon AZ 31.49549 109.89718

East Turkey Creek AZ 31.91743 109.23194

Florida Canyon AZ 31.75657 110.84362

French Joe Canyon AZ 31.81758 110.40917

Gardner Canyon AZ 31.70217 110.80051

Leslie Creek AZ 31.58916 109.50729

Madera Canyon AZ 31.72416 110.88071

Sierra Mazatán SON 29.09106 110.22091

Miller Canyon AZ 31.40691 110.28742

Cañon Nacapule SON 28.00898 111.09712

Ramsey Canyon AZ 31.4442 110.3171

Rattlesnake Creek AZ 32.69534 110.34395

Rucker Canyon AZ 31.75669 109.36433

Simpson Spring AZ 31.78932 110.47029

Sycamore Creek AZ 31.41914 111.19361

Vallecitos- La Madera SON 29.92156 109.51361

West Stronghold Canyon AZ 31.93385 109.99599

West Turkey Creek AZ 31.86102 109.33618

Seasonal ponds Boulder Pond AZ 31.76126 110.4856

East Willow Pond AZ 31.81503 109.05965

Exit 292 Pond AZ 31.97767 110.50866

Halfmoon Pond AZ 31.91235 109.97793

Rock Pond AZ 31.94488 109.11757

Stuck Pond NM 31.86468 109.03671

The Thing Pond AZ 32.08254 110.05257

Cattle troughs Bear Spring Tank AZ 31.77523 110.4616

Cottonwood Tanks AZ 31.48941 109.07038

Guindani Tank AZ 31.84151 110.36936

Kartchner Tank AZ 31.8835 110.35767

 Rock Tank AZ 31.93941 109.11703
USDA Forest Service Proceedings RMRS-P-67. 2013 279

Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure . . . Bogan

not exhibitreliable differences betweensites.Toexplore diurnalwater

temperature fluctuations, we deployed HOBO remote temperature

probes (Onset Computer Corporation, Pocasset, MA) at 10 cm water

depth in two ponds (East Willow and “Stuck”; table 1) and pools in

two streams (one pool each in N. Fk. Cave Creek and French Joe

Canyon; table 1). HOBO probes in the streams were deployed from

May 2004 through June 2005, but the seasonal pond sites began to dry

soon after the probes were deployed, so data are only available from

29 July-2 August 2004. Finally, we visually estimated percentages

of substrate types (bedrock, cobble, gravel, sand, and fines) at each

site. All insect collections and visual estimations were made by the

same individual (MTB) to minimize data collector error.

Statistical Methods

	 Because sample sizes and variances were unequal, we used nonparametricMann-Whitneyteststoexaminedifferencesintemperature,

conductivity, and pH between stream and pond sites. All tests were

performedinSYSTATv.11(SystatSoftwareInc.,SanJose,California).

Aquatic insect assemblage variation among siteswas visualized using

non-metric multidimensional scaling (NMS) on Sorensen distance

matricesin PC-ORD v5.0 (MjM Software, Glenden Beach, Oregon).

Because pond aquatic invertebrate sampling methods were designed

to maximize species detection, rather than quantifying relative abundances, all assemblage matrices were presence-absence transformed

priortostatistical analyses.WeusedPearson’srcorrelationstoexamine

relationships between species presence, environmental factors, and

axis scores. To test distinctness of stream and pond assemblages, we

used Multi-Response Permutation Procedure (MRPP) tests.	

Results

	 Pond temperatures during sampling visits (mean ± 1 SD: 23.5 °C

± 5.6) were significantly higher than stream pool temperatures(mean

± 1 SD: 20.1 °C ± 3.1; U = 225.5, p = 0.03), but conductivity and

pH did not differ significantly between streams and ponds (p = 0.47

and 0.46, respectively). The HOBO temperature probes deployed

from 29 July - 2 August 2004 indicated that, in addition to overall

higher temperatures in ponds over those 5 days, diurnal temperature

fluctuation was much higher in ponds than in stream pools (fig. 1).

Mean pond temperature over those 5 days was 25.6 ± 5.1 °C, while

mean stream pool temperature was 18.3 ± 2.3 °C. Pond substrate

was predominately fine silt (mean ± 1SD: 80% ± 16), while stream

pool substrate was variable (means ± 1SD: 38% ± 26 bedrock, 20%

± 22 fine, 16% ± 15 gravel, 16% ± 16 cobble, 10% ± 13 sand).

	 One-hundred and fifty species of Coleoptera and Hemiptera were

identified from the 25 stream, 7 seasonal pond, and 5 cattle trough

sites. Sixty-five species(43% of allspecies) were exclusive to stream

pools,while15species(10%of allspecies)were exclusive toseasonal

ponds. Forty-six species(31% of allspecies) were habitat generalists,

encountered in both pond and stream habitats (see tables 2 and 3).

No species were unique to cattle toughs. The remaining 24 species

(16% of all species) were only encountered at one site so we could

not confidently evaluate their habitat affinity.

	 TheNMSordinationconvergedona stable,2-Dsolution(finalstress

= 18.7%, final instability = 0.003; p = 0.02; fig. 2). The ordination

explained 81% of the variance in the original species matrix (axis

1: r2 = 0.37; axis 2: r2 = 0.44). Axis 1 was influenced by Aquarius 

remigis (r = –0.55), Pelocoris (r = 0.50), and Hydrophilus insularis

(r = 0.79) among other species while axis 2 was influenced by Notonecta lobata (r = –0.75), Thermonectus marmoratus (r = –0.69),

Desmopachria portmanni (r = –0.65), Gyrinus plicifer (r = –0.64),

and Corisella edulis (r = 0.71), among others. Axis 2 values were

positively associated with fine substrates (r = 0.75) and negatively

associatedwithbedrocksubstrates(r=–0.53).Latitudewasnegatively

associated with axis 1 (r = –0.45) and temperature was positively associated with axis 2 values (r = 0.42); however, both of these factors

appear to be associated with assemblage gradients described in part

by both axes (i.e., the vectors are diagonal to axes 1 and 2: fig. 2). 

	 Moststreampoolsitesformed a tightly clustered group in the NMS

graph (fig. 2). Stream pools located on the extreme southwestern

margin of the MSI (Mazatán, Nacapule and Alacrán; table 1) had

higher axis 1 scoresthan otherstreams pools, but were within the range

of axis 2 scoresfor all otherstreams. Seasonal pond assemblages varied widely, while cattle trough assemblages showed less variability

and were closer in composition to stream pool assemblages. MRPP

analyses confirmed the distinctness of stream pool, seasonal pond,

and cattle trough assemblages (A = 0.29, p < 0.0001). 

Discussion

	 Despitesuperficialsimilaritiesinhabitatcharacteristics,streampools

and seasonal pondsin the MSIsupport distinct aquatic beetle and bug

assemblages. Of the 150 species identified in this study, 80 species

were exclusive to eitherstream pools orseasonal ponds. Sixty-five of

these species were only found in the perennialstream pools, while the

remaining 15 were exclusively found in the environmentally variable

seasonal ponds (tables 2 and 3). The relatively low number of pond

exclusive species may be due to harsh environmental conditions in

ponds, but could also be due to the low number of ponds sampled (n

= 7) when compared with streams (n = 25). Fairy shrimp and other

non-insect invertebrates are often specialistsin seasonal ponds(King

and others 1996), so a broader taxonomic consideration of our study

ponds and poolswould likely only increase the community distinction

among these three habitat types.

	 Temperature has been identified as a major factor in aquatic

insect species distributions in numerous studies (e.g. Hawkins and

Figure 1—Diurnal water temperature fluctuations recorded at 15-minute 

intervals in two seasonal ponds (black lines) and two stream pools (grey 

lines) from 00:00h 30 July 2004 to 21:00h 2 August 2005. Each crosshatch on the x-axis represents the beginning of that day (00:00h). The 

lower stream pool line is from North Fork Cave Creek and the upper is 

from French Joe Canyon. The lower seasonal pond line is from East Willow 

pond and the upper line is from “Stuck” pond (see table 1).
280 USDA Forest Service Proceedings RMRS-P-67. 2013

Bogan Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure . . .

Table 2— Coleoptera species identified from Madrean Sky Island region ponds and stream pools with their habitat affinities. Only 

common taxa (occurrences in >10% of sites) are listed in order of percentage of sites at which they were found (%). Generalist 

species were found in at least two of the three habitat types sampled.

Stream pool exclusive % Seasonal pond exclusive % Generalist species %

Thermonectus marmoratus 81 Berosus stylifer 57 Laccophilus fasciatus 80

Gyrinus plicifer 73 Eretes sticticus 29 Berosus salvini 66

Dineutus sublineatus 69 Berosus cf. infuscatus 29 Rhantus gutticollis gutticollis 61

Rhantus atricolor 62 Berosus cf. miles 29 Stictotarsus striatellus 51

Sanfilippodytes 62 Neoclypeodytes sp. 14 Tropisternus ellipticus 51

Stictotarsus corvinus 62 Berosus peregrinus 14 Stictotarsus aequinoctialis 49

Berosus rugulosus 58 Tropisternus lateralis 49

Rhantus gutticollis mexicanus 38 Desmopachria portmanni 46

Postelichus immsi 35 Thermonectus nigrofasciatus 44

Helichus triangularis 31 Peltodytes dispersus 41

Hydraena sp. 31 Berosus moerens 41

Hydrochus sp. 31 Hygrotus sp. 39

Stictotarsus roffi 27 Laccophilus horni 39

Helichus suturalis 23 Liodessus obscurellus 37

Neoclypeodytes fryii 23 Agabus sp. 34

Laccobius sp. 23 Desmopachria mexicana 34

Dytiscus habilis 19 Laccophilus maculosus 29

Anacaena limbata 19 Enochrus sp. 27

Laccophilus oscillator 15 Hydrophilus insularis 22

Elmidae sp. 15 Berosus punctatissimus 17

Liodessus sp. 12 Laccophilus mexicanus 15

 Neoclypeodytes cinctellus 15

 Helophorus sp. 15

 Laccophilus pictus 12

Table 3—Hemiptera species identified from Madrean Sky Island region ponds and stream pools with their habitat affinities. Only 

common taxa (occurrences in >10% of habitats) are listed in order of percentage of sites at which they were found (%).Generalist 

species were found in at least two of the three habitat types sampled.

Stream pool exclusive % Seasonal pond exclusive % Generalist species %

Abedus herberti 77 Corisella edulis 100 Microvelia sp. 76

Ambrysus woodburyi 35 Gerris sp. 43 Graptocorixa abdominalis 71

Trepobates becki 31 Notonecta unifasciata 43 Notonecta lobata 63

Pelocoris sp. 31 Belostoma flumineum 29 Ranatra quadridentata 59

Notonecta kirbyi 23 Corisella tarsalis 14 Aquarius remigis 54

Platyvelia beameri 23 Corisella sp. 14 Buenoa arizonis 44

Rhagovelia varipes 23 Ramphocorixa acuminata 14 Graptocorixa serrulata 39

Hydrometra sp. 19 Notonecta undulata 14 Lethocerus medius 34

Curicta pronotata 19 Salda sp. 14 Buenoa arida 32

Graptocorixa gerhardi 12 Buenoa scimitra 29

Neocorixa snowi 12 Morphocorixa lundbladi 20

Gelastocoris sp. 12 Notonecta hoffmani 17

others 1997). Indeed, we found that ponds had significantly higher

temperatures than stream pools and also that increased temperature

was associated with the shift from stream pool to seasonal pond

assemblage types. Additionally, diurnal variability in temperature

appears to be much higher in seasonal ponds (fig. 2), though we did

not measure this factor at all sites. Temperature variability may be

a more important factor than mean temperature when examining

environmental drivers of observed aquatic community differences

(Palmer and others 1997). High and variable temperatures can impact

aquatic insect respiration and survival of early instars by changing

oxygen levels and water density (Williams 2005). As seasonal ponds

only occurred below 1200 m and stream pools only occurred above

1200 m, elevation is clearly a confounding factor in this study. Temperature and dissolved oxygen often co-vary with elevation, and we

are unable to disentangle the effects of water quality and elevation

in this study. We also did not measure nutrient levels, which may be

higher in seasonal ponds visited by livestock, and recommend that

such measurements be included in future studies. 

	 Local physical habitat characteristics (e.g. substrate, channel

morphology) can also strongly influence local aquatic invertebrate

assemblageswithin a given biogeographic region.Sanderson and others(2005) determined that a gradient from silt- to boulder-dominated

substrates wasthe most important physicalfactor determining stream

invertebrate abundancesinanorthernEnglandriversystem.Similarly,

we identified a gradient in local beetle and true bug assemblages

that was strongly associated with a shift from bedrock-dominated to
USDA Forest Service Proceedings RMRS-P-67. 2013 281

Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure . . . Bogan

fines-dominated substrate across all our study sites and habitat types

(NMS axis 2; fig. 2).

	 Williams (2005) noted that a short hydroperiod may be the most

constraining environmental factor for determining the distribution of

aquatic organisms. Although we did not measure exact hydroperiod

at our study sites, we know that ponds held water for less than 4

months during 2004. The habitat affinities of several species in our

study, however, may provide insight into how short hydroperiods

(as found in seasonal ponds) could act to constrain MSI beetle and

true bug assemblages. Several of the exclusively stream pool species

are either flightless (e.g. Abedus herberti; Lytle 2000) or have weak

dispersal abilities(e.g. Rhantus atricolor; R.L. Smith, personal commumication), so it is intuitive that these species would be excluded

fromtemporaryhabitats.Indeed,ZimmermanandSmith(1975)noted

that R. atricolor collections are almost exclusively from perennial

mountain streams. In contrast, many pond exclusive species are capable of far and frequent dispersal (e.g. Corisella edulis: Hungerford

1948) and may benefit from reduced competition and predation in

harsh seasonal environments (Wellborn and others 1996). The pond

exclusive dytiscid beetle Eretes sticticus appears to be especially

well adapted to seasonal habitats. It has an extremely rapid larval

development time (15 days: Kingsley 1985) and has been collected

almost exclusively from temporary desert ponds (Larson and others

2000).

	 Beetle andtruebugassemblagesincattle troughswere intermediate

between those from stream pools and seasonal ponds and were more

similar to one another than seasonal pond assemblages(fig. 2). Aquatic

insects must colonize these habitats after the troughs are constructed,

creating a strong dispersal filter that could lead to more self-similar

local assemblages. Since cattle troughs are filled by spring or well

water, they generally containwater year-round.Thislonger hydroperiod

may favor a distinct group of species with longer development times

buttolerant of occasional dry periods(seeWilliams 2005)when a pipe

breaks or the trough’s source fails. Despite the apparent preference

for human-constructed habitats by a number of dytiscid beetle species (Larson and others 2000) we did not find any species exclusive

to cattle troughs. 

	 Though widely distributed across a large geographic area (~30,000

km2),moststreampool assemblagesweremore similarto one another

thanwereseasonalpondassemblages(fig.2).Importantenvironmental

factors, such as relatively stable temperatures (fig. 1) and reliable

perennialwater,mayallowlonger-livedpredatorswithweakdispersal

abilities (e.g. top predator Abedus herberti: flightless, lifespan up to

3 years; Lytle 2000; Lytle and Bogan unpublished data) to persist in

perennial stream pools but not in seasonal ponds. These predators,

among otherspeciesill-suited to seasonal pond conditions but highly

competitive in perennial habitats, may contribute to the uniformity

of stream pool assemblages by exerting a strong biotic influence on

local communities (Wellborn and others 1996). 

	 Much ofthe variability among streampoolsiteswasrepresented by

axis 1 in the ordination, which was associated with latitude. Indeed,

thereappearstobealatitudinal assemblage shift,withsomewidespread

Neartic species(e.g. Aquarius remigis and Rhantus gutticollis) being

characteristic of northern sites while taxa with Neotropical centers

of distribution were characteristic of southern sites (e.g. Pelocoris, 

Hydrophilus insularis). The MSI are often described as a meeting

place for Nearctic and Neotropical species, so it is not surprising

that we detected a north-south gradient in assemblages while looking

over a large geographic range (28°-32° latitude). We did not sample

enough seasonal ponds in the southern part of the MSI (i.e. south of

31° latitude) to assess latitudinal gradients in pond assemblages.

	 While the results from this current study suggest that hydroperiod

is an influential factor in determining lentic beetle and true bug assemblages acrossthe MSI, hydroperiod may be even more important

in lotic systems. In a recentstudy, we compared lotic aquatic invertebrate community structure in a network of perennial and intermittent

stream reaches in the Huachuca Mountains (Bogan and others, in

review). We found that intermittent reaches were species-poor when

comparedtoperennialreaches,butthatthesespecies-poorcommunities

were composed mostly of unique species with adaptations to life in

temporary water and were not found in perennial reaches. The short

flow duration of intermittent reaches (<4 months, often dry for 1 or

more years at a time)strongly favorsspecies with rapid development

times and dormant egg or larval stages, such as winter stoneflies,

blackflies, and non-biting midges. Thus, despite being species-poor

relative to neighboring perennialreaches, intermittentstream reaches

are important contributors to regional species richness.

	 We hope that the results of these studies will help inform aquatic

conservation planning in theMSIregion.Temporarywaters and small

perennialstreams are often overlooked during conservation planning,

while focus is placed on larger streams or perennial lakes. We identified a number of aquatic species in the MSI, however, that appear

to prefer or be restricted to seasonal habitats. Many geographically

restricted, endemic invertebrate species are found in seasonal ponds

in other parts of the southwestern United States(e.g. King and others

1996). With sufficient taxonomic studies, new invertebrate species

endemic to MSIintermittentstreams and seasonal ponds would likely

be described. Additionally, numerous other species seem to require

Figure 2—NMS ordination of the beetle and true bug assemblages from 

26 stream pool sites (black circles), 7 seasonal valley pond sites (open 

triangle) and 5 constructed cattle tank sites (grey triangles). Environmental 

factors with Pearson’s correlations greater than 0.4 with either axis are 

illustrated as vectors. The length of each factor’s vector is proportional 

to that factor’s correlation values with the two axes.
282 USDA Forest Service Proceedings RMRS-P-67. 2013

Bogan Habitat Type and Permanence Determine Local Aquatic Invertebrate Community Structure . . .

the stable perennial conditions associated with some streams. These

species are increasingly threatened by drying climate and contracting

or vanishing perennial stream habitat across the MSI. We are already

witnessing stream-scale extirpations of important taxa (e.g. the top

predator Abedus herberti) due to unprecedented drying of perennial

habitats and increasing isolation of remaining perennial habitats

(Bogan and Lytle 2011). In order to preserve local endemic species

and regional aquatic diversity, aquatic conservation planning in the

MSI should include a wide range of habitat types, from perennial to

ephemeral and from lotic to lentic.

Acknowledgments

	 This work was conducted under USDA Forest Service Special

Uses Permit #SUP0092 (U.S.A.) and SEMERNAT Permiso #SGPA/

DGVS/04147 (Mexico). Scientific collection permits for Garden

Canyon (U.S. Army Fort Huachuca; Permit #ATZS-ISB 200-3) and

Leslie Canyon (U.S. Fish & Wildlife Service; Permit #22523-2004-

86) were made possible by S. Stone and B. Radke, respectively.

Collections from Kartchner Caverns State Park were made possible

by R. Toomey. Thanks to D. Gori and D. Turner of the Tucson office

of the Nature Conservancy for providing GIS and habitat data and

access to Ramsey Canyon. Special thanks to private landowners J.

and V. Austin for access to the Cajon Bonito, C. Wertz for access to

Chulo Canyon, and R. Monroy Leon and vaquero Juan for access to

and assistance at Vallecitos. Partial funding came from Oregon State

University and National Science Foundation grant DEB-0445366 to

D. A. Lytle. C. Olson allowed the use of lab space and access to the

aquatic insectreference collectionattheUniversityofArizona. Thanks

to F. Drake and J. Miles for assistance with preliminary sorting and

identification of samples at Oregon State University. C. Zugmeyer

provided essential logistical support during field work. 

References

Bilton, D.T., J.R. Freeland, and B. Okamura. 2001. Dispersal in freshwater

invertebrates. Annual Review of Ecological Systems 32:159-181.

Bogan, M.T. 2005. Diversity and community structure of aquatic insects in

isolated montane desert streams. M.S. thesis, Oregon State University,

Corvallis.

Bogan, M.T. and D.A. Lytle. 2007. Seasonal flow variation allows ‘timesharing’by disparate aquatic insect communitiesinmontane desertstreams.

Freshwater Biology 52:290-304.

Bogan, M.T. and D.A. Lytle. 2011. Severe drought drives novel community

trajectories in desert stream pools. Freshwater Biology 56: 2070-2081.

Bogan, M.T., K.S. Boersma, and D.A. Lytle. [In review]. Unique intermittent

streaminvertebrate communities alterlongitudinalgradientsofdiversityand

community structure in an aridland stream network. Freshwater Biology.

Hawkins, C.P., J.N. Hogue, L.M. Decker, and J.W. Feminella. 1997. Channel morphology, water temperature, and assemblage structure of stream

insects. Journal of the NorthAmerican Benthological Society 16:728-749.

Hungerford,H.B. 1948.TheCorixidae oftheWesternHemisphere.University

of Kansas Science Bulletin 32:1-827.

Huryn, A.D., J.B. Wallace, and N.H. Anderson. 2008. Habitat, life history,

secondary production, and behavioral adaptations of aquatic insects. Pages

55-103 In: An introduction to the aquatic insects of North America 4th

edition. (R.W. Merritt, K.W. Cummins, and M.B. Berg, editors). Kendell/

Hunt, Dubuque, Iowa.

King,J.L.,M.A.Simovich, andR.C.Brusca.1996.Speciesrichness, endemism

and ecology of crustacean assemblagesin northernCalifornia vernal pools.

Hydrobiologia 328: 85-116.

Kingsley, K.J. 1985. Eretes sticticus (L.) (Coleoptera: Dytiscidae): Life history observations and an account of a remarkable event of synchronous

emigration from a temporary desert pond. Coleopterists Bulletin 39:7-10.

Larson, D.J., Y. Alarie, and R.E. Roughgarden. 2000. Predaceous diving

beetles (Coleoptera: Dytiscidae) of the Nearctic Region, with an emphasis

on the fauna of Canada and Alaska. National Research Council of Canada

Press, Ottawa, Ontario. 

Lytle, D.A. 2000. Biotic and abiotic effects of flash flooding in a montane

desert stream. Archiv fuer Hydrobiologie 150:85-100.

McCune B. and J.B. Grace. 2002. Analysis of ecological communities. MjM

Software Design, Glenden Beach, OR, U.S.A.

Mielke P.W.,Jr., and K.J. Berry. 2001. Permutation methods:Adistance function approach. Springer, Berlin.

Palmer, M.A., C.C. Hakenkamp, and K. Nelson-Baker. 1997. Ecological

heterogeneity in streams: Why variance matters. Journal of the North

American Benthological Society 16:189-202.

Resh,V.H., D.B.Buchwalter, G.A.Lamberti, andC.H.Eriksen. 2008.Aquatic

insect respiration. Pages 39-54 In:An introduction to the aquatic insects of

North America. Fourth edition. (R.W. Merritt, K.W. Cummins, and M.B.

Berg, editors). Kendell/Hunt, Dubuque, IA.

Sanderson, R.A., M.D. Eyre, and S.P. Rushton. 2005. The influence ofstream

invertebrate compositionatneighboringsitesonlocal assemblage composition. Freshwater Biology 50: 221-231.

Vannote, R L. and B.W. Sweeney. 1980. Geographic analysis of thermal

equilibria: A conceptual model for evaluating the effect of natural and

modified thermal regimes on aquatic insect communities. American

Naturalist 115:667-695.

Wellborn, G.A., D.K. Skelly, and E.E. Warner. 1996. Mechanisms creating

community structure across a freshwater habitat gradient. Annual Review

of Ecological Systems 27:337-363.

Wiggins,G.B.,R.J.Mackay, andI.M.Smith.1980.Evolutionaryandecological

strategies of animals in annual temporary pools. Archiv für Hydrobiologie

Supplement 58:97-206.

Williams, D. D. 2005. The ecology of temporary waters. Oxford University

Press, New York.

Zimmerman, J. R. and R.L. Smith. 1975. The genus Rhantus (Coleoptera:

Dytiscidae) in North America. Part I. General account of the species.

Transactions of the American Entomological Society 101:33.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 283

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Gila Topminnow Interactions With Western Mosquitofish: 

An Update

Douglas K. Duncan

Arizona Ecological Services Office, U.S. Fish and Wildlife Service, Tucson, Arizona

Abstract—Western mosquitofish Gambusia affinis has major impacts on Gila topminnow Poeciliopsis occidentalis. Thirteen years have passed since information on negative impacts has been published. I will list 

and discuss additional examples where mosquitofish have impacted topminnow. In addition, it appears 

that long-term drought has a synergistic and negative effect on this relationship. Since the last publications 

detailing the loss of Gila topminnow populations to mosquitofish, two natural topminnow populations have 

been lost in the Santa Cruz River basin: Redrock Canyon and Sharp Spring. Drought and climate change 

likely exacerbated the impacts of mosquitofish on these two populations, speeding the decline and disappearance of topminnow from both. Dynamic conservation actions are required to conserve and recover 

the Gila topminnow.

Introduction

	 It haslong been known and thoroughly documented that Gambusia 

affinis western mosquitofish (mosquitofish) has major deleterious

effects on individual Poeciliopsis occidentalis Gila topminnow (topminnow) and their populations.It has been 13 yearssince information

on these negative impacts has been updated. In this paper I will list

and discuss additional examples where mosquitofish have impacted

topminnow. I will also summarize the apparent mechanisms of how

mosquitofish extirpate topminnow, and what the future oftopminnow

conservation might require.

	 The last peer-reviewed paper on topminnow-mosquitofish interaction was by W. L. Minckley (1999), and the last publication of any

sort discussing this was by Voeltz and Bettaso (2003). Further information can be found in Minckley and others (1977, 1991) and Meffe

and others (1983. These publications and others (Meffe and others

1982; Miller 1961) have made it abundantly clear that mosquitofish

negativelyimpacttopminnow, anddocumentedthe likelymechanisms

responsible (Meffe 1984, 1985; Schoenherr 1974). However, a brief

review of the mechanisms by which mosquitofish impact multiple

species, and topminnow populations in particular, are appropriate.

	 Mosquitofish was first collected in Arizona in the Phoenix area

in 1926 (Miller and Lowe 1964), in habitats that also held topminnow and other native fishes. Subsequently, mosquitofish came to be

distributed, most likely through purposeful release, not all legally,

throughout Arizona (Meffe 1985; Minckley 1973). They have also

beenreleasedworldwide(CourtenayandMeffe1989;ISSG2010), and

have been called the most introduced species in the world (Welcome

1988).

	 In addition to impacts on Gila topminnow, mosquitofish have

impacted other native fish in the western United States (Deacon and

others 1964; Meffe 1985; Whitmore 1997) and the rest of the world

(Arthington and Lloyd 1989; Glover 1989; Milton and Arthington

1983;UnmackandBrumley1991).Mosquitofishhavealsobeenshown

or suspected to prey on tadpoles (Goodsell and Kats 1999; Morgan

and Buttemer 1997; Webb and Joss 1997), newts (Gamradt and Kats

1996), frogs (Rosen and others 1995), and aquatic insects (Carchini

and others 2003; Englund 1999). Several studies have demonstrated

that mosquitofish (Crandall and Bowser 1982; Dykova and others

1994; Lom and Dykova 1995; Moravec 1998) and related species

(Perlmutter and Potter 1987) carry parasites and disease that can be

transferred into aquatic systems where they are released. Lastly, once

introduced, mosquitofish are also known to cause cascading changes

in the function of aquatic systems(Bence 1988; Hoy and others 1972;

Hurlbert and others 1972; McDowall 1990; Ortega-Mayagoita and

others 2002). Mosquitofish have been widely distributed by state

vector control agencies to control mosquitoes.

	 Thepredominantmechanismsofmosquitofisheffectontopminnow

are aggression and predation (Meffe and others 1983; Minckley and

Deacon 1968; Schoenherr 1977). Mosquitofish often replace topminnow in a matter of months (Meffe and others 1983; Schoenherr

1974), but with exceptions where the two species coexisted for years

(Minckley 1999). Possible means of coexistence have been shown to

be the species’differential response to flooding (Galat and Robertson

1988; Meffe 1984), or theorized greater habitat size and complexity

(Meffe1985).Anotherpotentialmechanismforprolongedcoexistence

is that mosquitofish do not invade springheads with extreme water

quality (Hubbs 1995), but are occupied by congeners (Hubbs 1957,

1971).Topminnowistolerantofextremespringheadconditions(Marsh

and Minckley 1990; Minckley 1973). The sites where mosquitofish

have extirpated topminnow in a few months to a few years have all

been small and simple habitats, generally reestablishment sites; in

natural habitats replacement takes longer.

	 Minckley (1999) also mentioned an additional alternative based

on work by Hubbs (1991, 1992, 1996) showing that some western

mosquitofish do not cannibalize their young. Minckley theorized that

certain stocks of mosquitofish could be less predatory, and that could
284 USDA Forest Service Proceedings RMRS-P-67. 2013

Duncan	 Gila Topminnow Interactions With Western Mosquitofish: An Update

be an explanation for coexistence of topminnow and mosquitofish.

Minckley (1999) also offered an alternative hypothesis involving

groundwater inflow. Groundwater inflow tends to be warmer than

surrounding waters, and may provide thermal refuge during periods

ofprolongedfreezing temperatures.However,topminnows havebeen

observed swimming below surface ice. Most perennial waters that

harbor or have harbored topminnow also are groundwater dependent.

However, I believe reliable water is the reason topminnows persist

at those sites, and not the thermal refuge provided there.

Results/Update

	 There are several additional topminnow sites with mosquitofish

where topminnow inArizona have been lostsince Minckley’sreview

(1999). Table 1 shows these sites, as well as updates the status of the

natural populations. The draft revised Gila topminnow recovery plan

(Weedman 1999:58) defines a natural population as “a population

which existed prior to fish transplants by humans, which existstoday

in its historic location free of knownmixing with other populations by

humans (Simons 1987).” While only a few sites have been recently

lost to mosquitofish, any loss of populations of an endangered species

is disconcerting, especially natural populations, which are crucial for

species survival (Weedman 1999).

	 There have been two natural populations lost since 1999. Sharp

Spring, a tributary headwater spring to the upper Santa Cruz River,

was where Meffe (1984) conducted field studies demonstrating that

Gila topminnow withstood floods better than mosquitofish. The last

year topminnow were found in Sharp Spring, in the San Rafael Valley, was 2001 (Voeltz and Bettaso 2003). Since 2001, no topminnow

have been found despite one to two surveys annually there, while

mosquitofish are usually numerous. Mosquitofish were first recorded

here in the late 1970s. Sharp Spring contained a large and vigorous

population of topminnow until mosquitofish accessed the system.

Since 1995, I have observed the continual decline in the amount of

aquatic habitat at Sharp Spring.

	 Redrock Canyon isthe other natural topminnow population lost to

mosquitofish since 1999. Redrock Canyon issignificant for Gila topminnowconservationfortworeasons:itisoneofonlytwopopulations

onpublic land(U.S.ForestService)(Rinne and others1980;Stefferud

and Stefferud 1995), and it was the one remaining site that may have

mimicked the historical expansion and contraction within basins and

subbasins (Minckley 1999). A single topminnow was captured here

in 2008 (Stefferud and Stefferud 2008). Mosquitofish are still found

in a small drainage from Cott Tank to Redrock Canyon where the

last Gila topminnow was found; longfin dace Agosia chrysogaster

are uncommon in lower Redrock Canyon although were common

throughout the drainage before the 2000’s. Reviews of management,

condition, and fish surveys in the Redrock Canyon watershed can be

found in Stefferud and Stefferud (1995), U.S. Bureau of Reclamation

and U. S. Forest Service (2008), and U.S. Fish and Wildlife Service

(1999:262-276). Redrock Canyon likely had more surface water historically, but has been dryer over the last decade (USBR and USFS

2008). The flow or extent of water in Redrock has not been regularly

measured.

	 I believe that mosquitofish impacts on topminnow, in combination

with long-term drought, was a factor in the extirpation of topminnow

atRedrockCanyon and Sharp Spring.Long-termdrought and climate

change also appear to be a factor in the extirpation or reduction of

other Gila topminnow populations(Bodner and others 2007; Duncan

and Garfin 2006; Voeltz and Bettaso 2003).

	 Arizona State Parks collected water level data at Sharp Spring

from 2000 to 2006 demonstrating reduced water quantity over time

(unpublished).Thoughwaterlevelswerehighlyvariable (fig.1),there

was an overall downward trend in water levels(fig. 1: trend lines). Of

the 10 pools measured, 8 had downward trending water levels, one

was static, and one had a slight upward trend. Seven pools went dry

at some time, and all of those were also dry in the last two measurements (fig. 1:2005-2006). The flow or extent of water in Redrock

has not been regularly measured. The best discussion of water in

the Redrock system is the Bureau of Reclamations’ and U.S. Forest

Service’s Redrock Barrier environmental assessment (USBR and

USFS 2008).

	 In addition to many of the sitesthat were reestablished in the 1980s

thatwere losttodesiccation(Minckley1999;Simons1987;Voeltz and

Bettaso 2003), I believe Gila topminnow at Heron Spring, Johnson

Wash Spring, and Willow Spring were extirpated (table 2) (Robinson

2010; Voeltz and Bettaso 2003), and populations at Cienega Creek

reduced, due to drying caused or exacerbated by the current drought

(Bodner and others 2007)

	 Heron Spring was stocked in 1981 with topminnow from nearby

Sharp Spring (Voeltz and Bettaso 2003). Heron Spring is a small

plunge pool that intercepts the locally high water table, with cienega

conditions extending as outflow for about 100m.Topminnow, though

rare,weregenerallyfoundintheplungepool(Voeltz andBettaso2003)

until 2007 when we captured no topminnow and measured dissolved

oxygenat1ppm(Ehret2009).Gila topminnowcansometimestolerate

that little dissolved oxygen, but the oxygen may have been less than

that before we measured it. The staff gauges maintained by Arizona

State Parks there showed no decline in water levels (unpublished

data). Discussions with others lead us to believe reduced subflow

into the spring, floating vegetation covering the pond surface, and

aquatic vegetation reducing outflow, combined to reduce dissolved

oxygen until topminnow could no longer survive in Heron Spring.

Table 1—Status of all natural Gila topminnow populations in the 

United States and Mexico. Sites in bold CAPS are ones where 

topminnow no longer occur (from Minckley 1999, Voeltz and 

Bettaso 2003, and USFWS files).

Site Extant?a Mosquitofish?

Bylas Spring S1 YESb NOc

Cienega Creek YES NO

Coal Mine Spring YES NO

Cocio Wash NO 1982 DRY

Cottonwood Spring YES NOd

Fresno Canyon YES NOe

Middle Spring S2 YES NOb,c

Monkey Spring YES NO

REDROCK CANYON NO 2005 YESf

Salt Creek S3 YES NOb,c

San Pedro River NO 1976 YES

Santa Cruz River

 San Rafael NOg YES

Tumacacori YES 2003 YES

SHARP SPRING NO 2001 YES

Sheehy Spring NO 1987 YES

Sonoita Creek YES YES

a

 If no, last year recorded.

b

 Renovated.

c

 None recently, they have been recorded.

d

 Recorded in spring run, never in head spring.

e

 Recorded downstream, below barrier. Renovated to remove green sunfish in 2007 

and is free of nonnatives; Sonoita Creek has many nonnatives.

f

 Slated for renovation.

g

 In Mexico 2006, United States in 1993.
USDA Forest Service Proceedings RMRS-P-67. 2013 285

Gila Topminnow Interactions With Western Mosquitofish: An Update Duncan

	 Cienega Creek has the largest occupied topminnow habitat in the

United States (Weedman 1999), though numbers in the upper perennial reach of Cienega Creek declined from 1989 to 2005 (Bodner and

others 2007). The actual location of the first pool has moved downstream (Bodner and others 2007; personal observation) since about

2001, indicating reduced groundwaterlevels and subflow in that area.

This has likely led to reduced dissolved oxygen in combination with

more vegetation (trees shading the water surface, biological oxygen

demand from decaying vegetation).

	 It appears that reduced volume and complexity of topminnow

habitat, when combined with presence of mosquitofish, is more

problematic for the persistence of topminnow, as opposed to the

presence of mosquitofish alone. Therefore, since climatologists expect the southwestern United States to be warmer and drier (Seager

and 2007), the impact of mosquitofish on topminnow populations is

expected to be greater in the coming decades as topminnow habitat

quality and quantity continues to decline.

Future Management

	 Actionsto conserve andrecovertheGila topminnowinArizona and

NewMexicoareongoingandadditionalactionsareplanned.Manynew

sites have been stocked with topminnow in the last 5 years(NMDGF

and others 2010; Robinson 2010; USFWS and others 2007), and only

time will tell if they persist. The involved management agencies and

landowners will continue with stockings, under the Arizona Game

andFishDepartmentSafeHarborAgreement(AGFD2007) and other

mechanisms. They will also continue stream maintenance, habitat

restoration, augmentation stocking, population and habitat monitoring, and outreach and education. Only vigorous conservation efforts

can lead to the conservation and recovery of topminnow in the face

of the threats they face. We are also exploring the use of topminnow

in vector control programs in place of mosquitofish. Lastly, all attention must be given to reducing the use, populations, and movement

of mosquitofish.

Acknowledgments

	 I would like to acknowledge all those individuals whose conversationsregardingtopminnowconservation,andspecificallymosquitofish

issues, have assisted my understanding of the issue; AGFD: Rob

Bettaso, Rebecca Davidson, Ross Timmons, Jeremy Voeltz, Dave

Weedman,KirkYoung;USFWS:SallyStefferud;BLM:HeidiBlasius,

Jeff Simms; ASU: Paul Marsh, W. L. Minckley; BR: Rob Clarkson;

USFS: Jerry Stefferud. I would also like to thank my peer reviewers,

Ross Timmons and Jerry Stefferud.

References

Arizona Game and Fish Department. 2007. Safe Harbor Agreement for topminnows and pupfish in Arizona. Phoenix, AZ. 36 p.

Figure 1—Averaged staff gauge measurements and trend line at Sharp Spring, Arizona (Arizona State Parks, 

unpublished).

Table 2—Status of some reestablished Gila topminnow 

populations in the United States. Site in bold CAPS is where 

topminnow no longer occur (from Minckley 1999, Voeltz and 

Bettaso 2003, and USFWS files).

Site Extant?a Mosquitofish?

HERON SPRING NO 2006 NO

JOHNSON WASH SPRING NO 2001 NO

Kayler Spring YES 2004 YES

WATSON WASH NO 1998 DRYb

WILLOW SPRING NO 2009C DRY

a

 If no, last year recorded.

b

 Well capped – mosquitofish present at capping.

c

 Topminnow released in 2009.
286 USDA Forest Service Proceedings RMRS-P-67. 2013

Duncan	 Gila Topminnow Interactions With Western Mosquitofish: An Update

Arthington, A.H.; Lloyd, L.L. 1989. Introduced poeciliids in Australia and

New Zealand. In: Meffe, G.K.; Snelson, F.F., eds. Ecology and evolution

of livebearing fishes (Poeciliidae). New Jersey: Prentice Hall: 333-348.

Bence, J.R. 1988. Indirect effects and biological control of mosquitoes by

mosquitofish. The Journal of Applied Ecology. 25(2): 505-521.

Bodner, Gita; Simms, Jeff; Gori, Dave. 2007. The state of the Las Cienegas

National ConservationArea: Gila topminnow population status and trends

1989-2005. Tucson, AZ: The Nature Conservancy in Arizona; Bureau of

Land Management, Tucson Field Office. 50 p.

Carchini G.; Di Domenico, M.; Pacione, T.; Solimini,A.G.; Tanzilli, C. 2003.

Species distribution and habitat features in lentic Odonata. Italian Journal

Zoology 70: 39-46.

Courtenay, W.R., Jr.; Meffe, G.K. 1989. Small fishes in strange places: A

review of

introduced poeciliids. In: Meffe, G.K.; Snelson, F.F., eds. Ecology and evolution oflivebearingfishes(Poeciliidae).NewJersey:PrenticeHall. 319-331.

Crandall, T.A.; Bowser, P.R. 1982. A microsporidian infection in mosquitofish, Gambusia affinis, from Orange County, California. California Fish

and Game. 68: 59-61.

Deacon, James E.; Hubbs, Clark; Zahuranec, Bernard J. 1964. Some effects

of introduced fishes on the native fish fauna in southern Nevada. Copeia.

1964: 384-388.

Duncan,Doug;Garfin,Gregg.2006.Nativefishconservation andclimate variabilityinsoutheasternArizona.In:Halvorson,Bill, ed.Borders,boundaries,

and time scales: proceedings ofthe 6th conference on research and resource

management in the southwestern deserts; extended abstracts. Tucson, AZ:

U. S. Geological Survey, Southwest Biological Science Center, Sonoran

Desert Research Station: 41-43

Dykova, Iva; Lom, Jin; Overstreet, Robin M. 1994. Myxosporean parasites

of the genus Kudoa Meglitsch, 1947 from some Gulf of Mexico fishes:

Description of two new species and notes on their ultrastructure. European

Journal of Parasitology. 30: 316-323.

Ehret, Suzanne. 2009.Upper SantaCruz trip report,June 22-23, 2009.Tucson,

AZ: Arizona Game and Fish Department, Region V Office. 6 p.

Englund, Ronald A. 1999. The impacts of introduced poeciliid fish and Odonata on the endemic Megalagrion (Odonata) damselflies of Oahu Island,

Hawaii. Journal of Insect Conservation. 3: 225-243.

Galat, David L.; Robertson, Ben. 1992. Response of endangered Poeciliopsis 

occidentalis sonoriensis in the Rio Yaqui drainage, Arizona, to introduced

Gambusia affinis. Environmental Biology of Fishes. 33: 249-264.

Gambradt, Seth C.; Kats, Lee B. 1996. Effect of introduced crayfish and mosquitofish on California newts. Consonservation Biolology. 10: 1155-1162

Glover,C.J.M.1989.Fishes.In:Zeidler,W.;Ponder,W.F.,eds.Naturalhistoryof

DalhousieSprings.Adelaide,Australia:SouthAustralianMuseum. 89-112.

Goodsell, Jeff A.; Kats, Lee B. 1999. Effect of introduced mosquitofish on

Pacific treefrogs and the role of alternate prey. Conservation Biology.

13(4): 921-924.

Hoy, J.B.; Kaufmann, E.E.; O’Berg, A.J. 1972. A large-scale field test of

Gambusia affinis and Chlorpyrifos for mosquito control. Mosquito News.

32(2): 163-171.

Hubbs, C. 1971. Competition and isolation mechanisms in the Gambusia 

affinis x G. heterochir hybrid swarm. Bulletin of the Texas Memorial

Museum. 8: 1-46.

Hubbs,C. 1996.Geographic variation ofGambusia life history traits.Proceedings of the Desert Fishes Council. 27: 1-21.

Hubbs, Clark. 1957. Gambusia heterochir, a new poeciliid fish from Texas,

with an account of its hybridization with G. affinis. Tulane Studies in

Zoology. 5: 3-16.

Hubbs, Clark. 1991. Intrageneric “cannibalism” in Gambusia. Southwestern

Naturalist. 36(2):153-157

Hubbs,Clark. 1992.Geographic variation in cannibalismof congeneric young

by Gambusia adults. Proceedings of the Desert Fishes Council. 22: 43-52.

Hubbs, Clark. 1995. Perspectives: springs and spring runs as unique aquatic

systems. Copeia. 1995: 989-991.

Hurlbert, Stuart H.; Zedler, Joy; Deborah Fairbanks. 1972. Ecosystem alteration by mosquitofish (Gambusia affinis) predation. Science. 175: 639-641.

Invasive Species Specialist Group. 2010. Gambusia affinis. International

Union for the Conservation of Nature, 2 p. Available: issg.org/database/

species/. [June 21, 2010].

Lom, J.; Dykova, I. 1995. Studies on protozoan parasites ofAustralian fishes.

Notes on coccidian parasites with description of three new species. Systematic Parasitology. 31: 147-156.

Marsh, Paul C.; Minckley, W.L. 1990. Management of endangered Sonoran

topminnow at Bylas Springs, Arizona. Description, critique and recommendations. Great Basin Naturalist. 50(3): 265-272.

McDowall, R. M. 1990. New Zealand freshwater fishes, a natural history and

guide. Heinemann Reed. 555 p.

Meffe, Gary K. 1983. Ecology ofspeciesreplacement in the Sonoran topminnow (Poeciliopsis occidentalis) and the mosquitofish (Gambusia affinis)in

Arizona. Tempe, AZ: Arizona State University. Dissertation.

Meffe,GaryK. 1984.Effectsof abioticdisturbanceoncoexistenceofpredatorprey fish species. Ecology. 65(5): 1525-1534.

Meffe, Gary K. 1985. Predation and species replacement in American southwestern fishes—a case-study. Southwestern Naturalist. 30(2): 173-187.

Meffe, Gary K.; Hendrickson, Dean A.; Minckley, W.L.; Rinne, John N.

1983. Factors resulting in decline of the endangered Sonoran topminnow

Poeciliopsis occidentalis (Atheriniformes:Poeciliidae)intheUnitedStates.

Biological Conservation. 25(1983): 135-159.

Meffe, Gary K.; Hendrickson, Dean A.; Rinne, John N. 1982. Description of

a new topminnow population inArizona, with observations on topminnow/

mosquitofish co-occurrence. Southwestern Naturalist. 27(2): 226-228.

Miller, Robert Rush. 1961. Man and the changing fish fauna of the American

Southwest. Papers of the Michigan Academy of Sciences, Arts and Letters. 46: 365-404.

Miller, Robert Rush; Lowe, Charles H. 1964. Fishes of Arizona. In: Lowe,

C. H., ed. The vertebrates of Arizona. Tucson, AZ: University of Arizona

Press: 133-151.

Milton, D.A.; Arthington, A.H. 1983. Reproductive biology of Gambusia 

affinis holbrooki Baird and Girard, Xiphorous helleri (Günther) and X. 

maculates(Heckel) (Pisces; Poeciliidae) in Queensland,Australia.Journal

of Fish Biology. 23: 23-41.

Minckley, W.L. 1973. Fishes of Arizona. Phoenix, AZ: Arizona Game and

Fish Department. 293 p.

Minckley, W.L. 1999. Ecological review and management recommendations

for recovery of the endangered Gila topminnow. Great Basin Naturalist.

59(3): 230-244.

Minckley, W.L.; Meffe, Gary K.; Soltz, David L. 1991. Conservation and

managementofshort-livedfishes:The cyprinodontoids.In:Minckley,W.L.;

Deacon, James E., eds. Battle against extinction—native fish management

in the American West. Tucson, AZ: University of Arizona Press: 247-282.

Minckley, W.L.; Rinne, John N.; Johnson, James E. 1977. Status of the Gila

topminnow and its co-occurrence with mosquitofish. Res. Pap. RM-198.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service. Rocky

Mountain Forest and Range Experiment Station. 8 p.

Minckley, W.L.; Deacon,James E. 1968. Southwestern fishes and the enigma

of “endangered species.”” Science. 159: 1424-1432.

Moravec, F. 1998. Nematodes of freshwater fishes of the Neotropical region.

Academia. 464 p.

Morgan, L.A.; Buttemer, W.W. 1997. Predation by the non-native fish Gambusia holbrooki on small Litoria aurea and L. dentata tadpoles. Australian

Zoology. 30: 143-149.

New Mexico Department of Game and Fish, Gila National Forest, and New

Mexico Fish and Wildlife Conservation Office. 2010. Gila River Basin

NativeFishesConservationProgram:NewMexicoactivities.Albuquerque,

NM: New Mexico Department of Game and Fish, U.S. Forest Service, U.S.

Fish and Wildlife Service. 7 p.

Ortega-Mayagoita,Elizabeth;Rojo,Carmen;Rodrigo,MariaA. 2002.Factors

masking the trophic cascade in shallow eutrophic wetlands: Evidence from

a microcosm study. Archiv Fur Hydrobiologie. 155(1): 43-63.

Perlmutter,Alfred; Potter, Hugh. 1987.Retrovirus-like particlesin embryonic

kidney tissue of the platyfish, Xiphophorus maculatus. Journal of Experimental Zoology. 243: 125-135.

Rinne,John N.; Rickel, Bryce; Hendrickson, Dean. 1980.Anew Gila topminnow locality in southern Arizona. Res. Note RM-382. Fort Collins, CO:

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest

and Range Experiment Station. 3 p.
USDA Forest Service Proceedings RMRS-P-67. 2013 287

Gila Topminnow Interactions With Western Mosquitofish: An Update Duncan

Robinson,Anthony. 2010.Arizona nativefishrecoveryand nonnativefishcontrol.Phoenix,AZ:ReporttoU.S.FishandWildlifeService,FWSAgreement

No. 201816J808: Final Report, Arizona Game and Fish Department. 34 p.

Rosen, Philip C.; Schwalbe, Cecil R.; Parizek, David A. Jr.; Holm, Peter A.;

Lowe, Charles H. 1995. Introduced aquatic vertebrates in the Chiricahua

region: Effects on declining native ranid frogs. In: DeBano, Leonard F.;

Ffolliott,PeterF.;Ortega-Rubio,Alfredo;[andothers], eds.Biodiversityand

management of the Madrean archipelago: the sky islands of southwestern

United States and northwestern Mexico, 1994 September 19-23; Tucson,

AZ. Gen. Tech. Rep. RM-GTR-264. Fort Collins, CO; U.S. Department

of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 251-261.

Schoenherr, Alan A. 1974. Life history of the topminnow, Poeciliopsis occidentalis (Baird and Girard) in Arizona, and an analysis of its interaction

with the mosquitofish Gambusia affinis (Baird and Girard). Tempe, AZ:

Arizona State University. Dissertation.

Schoenherr,AlanA. 1977.Density dependent and density independentregulationofreproductionintheGila topminnow,Poeciliopsis occidentalis(Baird

and Girard). Ecology. 58(2): 438-444.

Seager, Richard; Ting, Minfang; Held, Isaac; Kushnir, Yachanan; Lu, Jian;

Vecchi, Gabriel; Huang, Huei-Ping; Harnik, Nili; Leetmaa, Ants; Lau,

Ngar-Cheung; Li, Cuihua; Velez, Jennifer; Naik, Naomi. 2007. Model

projections of an imminent transition to amore arid climate in southwestern

North America. Science. 316: 1181–1184.

Simons, Lee H. 1987. Status ofthe Gila topminnow (Poeciliopsis occidentalis 

occidentalis) in the United States. Phoenix, AZ: Arizona Game and Fish

Department. 36 p.

Stefferud, Jerome A.; Stefferud, Sally E. 1995. Status of Gila topminnow and

results of monitoring the fish community in Redrock Canyon, Coronado

National Forest, 1979-1993. In: DeBano, Leonard F.; Ffolliott, Peter F.;

Ortega-Rubio,Alfredo;[and others], eds.,Biodiversity andmanagement of

theMadreanarchipelago:The skyislandsofsouthwesternUnitedStates and

northwesternMexico, 1994September 19-23;Tucson,AZ.Gen.Tech.Rep.

RM-GTR-264. Fort Collins, CO; U.S. Department of Agriculture, Forest

Service, Rocky Mountain Forest and Range Experiment Station: 361-369.

Stefferud, Sally E.; Stefferud,JeromeA. 2008. Report of activitiesin calendar

year 2008 under Permit number TE053109-0. Phoenix, AZ. 4 p.

U.S. Fish and Wildlife Service. 1999. Biological opinion: proposed on-going

and long-term grazing on the Coronado National Forest. 2-21-98-F-399,

Tucson, AZ: Arizona Ecological Services Field Office. 370 p.

U.S.FishandWildlifeService,U.S.BureauofReclamation,NewMexicoGame

and Fish Department, and Arizona Game and Fish Department. 2007Gila

River Basin Native Fishes Conservation Program: revised strategic plan

2008-2012. Phoenix, AZ. 16 p.

U.S. Bureau of Reclamation and U. S. Forest Service. 2008. Final EnvironmentalAssessment: native fish restoration in Redrock Canyon, Santa Cruz

County, Arizona. Phoenix, AZ: Phoenix Area Office, U.S. Department of

the Interior;Tucson,AZ: Coronado National Forest, Southwestern Region,

U.S. Department of Agriculture. 171 p.

Unmack, P.; Brumley, C. 1991. Initial observations on the spawning and

conservation status of the redfinned blue-eye, (Scaturiginichthys vermeilipinnis). Fishes of Sahul. 6(4): 282-284. (Journal of the Australian New

Guinea Fishes Association, Australia).

Voeltz, Jeremy B.; Bettaso, Robert H. 2003. 2003 status of the Gila topminnow and desert pupfish in Arizona. Phoenix, AZ: Arizona Game and Fish

Department. 124 p.

Webb, C.; Joss, J. 1997. Does predation by the fish Gambusia holbrooki

(Atheriniformes: Poeciliidae) contribute to declining frog populations?

Australian Zoologist. 30: 316-324.

Weedman,DavidA.1999.Gilatopminnow,Poeciliopsis occidentalis occidentalis,revisedrecoveryplan.Albuquerque,NM:U.S.FishandWildlifeService.

Welcome, R.L., comp. 1988. International introductions of inland aquatic

species. FAO Fish. Tech. Pap. 294. 318 p.

Whitmore,S.1997.AquaticnuisancespeciesinRegion6oftheFishandWildlife

Service, Pierre, SD: U.S. Fish and Wildlife Service, Great Plains Fish and

Wildlife ManagementAssistance Office, Fish andWildlife Service Report.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
288 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Native Aquatic Vertebrates: Conservation and 

Management in the Río Sonoyta Basin, Sonora, Mexico

C. Minckley

Cuenca los Ojos, Flagstaff, Arizona

Izar Izaguirre Pompa

Reserva de la Biosfera del Pinacate y Gran Desierto de Altar, Puerto Peñasco, Sonora, Mexico

Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Ross Timmons

Arizona Game and Fish Department, Phoenix, Arizona

Dennis Caldwell

Caldwell Design, Tucson, Arizona

Jaime López Méndez

Colegio de Bachilleres del Estado de Sonora, Sonoyta, Sonora, Mexico

Phil Rosen

University of Arizona, Tucson, Arizona

Abstract—The Río Sonoyta in northern Sonora is an important aquatic ecosystem that is disappearing because of drought and groundwater withdrawal. Its native species are also threatened by introduced species. 

The only watered reach is an intermittent segment (<1 km, Agua Dulce), found just across the International 

Border from Organ Pipe Cactus National Monument. The native fish present in the river include the endangered Sonoyta pupfish and the indigenous longfin dace. In the wild, the pupfish occurs only at Quitobaquito 

Springs and at Agua Dulce. The longfin dace may be extirpated. A partnership formed and led by the La 

Reserva de la Biosfera El Pinacate y Gran Desierto de Altar included partners from the United States and 

Mexico to create fish refuges in Mexico. We summarize conservation efforts to maintain native fishes in refuge 

ponds, report on their status, and offer suggestions for future management. We also present information 

on the future establishment of refuges for longfin dace and augmentation of pupfish refuges in the United 

States and Mexico. We briefly discuss the impact of a new wastewater treatment plant on the Sonoyta River.

Introduction

	 The native aquatic vertebrates of the southwestern United State

and northwestern Mexico are in trouble. This is particularly true for

native fishes that have been recognized as endangered, threatened,

or rapidly declining for decades, a trend also increasingly observed

in many aquatic invertebrates, amphibians and reptiles(Hershler and

Landye 1988; Miller 1961; Miller and others 2005; Minckley 1973;

Minckley and Deacon 1991; Minckley and Marsh 2009; Rosen and

Melendez 2010). These declines result from effects of introduced

fishes, drought, and groundwater pumping, all of which dramatically affect native aquatic habitats. This paper presents information

on the recent conservation activities implemented for native aquatic

vertebrates of the Río Sonoyta.

Description of the Río Sonoyta

	 The Río Sonoyta basin of Arizona-Sonora covers roughly 3,160

km2 and was formerly a tributary of the Colorado River system.

This former connection was diverted to the Sea of Cortez during

the Pleistocene by the Pinacate lava flows (Donnelly 1974; Miller

and Fuiman 1987). The drainage originates near the Sierra del Pozo

Verde in Mexico, running northwest in Sonora and crossing into the

United States where it turns west as Vamori Wash on the Tohono

O’odham Nation. Vamori Wash then joins San Simon Wash, draining the western Tohono O’odham Nation and joining Sonoran parts

of the basin as it crosses back into Mexico. It forms the Río Sonoyta

near La Nariz. From there the river continues west through Sonoyta

paralleling the border. The river then turns south along the east side

of the Pinacate volcanic shield, passing through the eastern fringe of

the Gran Desierto before reaching the Sea of Cortez east of Puerto

Peñasco.
USDA Forest Service Proceedings RMRS-P-67. 2013 289

Native Aquatic Vertebrates: Conservation and Management in the Río Sonoyta Basin . . . Minckley and others

	 Before the 19th and 20th centuries, perennial surface flow and

cienegas occurred along portions of the river. Channel down-cutting

in the mid 1800s degraded much of the upper drainage due primarily

to groundwater pumping and livestock grazing (Hendrickson and

Varela Romero 1989; Miller and Fuiman 1987; Rosen and others

2010; Shoenherr, 1988).

	 Today,the river continuesthistrend of degradation and desiccation,

still driven by the burgeoning demands for water by an increasing

human population (Hendrickson and Minckley 1985; Miller and

others 2005; Minckley and Marsh 2009). Reduced perennial surface

flow during summer(<1 km)is usually present in the privately owned

Agua Dulce or “Papalote” reach 17 km downstream from Sonoyta,

where flow is maintained by shallow groundwater discharge. The

federal government considers the river channel federal land but has

an agreement with the landowner to provide access to the property.

Río Sonoyta Fishes

	 FishsurveysoftheRíoSonoytahavebeenwelldocumentedstarting

in the 1800s and continuing into the present (McMahon and Miller

1985;Miller and Fuiman 1987;Minckley andMarsh 2009;Rosen and

others 2010; Snyder 1915). The only native fishes collected from this

desert river have been the Sonoyta pupfish (Cyprinodon eremus) and

longfindace (Agosia chrysogaster)(Miller andothers2005;Minckley

and Marsh 2009). The pupfish also occurs at Quitobaquito Spring,

just across the International Border on Organ Pipe Cactus National

Monument (QSWG 2011). The two pupfish populations have been

isolated long enough to differ genetically and are managed separately

(Loftis and others 2009). The Sonoyta pupfish islisted as endangered

by the United States (USFWS 1986).

	 The longfin dace is a widespread and genetically diverse species,

occurring in the Bill Williams, Gila, and Santa Cruz rivers of the

United States, and in the Ríos Santa Cruz, de la Concepción, and

Sonoyta in Sonora, Mexico (Minckley and Marsh 2009). The Ríos

Sonoyta and de la Concepción populations display some overlap or

gradation of morphological characters with populations of the more

southernandeasterly“Agosiasp.” indrainagestothe eastandsoutheast

(Hendrickson 1987; Minckley and Marsh 2009; Tibbets 1993). The

longfin dace is listed as threatened by Mexico (SEDESOL 2010).

	 Records of introduced fishes from the river include mosquitofish

(Gambusia affinis), black bullhead (Amerius melas), andGila topminnow (Poeciliopsis occidentalis) (Bagley 1997; McMahon and Miller

1985; Rosen and others 2010; Snyder 1915). An unidentified tilapia

was reported in 2003 but not collected (Knowles 2010). The blue

tilapia (Oreochromis aureus) has been introduced into the springs

and pond at Ejido Quitovac, an isolated spring southeast of Sonoyta

(this species may have been misidentified as Tilapia zilli in Rosen

and others 2010). There are also commercial fish farms in the drainage near the town of Sonoyta, which reportedly raise some species

of tilapia that have not yet been sampled or identified by any of the

authors.

	 Recently, riverine populations of the Sonoyta pupfish have plummeted. Longfin dace, abundant in the system asrecently as 2005, now

appear extirpated from the river and have not been collected there

since 2008 (Minckley and Izaguirre Pompa 2010; Rosen and others

2010).

Río Sonoyta Amphibians and Reptiles

	 There are six species of anuran amphibians (toads) in the Río

Sonoyta valley and a population of endemic Sonoyta mud turtle

Kinosternon sonoriense longifemorale, a candidate speciesforlisting

under the U.S. Endangered Species Act (Knowles and others 2004;

Rosen 2007; Rosen and others USFWS 2011). Little is known about

the amphibians of this river or their use of it.

	 The Sonoyta mud turtle was once widespread in the Río Sonoyta

watershed and is still common today at Quitovac and Quitobaquito

Springs (Rosen and others 2010; Rosen and Melendez 2010). However, since downcutting of the cienega at Sonoyta and the advent of

increased water pumping in the 1970s, populations of this endemic

turtle have declined markedly. Currently, a population can be found

in the sewage lagoon and associated wastewater effluent discharge

from the town of Sonoyta (Rosen and others 2010). Populations of

this turtle also occur at the Presa Xochimilco dam site in Sonoyta

and in the wetted portion of Río Sonoyta at Agua Dulce (Rosen and

others 2010). Small numbers of individuals have been found in the

river at Santo Domingo and within 4 km upstream of Sonoyta.

Conservation Actions

	 The future plight of native fishes in the arid southwest was recognized by ichthyologists working on native fishes in the region by the

1940s (Minckley and Deacon 1991). The first recovery actions were

also carried out by these individuals who moved fish they thought

were in danger of extirpation to sites where they were protected from

immediatelyobviousthreats andbelievedlikelytosurvive.Beginning

in the 1960s and continuing to the present, government agencies have

developed recovery programs that include natural sites and artificial

refuges for listed fish species (Section V, Minckley and Deacon eds.

1991). By 1993, the Desert Pupfish Recovery Plan (USFWS 1993)

listed over 100 attempts at establishing refuges,including theSonoyta

(Quitobaquito) pupfish (USFWS 1993). The U.S. Fish and Wildlife

Service received internal funding to build pupfish refugesin northern

Sonora in2006.TheUSFWSPreventingExtinctiongrantwasdeemed

necessary due to the on-going drought and groundwater pumping

from the Río Sonoyta, which threatened to extirpate the Río Sonoyta

population of the pupfish.

	 The design of the refuge ponds constructed in Sonora is based on

the ideas and plans developed by individuals and agencies working

on native aquatic vertebrate’s recovery in Arizona. The ponds are

lined with a Geo-PadTM polypropylene felt underliner covered by

a 45 mil EPDM PondguardTM liner, and are 3 by 4 M in area, hold

~ 5600 L of water and are ~1 M deep. Each pond has an inflow that

passes through a short stream habitat before entering the main pond.

Water is recirculated and leaves the pond through a filtered outflow,

which is maintained using an electric pump. Pond substrate is fine

sand and gravel. The refuge ponds were designed to accommodate

both native fish species although the stream habitat was designed

specifically to provide spawning habitat for the longfin dace.

	 Fish used to stock the ponds were obtained in 2007 from the Agua

Dulce reach of the Río Sonoyta by seining and netting. Particular

emphasis was placed on sorting the fish during capture and before

stocking to prevent introduction of mosquitofish into refuges. Both

species were collected and placed into coolers containing clean aerated water. Marine salts and a commercial conditioner were added

to the water in all of the transport coolers to reduce the stress caused

by handling and transport. The fishes were transported directly to

the ponds, gradually acclimated to the pond water temperature, and

stocked.

	 By 2011, five refuges were established and stocked with pupfish

and longfin dace. In 2007, two refuge ponds were constructed at the

Pinacate Biosphere Reserve headquarters and at the Centro Intercultural de Estudios de Desiertos y Oceanos(CEDO) in Puerto Peñasco.
290 USDA Forest Service Proceedings RMRS-P-67. 2013

Minckley and others Native Aquatic Vertebrates: Conservation and Management in the Río Sonoyta Basin . . .

Fish were also stocked into the springs at Ejido Quitovac. The fourth

refuge was completed and stocked in 2008, at the Colegio de Bachilleres del Estado de Sonora high school (COBACH) in the town of

Sonoyta (López Méndez 2011); this pond also contains aRío Sonoyta

mud turtle (introduced by students at the school). A fifth refuge was

constructed in 2011 at the new Pinacate Biosphere Visitors Center

by staff.

	 Conservation activities for the Sonoyta mud turtle are centered on

salvage of turtles at Sonoyta from effluent waters that are, or will be

desiccated as the antiquated sewage system of the town is updated.

A cooperative agreement involving the Municipio of Sonoyta, local ejidos, the U.S. EPA (which is providing partial funding), and

SEMARNAT, the Mexican national conservation authority, includes

conservation considerations in the development of a new sewage

treatment plant for the town of Sonoyta (Rosen 2009). This facility

will have one or more ponds that will provide turtle habitat, which

when combined with habitat created in the river channel by effluent

discharge from the plantshould allow thisspeciesto persist in Sonoyta.

Once the sewage plant is operational the turtles present at the current

sewage lagoonandnearPresaXochimilcowillbe salvagedandmoved

to the new facility. Water in the new effluent reach of Río Sonoyta

may be suitable for establishment of native fishes as well (USEPA

2008).

Summary

	 Construction of refuge ponds for Río Sonoyta native fishes has

been successful to date, and appears to have successfully secured

small populations of the Río Sonoyta forms of Sonoyta pupfish and

longfin dace.Today, pupfish are present in all ofthe ponds and a small

population of longfin dace persists at the refuge at COBACH. As the

water levelsin Río Sonoyta continue to be unreliable—it is generally

dry through most of its length and the remaining wetted portion has

gone nearly or completely dry during recent droughts—these refuges

are necessary to continue to provide a place to maintain these species

for the foreseeable future. These habitats lack careful management

and remain precarious due to silting in by blowing sand, maintenance

issues, water supply issues, and funding for replacement of parts. If

flows are returned to the river these refuge populations can be used

to re-establish populations in their historical habitat.

	 In addition to construction of the refuge pond at COBACH, development of a curriculum using the pond as a research and study

tool is in progress. Plans are to have the students explore different

aspectsof conservation, ecology, andaquaticbiologyusingahands-on

approach, while managing their pond for the security and continued

existence of endangered and threatened species. During the past year

students have given presentations at the annual meeting of the Desert

FishesCouncil in Hermosillo, Sonora and at the 2012 Sonoran Desert

Symposium.Futureplansinclude communityoutreachtofurtherraise

awareness about the plight of native aquatic vertebrates and the Río

Sonoyta.

	 There are also tentative plans to move between 200 and 500 individuals of both native fish speciesto refugesin the United States. The

Sonoyta pupfish from Mexico would be used to provide immigrants

into genetically bottlenecked populations of this species that were

established in the United Statesin 1976 (AGFD 2001).Today, current

refuges such as the one at Organ Pipe Cactus National Monument

introducewildindividualsintotheir artificialrefugepondstomaintain

genetic diversity. Longfin dace from the refuges would be used to

establish a refuge population at the Arizona-Sonora Desert Museum

as a hedge against extinction in Mexico.

References 

ArizonaGame and FishDepartment. 2001.Quitobaquito pupfish,Cyprinodon 

eremus. Unpublished abstract compiled and edited by the Heritage Data

ManagementSystem,ArizonaGame andFishDepartment,Phoenix,AZ.4p.

Bagley, B. 1997. Desert pupfish collections in Baja California and Sonora,

Mexico. January 10-17, 1997. Trip Report, Arizona State University,

Tempe, AZ. 13 p.

Donnelly,M.F.1974.GeologyoftheSierradelPinacatevolcanicfield,northern

Sonora, Mexico and southern Arizona, U.S.A. Unpubl. Ph.D. dissertation,

Stanford University, Stanford, CA.

Hendrickson, D. A. 1987. Geographic variation in morphology of Agosia 

chrysogaster, a Sonoran Desert cyprinid fish. PhD dissertation, Arizona

State University, Tempe, AZ.

Hendrickson, D. A., and W. L. Minckley. 1985. Cienegas: Vanishing climax

communities of theAmerican Southwest. Desert Plants(Boyce Thompson

Arboretum, Superior, AZ) 6 no 3 (1984): 131-175.

Hendrickson, D. A., and A. Varela-Romero. 1989. Conservation status of

desert pupfish, Cyprinodon macularius, in Mexico and Arizona. Copeia

1989:478-483.

Hershler, R., and J. J. Landye. 1988. Arizona Hydrobiidae (Prosobranchia:

Rissoacea). Smithsonian Contributions to Zoology, Number 459:1-63.

Knowles, G. 2010. E-mail to C. O. Minckley documenting the observation of

a tilapia in the Río Sonoyta in the Agua Dulce reach on 15 October 2003.

Knowles, G. W., J. Rorabaugh, R. Paredes Aguilar, P.C. Rosen, D.H. Hall,

and D. Riedle. 2004. Distribution, status and conservation of the Sonoyta

mud turtle (Kinosternon sonoriense longifemorale). Hendrickson, D. A.,

and L. T. Findley, eds., Proc. of the Desert Fishes Council Vol XXXVII

(2002):28-29.

Loftis, D. G., A. A. Echelle, H. Koike, R. A. Van Den Bussche, and C. O.

Minckley. 2009. Genetic structure of wild populations of the endangered

Desert Pupfish complex (Cyprinodontidae: Cyprinodon). Conservation

Genetics. 10:453-463.

López Méndez, J. 2011. Refuge populations of Río Sonoyta pupfish (Cyprinodon eremus) and longfin dace (Agosia chrysogaster) in COBACH

Campus Sonoyta. Presentation to 43 Annual Meeting of the Desert Fishes

Council, Hermosillo, MX.

McMahon,T. E., andR.R. Miller. 1985. Status ofthe fishes oftheRío Sonoyta

Basin, Arizona and Mexico. Pister, E. P. ed., Proceedings of the Desert

Fishes Council XIV (1982): 237-245.

Miller, R. R. 1961. Man and the changing fish fauna of the American Southwest. Pap. Michigan Acad. Sci., Arts, Lett. 46: 365-404,

Miller, R.R., and L. A. Fuiman. 1987. Description and conservation status of

Cyprinodon macularius eremus, a new subspecies of pupfish from Organ

Pipe Cactus National Monument, Arizona. Copeia 1987: 593-609.

Miller, R.R., W.L. Minckley, and S.M. Norris. 2005. Freshwater fishes of

Mexico. University of Chicago Press, Chicago, Illinois. 490 p.

Minckley, C. O. and I. S. Izaguirre Pompa. 2010. Fishes of the Río Sonoyta—

Past, Present, Future? Presentation at Sonoran Desert Symposium, Ajo,

AZ. March 2010.

Minckley, W. L. 1973. Fishes of Arizona. Arizona Game and Fish Department, Phoenix, Arizona.

Minckley, W. L., and J. E. Deacon, eds. 991. Battle against extinction: native fish management in the American West. University of Arizona Press,

Tucson, AZ.

Minckley,W. L., and P.C. Marsh. 2009.Inland fishes ofthe greatersouthwest:

chronicle of a vanishing biota. University of Arizona Press, Tucson, AZ.

QuitobaquitoSpringsWorkingGroup.2011.DRAFTConservationagreement

and rangewide conservation assessment and strategy for the Sonoyta mud

turtle, Sonoyta pupfish, Quitobaquito spring snail, and longfin dace in

Quitobaquito and Río Sonoyta.

Rosen, P. C. 2007. The amphibian and reptiles of the dry borderlands of

northwestern Mexico and southwestern Arizona. Pages 310-337 in Felger,

R. S., and B. Broylesm, eds., Dry Borderlands, University of Utah Press,

Salt Lake City, UT.

Rosen, P. C. 2009. Summary of water quality, quantity, and habitat location

needs for native fish and turtles associated with new wastewater treatment
USDA Forest Service Proceedings RMRS-P-67. 2013 291

Native Aquatic Vertebrates: Conservation and Management in the Río Sonoyta Basin . . . Minckley and others

infrastructure at Sonoyta, Sonora, Mexico. School of Natural Resources,

University of Arizona, Tucson, AZ.

Rosen, P. C., and C. Melendez. 2010. Observations on the status of aquatic

turtles and the occurrence of ranid frogs and other aquatic vertebrates in

northeasternMexico. Pages 205-224 inHalvorson,W.,C.Schwalbe, andC.

Van Riper III, eds., Southwestern Desert Resources, University ofArizona

Press, Tucson, AZ. 360 p.

Rosen, P.C., C. Melendez, J.D. Riedle, A. C. Pate, and E. Fernandez. 2010.

EcologyandConservationintheSonoytaValley,Arizona andSonora.Pages

143-160 in Halvorson, W., C. Schwalbe, and C. Van Riper III, eds., SouthwesternDesertResources,UniversityofArizonaPress,Tucson,AZ.360pp.

Schoenherr, A. A. 1988. A review of the life history and status of the desert

pupfish, Cyprinodon macularius. Bull. So. Calif. Acad. Sci. 87:104-134.

SEDESOL. 2010. NORMAOficial Mexicana NOM-059-SEMARNAT-2010.

Protecciónambiental-EspeciesnativasdeMéxicodeflorayfauna silvestresCategorías de riesgo y especificaciones para su inclusión, exclusión o

cambio-Listade especies en riesgo.

Snyder, J. O. 1915. Notes on a collection of fishes made by Dr. Edgar A.

Mearns from rivers tributary to the Gulf of California. Proceedings, U.S.

National Museum 49 No 2125.

Tibbets, C. A. 1998. Evolutionary genetics of Agosia chrysogaster. PhD dissertation. Arizona State University, Tempe, AZ.

U.S. Environmental Protection Agency. 2008. Environmental assessment:

improvements to the wastewater collection and treatment systems for the

City of Sonoyta, Sonora, Mexico. Region IX San Francisco, CA. 50 p

U.S. Fish and Wildlife Service.1986. Endangered and threatened wildlife and

plants:Determination of endangered status and critical habitatforthe desert

pupfish. Federal Register 51 (61):10842:10850l.

U.S. Fish and Wildlife Service. 1993. Desert pupfish recovery plan. U.S. Fish

and Wildlife Service, Phoenix, AZ. 67 p.

U.S. Fish and Wildlife Service. 2011. Endangered and threatened wildlife

and plants; review of native species that are candidates for listing as

endangered or threatened; annual notice of findings on resubmitted petitions; annual description of progress on listing actions. Federal Register

76(207):66370- 66439.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
292 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Documenting the Biodiversity of the Madrean 

Archipelago: An Analysis of a Virtual Flora and Fauna

Nicholas S. Deyo and Thomas R. Van Devender

Sky Island Alliance, Tucson Arizona

Alex Smith

School of Natural Resources and the Environment, University of Arizona, Tucson Arizona

Edward Gilbert

SEINet Flora Database, Tucson, Arizona

Abstract—The Madrean Archipelago Biodiversity Assessment (MABA) of Sky Island Alliance is an ambitious project to document the distributions of all species of animals and plants in the Madrean Archipelago, 

focusing particularly on northeastern Sonora and northwestern Chihuahua, Mexico. The information is 

made available through MABA’s online database (madrean.org). The sources of these records are museum 

collections, herbaria, scientific literature, other databases, observations, and field notes. Many observations 

come from MABA expeditions to isolated mountain ranges in northeastern Sonora involving taxonomic 

specialists, professors, students, and others from universities in Arizona and Sonora; and personnel from 

Mexican agencies. This paper provides an analysis of the current status of species records contained within 

the MABA flora and fauna databases and the significance of these records for future collection, research, 

and conservation planning. 

Introduction

Project Background

	 Since 2009, the Sky IslandAlliance’s MadreanArchipelagoBiodiversity Assessment (MABA) has been documenting the biodiversity

of the Sky Island region with the aim of supporting conservation and

scientific inquiry.The MABAproject is a direct result of the Madrean

Conferences held in 1994 and 2004. These conferences convened

scientists and natural resource managers from the United States and

Mexico to discussthe biodiversity and naturalresourcesmanagement

of the Sky Island region (DeBano and others 1995; Gottfried and

others 2005). One conclusion from these conferences wasthe lack of

biodiversity data needed to supportthemanagement and conservation

of the Madrean Archipelago, particularly in Mexico. Documenting

the biodiversity of the Sky Island region is challenging, as the area

contains over 50 mountain ranges/mountain complexes, hundreds of

rare and localized species, and numerous habitats dispersed between

the Rocky Mountains to the north, the Sierra Madre Occidental to

the south, and the Sonoran and Chihuahuan deserts to the west and

east. Sky Island Alliance worked with programmer Ed Gilbert, the

SEINet flora database, Symbiota, and the Veolia Foundation to create the MABA online database (http://www.madrean.org); since its

creation, the database has become a large repository of information

for many groups of species in the Sky Island region and includes

both vouchered specimens and observations from herbaria, museum

collections, scientific literature, and MABA’s own field expeditions.

This paper provides an analysis of the current status of species

records contained within the MABA flora and fauna databases and

the significance of these records for future collection, research, and

conservation planning.

Virtual Flora and Fauna

	 TheMABAonlinedatabase is considereda “virtualflora andfauna,”

a type of digital resource that is revolutionizing the way biodiversity

information is documented and shared. Floras and faunasreferto both

the species of plants and animals that inhabit a region as well as the

publications that describe them (Heidorn 2004). Historically, floras

and faunas have taken the formof printmedia and contain information

such as scientific classifications, species descriptions, illustrations,

photos, distribution maps, and identification keys. There are limitationsto printedmedia when it comesto documenting biodiversity, not

least of which are the sheer number of organisms in some taxonomic

groups; given that there are an estimated 5,300 species of flowering

plants in Madrean Pine-Oak Woodlands (Conservation International

2011), a print flora for the entire region would be impractical. Additionally, the cost and labor associated with publishing regional floras

and faunas also limit production and distribution of these works; in

2003 the estimated cost of producing a flora with 2,104 species was

$1,579,946 (Heidorn 2004).
USDA Forest Service Proceedings RMRS-P-67. 2013 293

Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna Deyo

	 Web-based virtual floras overcome many of these problems: they

are less expensive, support unlimited images and species records,

allow for complex data queries, and can be corrected and updated

continually (Gilbert, unpublished presentation).Web-based flora and

fauna have taken databases, once the private domain of individual

researchers, and have extended them to be a medium for collaboration between researchers and citizen scientists (Schnase and others

1997). Finally, web-based floras and faunas increase the speed at

which biodiversity data is collected and made available, a critical

factor considering how quickly species and habitats are being lost to

human activities (Heidorn 2004).

	 The last two decades have seen a worldwide effort to digitize herbaria and museum collections, resulting in flora and fauna databases

containing on the order of 104 and 106 records (Soberón and others

2002).Accesstosuchlargedatasetshasspurredtheburgeoningfieldof

biodiversity informatics(Soberón and Peterson 2004), and has led to

scientific applicationssuch asmodeling speciesranges and evolutionary analyses (Petersen and others1999; Zhong 1999); locating areas

for biodiversity exploration (Jones and Gladkov 1999; Soberón and

others 2004); prioritizing for protected areas (Godown and Peterson

2000; Kelley and others 2002), and modeling routes for the spread

of invasive species (Higgins and others 1999; Peterson 2004).

	 No matter how sophisticated the design of a virtual flora orfauna, it

isonlyasgoodasthedata it contains.Adatabasewitha comprehensive

set of species records allows for comparison of species lists across

sites; observation ofspeciesrichness over time, including before and

after habitat modifications; and analyses such as those mentioned in

the previous paragraph (Soberón and others 2004). Adding records

to the MABA database has been ongoing since the program began in

2009. Thisresearch reflectsthe status of the MABAdatabase through

analyzing species records in reference to Sky Islands/complexes,

reporting the sources of records, and identifying which time periods

have been the most productive for collection efforts. The results of

this paper are useful for targeting areas for future collection and data

acquisition efforts and better understanding how the MABAdatabase

can be used for research and conservation planning.

Methods

Defining Sky Island Ranges and Complexes

	 In order to perform count and density analyses on species data for

each Sky Island/complex, a GIS polygon layer was created defining

the physiographic boundaries of each range or complex containing

oak woodland. Generally, the boundaries of ranges or complexes

were digitized using physiographically defined mountainousfeatures

containing contiguous or closely clustered areas of oak woodland that

were isolated from the main Sierra Madre and Rocky Mountains.

Previously created maps of the region have classified Sky Islands

exclusively on the presence of vegetation (Marshall 1957; Brown

and Lowe 1980). Range/complex borders were defined by drawing along a 7 percent slope gradient threshold (see fig. 1). In areas

of ambiguous slope, borders were drawn to include 50 percent or

greater digital elevation model (DEM) cells with a 7 percent slope

gradient or higher. These boundaries emphasize the physiographic

continuity of complexes or ranges, including areas outside of the

oak woodland boundaries that are still mountainous. Physiographic

ranges and complexes were subdivided when they showed significant

gaps in oak woodland presence. These divisions were drawn along

low points of elevation in those gaps. Additionally, two ranges that

did not show oak woodlands in the Geographic Information System

(GIS) vegetation database used in this analysis, the Mule Mountains

and theCerroCobachi, were included based on the field experience of

Dr.TomVan Devender.Asmall number of ranges and complexesthat

show oak woodland connections to the Sierra Madre were included

in this analysis because of historical precedent. These “peninsulas”

of the Sierra Madre are noted in table 1.

	 Names of complexes were determined by combining the northernmost and southernmost range names. Range names were acquired

from U.S. Geological Survey (USGS, 2002) and Mexican National

Institute of Statistic and Geography (INEGI) topographic maps. In

some cases, a more commonly known name was chosen over the

naming convention above.Asingle unhyphenated name was used for

isolated ranges and smaller subdivisions of Sky Island complexes.

Figure 2 provides a map of the Sky Island ranges/complexes used in

this analysis.

Data Acquisition and Manipulation

	 Species point locations and associated data were obtained by SQL

queries to the separate MABA flora and fauna databases. Queries

were made using a web-based MySQL interface to facilitate downloading large datasets; however, the same data can be accessed using

the MABA online database (http://www.madrean.org). Latitude and

longitude coordinatesboundedqueriestoa regioncontainingArizona,

New Mexico, Sonora, and Chihuahua. Records lacking geographic

coordinateswere eliminated.Eachacquireddatasetwasthenimported

into a GIS and projected into the Albers Conformal Conic projection

using the NAD 83 datum. A spatial join was then created to link species data with Sky Islands/complexes boundaries.

	 Queries were made for the six separate classes of animals: Arthropoda, Aves, Mammalia, Osteichthyes, Amphibia, and Reptilia.

Amphibia andReptilia classeswere then combined.For plantrecords,

the entire database was downloaded and classified by family name

into three separate datasetsfor angiosperms, gymnosperms, and ferns

Figure 1—This image shows areas above 7 percent grade (white) that were 

used to help digitize the boundary of Sky Island ranges and complexes.
294 USDA Forest Service Proceedings RMRS-P-67. 2013

Deyo Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna

Table 1—Species records by Sky Island or Sky Island complex.
USDA Forest Service Proceedings RMRS-P-67. 2013 295

Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna Deyo

Figure 2—Map depicting the Sky Island ranges and complexes used to the distribution of species record data in the Madrean Archipelago. 
296 USDA Forest Service Proceedings RMRS-P-67. 2013

Deyo Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna

and fern allies. Observations of members of the families Asteraceae,

Poaceae, and Fabaceae were then isolated from the angiosperm data.

Analysis of contributing institutions and record collection dates was

performed using data downloads from separate SQL queries. This

separate analysis contained some discrepancies between the two

querying efforts. This may be attributed to the addition of new sets

of data since the first queries were performed.

	 The Nature Conservancy’s Apache Highlands Ecoregion conservation priorities analysis provided vegetation data for Arizona, New

Mexico, and northern Sonora and Chihuahua (Marshall and others

2004). This data combined Gap Analysis Program (GAP) vegetation

data for areas in the United States with Forest Inventory 2000 layers

for areas in Mexico. As this data terminates in the middle of Sonora,

data for more southern regions was acquired from an INEGI layer

entitled “uso delsuelo y vegetación” (2005).Additionally, a digitized

layer of Brown and Lowe’s Biotic Communities of the Southwest

classification map (1980) was used to supplement the more modern,

digitally mapped layers. Where oak woodland data differed between

sources,the combined area of oakwoodlandsin alllayersforthe given

region was used to define Sky Islands/complexes. Digital elevation

model (DEM) data used to create the slope analysislayer is a stitched

mosaic of 90 m resolution raster imagery from NASA’s 2000 Shuttle

Radar Topography Mission (USGS 2002).

Results

General Statistics 

	 Atotalof596,681plant and121,355animalrecordswithgeographic

coordinates were queried from the MABA flora and fauna databases

for the States of Arizona, New Mexico, Sonora, and Chihuahua. Of

these records, 87,134 plant and 13,278 animalrecords were identified

within the Sky Island/complex polygons created for this analysis.

These figures represent only 14 percednt of all records within the

states mentioned. However, as the Sky Islands/complexes only

comprise 4 percent of the area of those states, these figures show a

greater number of records within Sky Islands/complexes. Across all

Sky Islands/complexes, the average sampling effort (the number of

records/area in km2)for plantswas 2.18 records/km2 and 0.22 records/

km2 for animals, representing a tenfold difference in sampling effort

between the MABAflora and fauna databases. When sampling effort

was compared between Sky Islands/complexes in Mexico versus

the United States, the results differed between the animal and plant

records. Note that the country label of Sky Islands/complexes that

straddled the border refers to the location of the polygon’s centroid.

Plants were better represented in the United States with an average

sampling effort of 3.98 records/km2 compared with 0.87 records/km2

inMexico.AnimalswerebetterrepresentedinMexicowithanaverage

sampling density of 0.35 records/km2, while the United States had

an average of 0.04 records/km2. Table 1 provides an overview of the

data generated in this analysis.

Contributing Institutions

	 Thedatabaseswereanalyzedtobetterunderstandwherethemajority

ofrecordsoriginated.Twenty-twoinstitutionscontributedtotheMABA

flora database. The University of Arizona Herbarium contributed the

most records, followed by the Arizona State University Herbarium,

MABA observations, Sonoran Desert Plants: An Ecological Atlas,

and the Desert Botanical Garden. Thirty-six institutions contributed

to the MABAfauna database. MABAcontributed 13,198 records, by

far the greatest number of observations, followed by the University

of Arizona, Arizona State University, eBird, and the University of

Texas atArlington. Figure 3 showsthe distribution of records among

contributing institutions.

Temporal Distribution of Records

	 Examining collection dates for species records revealed that both

the MABA flora and fauna databases were right skewed (fig. 4)

toward more modern collection dates. Seventy-five percent of all

plant records were collected after 1962 while the same percentage

of animal records was collected after 1976. The earliest plant record

in the database was an Acacia texensis collected by G. Thurber in

1851. The earliest animal record was a Rana yavapaiensis collected

by Edgar A. Mearns in 1893.

	 Within the MABA flora database, most ranges/complexes show

recent collection efforts; however, it appears that the Cerro Cobachi,

Sierrita, Big Hatchet, El Maviro, Las Calabazas-Martinez, and Santo

Niño ranges have not seen collection efforts since the late to mid1990s. Fewer ranges within the MABA fauna database show recent

collection efforts; the Sierrita, Tortolita, Pinaleño, Cerro El Tiznado,

Mule,CerroCobachi, andAnimasrangesdonotshowanynewrecords

since the late 1990s, and the Sierrita, Tortolita, and Pinaleño ranges

do not show any records since the late 1970s. It should be noted that

the Sierrita range only showed two animal records revealed in a later

query of the database. The statistical mode of record collection dates

was calculated for each individual Sky Island/complex for flora and

fauna recordsseparately. For plant records, 14 of the 33 Mexican Sky

Islands/complexes had modes more recent than 2000, while only 4 of

the 22 United States Sky Islands/complexes had modes more recent

than 2000. This indicates that sampling efforts in Mexico have been

more vigorous recently than those in the United States. For animal

records, 12 of the 33 Mexican Sky Islands/complexes had modes

more recent than 2000, while only 3 of the 22 United States Sky

Islands/complexes had modes more recent then than 2000. As with

plant records, this shows that the collection effort for Mexico has

been more vigorous recently.

Records by Taxa

	 The databases were queried based on major taxonomic groups to

better understand which groups were bettersampled; conversely, this

analysis also revealed which Sky Island ranges/complexes have the

fewest number of records. In terms of overall records (both animals

andplants),theHuachuca-Patagoniacomplexyieldedthemostrecords

with 11,104 observations. Not surprisingly, other ranges/complexes

with large numbers of records were the Dos Cabezas-Chiracahuas

(10,881 records), the Santa Catalinas(9,994 records), the PeloncilloPan Duros (7,512 records), and the Santa Ritas (6,748 records). The

most poorly represented ranges/complexeswereBatamote,Enmedio,

San Juan, Cerro Cobachi, and Cerro El Tiznado. Of these, Batamote

and Enmedio had no records and the remainder all had fewer than

50 records.

	 The animal taxa queried in the MABA fauna database were Arthropoda,Aves, Mammalia, Osteichthyes(bony fish), andAmphibia/

Reptilia. There were more records for Aves than for any other class

of animals with 8,690 records,followed byAmphibia/Reptilia (2,326

records),Arthropoda (1,683 records), Osteichthyes(362 records) and

Mammalia (217 records).The top five ranges/complexesforArthropoda were the LosAjos-La Madera, Peloncillo-Pan Duro, El Gato, San

Javier, and SanAntonio-Azul. Of the 55 ranges/complexesidentified
USDA Forest Service Proceedings RMRS-P-67. 2013 297

Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna Deyo

Figure 3—Major contributing institution for the MABA fauna database (a) and the MABA flora database (b). 

Note: not all contributing institutions are depicted here.

here, 40 have 10 records or fewer. The top five ranges/complexes for

Aves were the Peloncillo-Pan Duro, Los Ajos-La Madera, El Tigre,

San Antonio-Azul, and Mariquita-Elenita. While Aves was better

represented than othertaxa, 29 ranges/complexesremainwithout bird

records. For Amphibia and Reptilia combined, the best-represented

ranges/complexes were El Tigre, Peloncillo-Pan Duro, Los Ajos-La

Madera, El Pinito, and La Huerta. Thirty-eight ranges/complexes

had 10 or fewer records for reptiles and amphibians. Osteichthyes

and Mammalia showed very few records, so comparisons between

ranges are excluded here (see table 1).

	 The plant groups queried in the MABA flora database were the

divisions for angiosperms, gymnosperms, and ferns and fern allies.

Angiosperms were further divided into the familiesAsteraceae, Poaceae, and Fabaceae. All divisions had several thousand records each.

Angiosperms were best represented with 81,861 records (expected

since angiosperms include many more species than other groups),

followed by ferns and fern allies (3,135 records) and gymnosperms

(2,183 records). The angiosperm families were represented with

13,458 Asteraceae, 10,361 Poaceae, and 8,996 Fabaceae. Gymnosperms were least well represented with 34 ranges/complexes having

fewer than 10 records, while ferns and fern alliesreflected 31 ranges/

complexes with fewer than 10 records. Angiosperms had at least 10

records in all ranges except 3: San Juan (2 records), Batamote, and

Enmedio. Of the 55 Sky Island ranges and complexes, only 9 had

records for each taxonomic group (both animals and plants); these

were the Huachuca-Patagonias, Atascosa-Cibutas, Peloncillo-Pan

Duros,Los-Ajos-LaMaderas,Galiuros,SanAntonio-Azuls,ElTigre,

El Gato, and Aconchi.

Discussion

	 Biodiversity databases that are “extensive and exhaustive” are

important because they allow for the application of biodiversity

data to basic research and conservation planning purposes (Soberón

and others 2004). This analysis was designed to help the Sky Island

Alliance’s MABAprogrambetter understand the strengths and weaknesses of its flora and fauna biodiversity databases, and to inform

the next steps to improve the coverage and quality of species data. It

is also an opportunity to consider how these data can be applied for

conservation planning purposes.

	 Specimen records are typically biased by the spatial and temporal

distributionof collectionefforts(SoberónandPeterson2004;Soberón

and others 2004). Biases in species location data are often attributed
298 USDA Forest Service Proceedings RMRS-P-67. 2013

Deyo Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna

Figure 4—The distribution of species records collection dates for 5-year intervals for the MABA fauna 

database (a) and the MABA flora database (b).

to proximity to roads, research stations, educational institutions, land

tenure, and cities (Soberón and others 2000). This analysis revealed

that many Sky Islands and complexes have no or very little species

data; only nine ranges contained records for all of the taxonomic

groups queried here. Additionally, there remains a large disparity in

sampling effort between the United States and Mexico. Plants are

better represented in the United States by a factor of nearly five to

one, which islikely due to actualsampling effort because of the large

network of herbaria that have provided data.Animalrecords are better

represented in Mexico by a factor of eight to one. This discrepancy

along with the overall better representation of plants in the database

indicates that a large number of existing animal records in museum

collections are not included in the MABA database. Therefore, adding additional animal collections from museums and other sources

to the MABA database should be a priority. Additionally, any Sky

Islands/complexes lacking in species data across multiple groups of

taxa should be considered for future field collection efforts.

	 Biodiversity databases rely on the species collections contained in

museums and herbaria. Natural history data may become antiquated

and less useful for contemporary applications if collections are not

continually updated (Peterson and others 1998; Winker 1996). At

first glance, the distributions ofrecord collection dates appeared to be

relatively contemporary, with 75 percent of records being collected

in the last 50 years. However, this does not account for biases in the

collection of individual taxa, which may occur when institutions

receive funding to study specific taxonomic groups or when certain

taxa are treated with particularinterest in a given period (Soberón and

others 2004). Individual Sky Islands/complexes that do not contain

records from recent years are due for new collection efforts.

	 The importance of continuing the collection and compilation of

species data cannot be understated. Species records from herbaria,

museum collections, and other sources are the foundation of our understanding of the biodiversity of the planet and how to conserve it

(Winker 1996).While there are remote sensing methodsfor assessing

areas for high biodiversity and conservation priority, the failure to

include speciesdata inthese analysesmayleaveunique andvulnerable

speciesunprotected(Brooks andothers2004).Furthermore,increased

accessto species data through online databaseslike MABA’s, as well

asthe huge increase in othersources of environmental data, hasled to

innovative methods of modeling species distributionsforresearching

biodiversity and conservation planning purposes (Lobo and others

1997;Jones and Gladkov 1999; Soberón and others 2004;Rebelo and

Siegfried 1992; Csuti and others 1997; Godown & Peterson 2000;

Peterson and others 2000; Kelley and others 2002;Burgess and others

2002; Chen & Peterson 2002). Combining species data with data on

climate, physiography, and other characteristics allowsthe modeling

of ecological niches (Soberón and Peterson 2004). The use of these

methods would help overcome some of the gaps and biasesin species

data now available for the Sky Island region and contribute to a better

understanding of conservation priorities in the region.

	 However, before data can be used for the applications mentioned

above its quality must be assessed. Large mixed source databases
USDA Forest Service Proceedings RMRS-P-67. 2013 299

Documenting the Biodiversity of the Madrean Archipelago: An Analysis of a Virtual Flora and Fauna Deyo

such as MABA’s often face difficulties in maintaining data quality.

The errors in most need of review are those related to taxonomy and

incorrect georeferencing. Taxonomic errors are somewhat mitigated

by taxonomic thesauruses that check for antiquated scientific names,

a continual challenge in a world of rapidly changing taxonomy. The

correction of georeferencing errors is more laborious and requires

cross-referencing species’locality descriptions with their geographic

coordinates (Soberón and others 2002); although there are some

statistical tools available to help with this process (Chapman and

Busby 1994). Some assurance of quality control would improve the

confidence with which researchers and lay people could use MABA

data. Without quality control measures,

	 TheMABAprojecthasmadegreatstridesincompilingbiodiversity

data within the Madrean Archipelago, both from existing collections

and from field collection efforts. There still remain numerous gaps

in the biodiversity data available for the region, further emphasizing the need for more collection efforts. Despite the lack of data for

specific areas, there remains a great deal of data that is available for

conservation and research efforts. Understanding the biodiversity of

the Madrean Archipelago will help highlight the uniqueness of the

region and protect its species for future generations.

References

Brooks, Thomas; Fonseca, Gustavo; Rodrigues, A. 2004. Species, data, and

conservation planning. Conservation Biology 18 (6): 1682-1688.

Brown, David E.; Lowe, Charles H. 1980. Biotic communities of the Southwest. Gen. Tech. Rep. RM-78 (map) [Revision 1983]. Fort Collins, CO:

U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest

and Range Experimental Station.

Burgess,N.D.;Rahbek,C.;Williams,P.;Larsen,F.W; and Balmford,A. 2002.

How much of the vertebrate diversity of sub-SaharanAfrica is represented

by recent conservation proposals? Biological Conservation. 107: 327-339.

Chapman,A.;BusbyJ.R.1994.Linkingplantspeciesinformationtocontinental

biodiversity inventory, climate modeling and environmental monitoring.

In: R.I. Miller (editor) Mapping the diversity of nature. Chapman & Hall,

London.

Chen, G.; Peterson,A.T. 2002. Prioritization of areasin China for biodiversity

conservation based on the distribution of endangered bird species. Bird

Conservation International. 12: 197-209.

ConservationInternational.2011.Madreanpine-oakwoodlands.Conservation

International Website. Online: http://www.conservation.org/where/priority_areas/hotspots/north_central_america/madrean-pine-oak-woodlands/

Pages/default.aspx. [2011].

Csuti, B.; [and others]. 1997. A comparison of reserve selection algorithms

using data on terrestrial vertebrates in Oregon. Biological Conservation.

80: 83-97.

DeBano, Leonard H.; Ffolliott, Peter F.; Ortega-Rubio,Alfredo.; [and others],

tech. coords. 1995. Biodiversity and management of the Madrean Archipelago: The sky islands of southwestern United States and northwestern

Mexico. 1994 Sept. 19-23; Tucson, AZ. Gen. Tech. Rep. RM-GTR-264.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Forest and Range Experiment Station. 669 p.

Gilbert, Edward. 2011. Symbiota: Specimen-based virtual flora software

platform. Unpublished presentation on file at: Sky Island Alliance office,

Tucson, AZ. 45 p.

Godown, M.E.; Peterson, A.T. 2000. Preliminary distributional analysis of

U.S. endangered bird species. Biodiversity Conservation. 9: 1313-1322.

Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane G.; Edminster, Carleton

B., comps. 2005. Connecting mountain islands and desert seas: Biodiversity and management of the Madrean Archipelago II; 2004 May 11-15;

Tucson,AZ. Proceedings RMRS-P-36. Fort Collins, CO: U.S. Department

of Agriculture, Forest Service, Rocky Mountain Research Station. 631 p.

Heidorn, P.B. 2004. Publishing virtual floras and faunas. Bulletin of the

American Society for Information and Technology. 30(2): 8-11.

Higgins, S.I.; Richardson, D.M.; Cowling, R.M.; Trindersmith, T.H. 1999.

Predicting the landscape-scale distribution of alien plants and their threat

to plant diversity. Conservation. Biology. 13: 303-313.

Jones, P.G.; Gladkov, A. 1999. FLORAMAP: A computer tool for predicting

the distribution of plants and other organisms in the wild. Cali, Colombia:

Centro Internacional de Agricultura Tropical.

Kelly, C.; Garison, J.; Aggarwal, A.; Sarkar, S. 2002. Place prioritization for

biodiversity reserve network design: A comparison of SITES and ResNet

software packages for coverage and efficiency. Diversity and Distributions. 8: 297-306.

Lobo,J.M.; Lumaret,J.P.; Jay-Robert, P. 1997. Taxonomic databases astools

in spatial biodiversity research. Annals of the Society of Entomology of

France. 33: 129-138.

Marshall, J. 1957. The birds of the pine oak woodland in southern Arizona

and adjacent Mexico. Pacific Coast. Avifauna No.32.

Marshall, R.M.; Turner D.; Gondor A.; [and others]. 2004. An ecological

analysis of conservation priorities in the Apache Highlands Ecoregion.

Prepared by The Nature Conservancy of Arizona, Instituto del Medio

Ambiente y el Desarrollo Sustentable del Estado de Sonora, agency and

institutional partners. 152 p.

Mexican National Institute of Statistic and Geography (INEGI). 2005. Uso

del Suelo y Vegetación Dataset. Scale: 1:250 000. Mexico City, Mexico.

Peterson, A.T.; Navarro-Siguenza , Adolfo G.; Hesiquio, B. 1998. The need

for continued scientific collecting; a geographic analysis of Mexican bird

specimens. Ibis. 140: 288-294.

Peterson, A.T. 2004. Predictability of the geography of species’invasions via

ecological niche modeling.The QuarterlyReview ofBiology. 78: 419-433.

Peterson, A.T.; Egbert, S.L.; Sanchez-Cordero, V.; Price, K.P. 2000. Geographic analysis of conservation priorities using distributional modelling

and complementarity: Endemic birds and mammals in Veracruz, Mexico.

Biological Conservation. 93: 85-94.

Peterson, A.T.; Soberon, J.; Sanchez-Cordero, V. 1999. Conservatism of

ecological niches in evolutionary time. Science. 285: 1265-1267.

Rebelo, A.; Siegfried, W.R. 1992. Where should nature reserves be located in

theCapeFloristicRegion,SouthAfrica?Modelsforthe spatial configuration

of a reserve network aimed at maximizing the protection of floral diversity.

Conservation Biology. 6: 243-252.

Schnase, John L.; Kama, Deborah L.; Tomlinson, Kay L.; [and others]. 1997.

The Flora of North America digital library: A case study in biodiversity

database publishing. Journal of Network and Computer Applications.

1997(20): 87–103.

Soberón,Jorge; Peterson,A.T. 2004. Biodiversity informatics: Managing and

applyingprimarybiodiversity data.PhilosophicalTransactions oftheRoyal

Society of London - Series B: Biological Sciences. 359(1444): 689-98.

Soberón, Jorge; Arriaga, Laura; Lara, Liliana. 2002. Issues of quality control

in large, mixed-origin entomological databases. In: Towards a global biological information infrastructure. European Environment Agency. 17-22.

Soberón, Jorge; Davila, M.P.; Golubov, J. 2004 Targeting sites for biological

collections.In: Seed storage:Turning science into practice (ed. H. Pritchard

& M. Way). Richmond: Kew Royal Botanic Gardens.

Soberón,Jorge;Llorente,JorgeB.; Oñate,Leonor. 2000.The use ofspecimenlabel databases for conservation purposes: An example using Mexican PapilionidandPieridbutterflies.BiodiversityandConservation.9:1441–1466.

U.S. Geological Survey (USGS). 2002. Shuttle Radar Topography Mission

(SRTM) Elevation Dataset [BIL raster images]. 90m spatial resolution.

Sioux Falls, SD.

Winker, K. 1996. The crumbling infrastructure of biodiversity: The avian

example. Conservation Biology. 10: 703-707.

Zhong, Y.; Luo, Y.; Pramanik, S.; Beaman, J.H. 1999. HICLAS: A taxonomic

database system for displaying and comparing biological classification and

phylogenetic trees. Bioinformatics. 15(2): 149-156.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
300 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 301

Restoration
302 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 303

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Agricultural Field Reclamation Utilizing Native Grass Crop 

Production

J. Curé

Polytechnic Campus, Arizona State University, Mesa, Arizona

Abstract—Developing a method of agricultural field reclamation to native grasses in the Lower San Pedro 

Watershed could prove to be a valuable tool for educational and practical purposes. Agricultural field 

reclamation utilizing native grass crop production will address water table depletion, soil degradation and 

the economic viability of the communities within the watershed. The focus of this study was to analyze the 

conversion of agricultural fields to native grass crops of three representative sites. Study sites include; a farm 

with fields that were seeded and allowed to grow for a period of time with subsequent termination of irrigation, a ranch that converted approximately 18 acres to native grasses for production of grass-fed beef and a 

ranch that is working towards a goal of restoring the giant sacaton habitat. Each site was analyzed utilizing a 

matrix of questions. Personal interviews and site visits were sources for information to complete the matrix. 

Conclusions were drawn through analysis of the matrix and recommendations developed to formulate the 

reclamation method most appropriate to achieve the anticipated goals of each site. This study developed a 

recommended method for agricultural field reclamation to native grasses in the Lower San Pedro Watershed 

to be used as a tool for educational and practical purposes utilizing the information obtained. The recommended method includes the processes for initial plan development, field preparation, and seeding. The 

methods are discussed within the project and prepared as an educational flier.

Introduction

	 Water table depletion, soil degradation and economic viability

are major issues in the borderland agricultural communities located

in Southern Arizona and Northern Mexico (Anderson and others

2006). Agricultural production of cotton and alfalfa, traditional

commercial crops, results in lower water tables and reduced flows

in related rivers with the ultimate impact on future sustainability of

the region (Anderson and others 2006). The Sonoran Desert, located

in Southern Arizona and Northern Mexico, has several river systems

and related watersheds in the United States and Mexico impacted by

water table depletion and the adverse effects of extensive high water

use agriculture (Collins and Bolin 2007).

	 Traditional agricultural crops require substantial amounts of water with the resultant water table drawdown. Agricultural practices

utilizing traditional commercial crops are not sustainable in the San

Pedro River Watershed as they exceed the recharge capacity of the

aquifer with the result of reduced flow in the river (Morehouse and

others 2008). The deterioration of the riparian area related to the river

is jeopardizing indigenous species of flora and fauna, such as the

Southwestern Willow Flycatcher (Empidonax traillii extimus) that

is on the endangered species list (Price and others 2005; U.S. Fish

and Wildlife Service 1995).

	 The goal of this study is to develop a plan and process to reclaim

agricultural fields to native grasses within the study area. Many attempts have been made at field reclamation with varying degrees of

success.Literature review,farmmanagerinterviews, and site analysis

were incorporated into the plan analysis and development.The goal of

the study isto provide a prescription forsuccessful field reclamation.

Background

	 The Lower San Pedro River has experienced decreased flow due to

irrigation, mining, stock grazing, public water supply, and domestic

and commercial/industrial wells. Drawdown increased from 10,000

acre-feet per year in the 1940s to more than 60,000 acre-feet per

year in 1990 (Katz and others 2009). The reclamation of agricultural

fields to native grasses will decrease water use, therefore, allowing

the restoration of the water table and increase flow in the river and

improvement in the related habitat(Baker n.d.).To encourage change

from traditional crops to native grasses, it is essential to document

the scientific aspects of the conversion. Literature review and site

inspections show that many small pieces of the bigger puzzle exist

but a comprehensive document that accurately describesthe detailsis

lacking.Acomprehensivedocumentthat establishestheprocedure for

change will enable stakeholders to make value judgment, educators

to have a tool for demonstration and activist supportive research.

	 The primary research question that this study addresses is how to

approach the reclamation of agricultural fields to native grasses and

ensure long term success. The objective of the study was to look

at three sites within the Lower San Pedro River watershed, review

supporting literature, and document the results. The development of

a reclamation process will provide a guide that allows for successful

reclamationof agriculturalfieldstonativegrasses.The encouragement

for stakeholders to convert to native grasses must be supported with
304 USDA Forest Service Proceedings RMRS-P-67. 2013

Curé Agricultural Field Reclamation Utilizing Native Grass Crop Production

evidence that it is an economically viable process for the present and

the future.

	 The initial fact finding started with a visit to the Natural Resources

Conservation Service (NRCS) office inTucson and an interview with

Ms. Kristen Egen, District Conservationist, in November of 2010.

Ms. Egen provided the standard procedure available from the NRCS

for the process of planting native grasses in the region. The process

includes research involving historic vegetation and soil analysis

utilizing the U.S. Department of Agriculture (USDA) NRCS Soil

Survey website. The preceding information allows determination of

the historical grasses in the area and, with that, a selection for the

development of the seed mix to be used. The NRCS website (http://

websoilsurvey.nrcs.usda.gov) is an important resource in the process

offieldconversionwithinformationaboutsoiltype andrecommended

grasses for the area under question.

Study Sites

	 Three representative sites within the San Pedro Watershed were

selected for the study: Black Farm Preserve (BFP) owned by the Salt

River Project (SRP), Camp Stool Ranch (CSR) owned by the Mercer

Family, and Cobra Ranch (CR) owned by The Nature Conservancy

(TNC). These sites were chosen on the basis of known qualities

including available data, access to the site, and available information and they represent a cross section of sites within the watershed.

Interviews and site visits along with written documentation of events

on the sites compose the majority of information documented for the

results analysis (table 1).

	 Black Farm Preserve (BFP) is owned by the Salt River Project and

managed by The Nature Conservancy. The fields planted to native

grasses totaled 101 acres. Ruth Valencia, Senior Environmental ScientistforSRP, provided information concerningBFPand themethods

used for conversion to native grasses and the subsequent results. SRP

worked with theTucson Plant Materials Centerstaff for development

ofthe seedmix to be utilized at theBFP.InterviewswithDanWolgast,

the farm manager at BFP, and Molly Hanson with TNC Lower San

PedroRiver Programprovided additional information concerning the

process of conversion at BFP and subsequent results.

	 Camp Stool Ranch (CSR), which is owned and operated by the

Mercer Family, is an operating cattle ranch. The field area planted

to native grasses totaled 18.7 acres. The information on this ranch

was obtained from interviews with Mike Mercer and Kristen Egen

along with documentation from the NRCS. Several site visits were

made to view the fieldsin the fall and spring. Dan Walgast and Molly

Hanson provided additional information concerning the grasses used

and history of the site through interview questioning at the site.

	 Cobra Ranch (CR) is a Nature Conservancy property and part of

the Aravaipa Canyon Preserve located at the headwaters of Aravaipa

Creek, part of the Lower San Pedro Watershed. The initial native

grass plantings encompassed 20 acres. Mark Haberstich, the preserve

manager, provided excellent historical documentation of the efforts

in restoring native grass to the major agricultural fields at the ranch.

Table 2 presents information on the scope of reclamation effort at

each site.

	 The key questions presented for each of the sites are listed in

table 3. The people interviewed were able to provide information to

answer the questions and are provided in the “Results” section. The

information that was provided is ofsignificant value with application

to this study, the results and recommendations for further research.

Results

	 Q: What were the existing conditions prior to change from agricultural fields to native grass species?

	 A:The existing conditionsforthe three study locations varied from

current agriculture crops,including cotton,Sudan grass, andwheat,to

fallow for several years. BFP fields in the previous years were under

a rotation of wheat and cotton. CSR had been growing Sudan grass

and winter rye the prior season. CR fields were fallow the preceding

5 years with Sudan grass as the crop in the year 2002.

Table 1—Restoration sites, lower San Pedro watershed.

Site-Agency Contact Address Phone and email

Black Farm Preserve Ruth Valencia Mail Station PAB352 (602) 236-2830

Senior Environmental PO Box 52025 Ruth.Valencia@srpnet.com

Scientist Phoenix, AZ 85072-

 2025

Camp Stool Ranch Mike and Lori Mercer Mike and Lori Mercer (5200 487-2021

Owners Box 157 mercer2021@msn.com

Mammoth, AZ 85618

Cobra Ranch Mark Haberstich 41099 W. Aravaipa (928) 828-3443

Arivaipa Canyon Preserve Manager Canyon Rd. Klondyke, mhaberstich@tnc.org

Preserve AZ 85643

The Nature Molly Hanson 1510 E. Fort Lowell (520) 357-6076

Conservancy Lower San Padro Rd., Tucson, AZ mhanson@tnc.org

River Program 85710

The Nature Dan Wolgast 80444 E. Pelayo Rd. (520) 237-6064

Conservancy Preserve Manager Winkleman, AZ 85192 dwolgast@tnc.org

The Nature Gita Bodner, PhD 1510 E. Fort Lowell (520) 622-3861

Conservancy Conservation Ecologist Rd., Tucson, AZ gbodner@tnc.org

 85710

NRCS USDA Kristen Egen 3241 N. Romero Rd. (520) 292-2999

District Conservationist Tucson, AZ 85705 Kristen.egen@az.usda.gov
USDA Forest Service Proceedings RMRS-P-67. 2013 305

Agricultural Field Reclamation Utilizing Native Grass Crop Production Curé

	 Q: What was the process used for preparing the fields?

	 A: All three sites used disc harrowing as part of the process used

for preparing the fieldsforseeding native grasses.In addition, the first

step in the process at BFP wasto plow under the fields upon purchase

of the property and then furrows were established for irrigation. CSR

discedthefieldareastobe seeded.Noothertreatment accompaniedthe

initialseeding. Later, wood chip mulch was added to try and improve

the organic material content of the soils. CR disc harrowed, leveled, and

dragged the fields. Seed drilling without disc harrowing was a followup method at CSR and seed drilling with imprinting was a follow-up

method at CR. The addition of soil amendments including manure at

CSR and CR, wood chips at CSR, and beneficial mycorrhizal fungi

at CR was a part of the preparation process for subsequent seed applications. The 2012 planting at CR will use a riphook and plow to

prep the field. The theory behind this method is that it will disturb

the gophers and allow the plants to grow enough to keep ahead of

the gopher population and subsequent destruction of the plant root

system.

	 Q: Was soil weed seed load addressed prior to seeding?

	 A: Soil seed load is one of the major problems inherent with the

reclamation project,substantiated by literature review, and field interviewswith the stakeholders.BFPpretreatedwith the use of herbicides

Table 2—Scope of site reclamation. 

BFP CSR CR

Date of initial reclamation 2004 2008 2007

Approximate acreage initially involved 33 18.5 20

Approximate total acreage available for reclamation 101 18.5 50

Table 3—Analysis matrix.

BFP CSR CR

1. What were the existing Rotation of wheat and Sudan grass and winter Sudan grass-2002-

conditions prior to cotton-prior season rye-prior season -fallow for 5 years prior to

change over from planting

agricultureal fields to

native grasses?

2. What was the process Disc harrowed, furrowed, Disc harrowed Disc harrowed leveled 

used for preparing the pre-irrigated with field cultivator and

fields? dragged

3. Was soil weed seed Yes, pretreated with No Yes, irrigated to stimulate

load addressed prior to herbicides germination and then

seeding? (pre or ost emergent mowed once

unknown)

4. What were the seeding Mechanical broadcast Drill seeder, hand Drill, mechanical

techniques? broadcast broadcast and hand

broadcast

5. Was plugging used for No Yes Yes

Sacaton?

6. What was the seed See detail listing See detail listing See detail listing

mix used?

7. How successful was Varied with seed type, Varied with seed type, Varried with seed type,

germination? soil conditions, time of soil conditions, time of soil conditions, time of

year applied and year applied and year applied and

available water available water available water

8. What was the follow- Irrigation provided until Consistent application to Sporadic to maintain

up for watering and field server and transfer maximize production cover and minimal

maintenance? complete growth

9. What is the current Weed populations with Abundant grasses that Native grass populations

condition of the fields? an abundance of native provide substantial increasing providing hay

grasses forage and 62-1 ton 

bales of hay/17.8 acres

10. What are the water Total server and transfer Aproximate 50% Undocumented use,

savings? reduction greater infiltration rate

11. Is graass-fed beef No Yes No

produced?
306 USDA Forest Service Proceedings RMRS-P-67. 2013

Curé Agricultural Field Reclamation Utilizing Native Grass Crop Production

for broadleaf weeds prior to seeding. It is unknown at the time of

writing the results if the pretreatment was a pre- or post-emergent

herbicide. CR plowed, irrigated, and mowed prior to initial seeding

of native grasses.The field atCSRhad been under cultivation without

an excessive weed seed load, and, therefore, the native grasses were

seeded without treatment for a weed seed load prior to seeding. CSR

andCR mowed to control weeds but did not use application of chemicals to control weed seeds. The three sites continue to use mowing

as a method of weed control. Timely mowing is critical for success.

The weeds must be allowed to mature to the point that flowers are

forming or have formed, but seed has not yet set.

	 Q: What were the seeding techniques utilized?

	 A: The seeding techniques varied among the sites. BFP used a

mechanical broadcaster initially but ran into problems with the seed

clogging the equipment. The range drill proved to be the most effective for the majority of the seeding process. CSR and CR used

a combination of drill seeder, mechanical broadcaster, and hand

broadcasting. The native seeds vary in shape and size leading to

difficulty with distribution mechanically by drill or broadcast. Close

attention to the calibration and distribution of the seed is essential for

good coverage and subsequent success of the crop. The range drill

proved to be the preferred method of application with ease of use and

successful distribution of seed for the larger seeds at all three sites.

When the practitioners were questioned on the use of hydroseeding,

they indicated that the process was too expensive to utilize although

labor and projectsuccess were notfactorsin the expense calculations;

only the initial rental cost of the equipment was considered.

	 Q: Was plugging used for Sacaton?

	 A: Plugging was not used at BFP but was used to establish Sacaton

with later plantings at the CSR and CR sites.

	 Q: What was the native seed mix used?

	 A: The native seed mix varied with each site. NRCS used soil and

historic vegetation analysisto help develop the seed list for each site.

The actual seed mix varied due to the historic vegetation, soils, seed

collection sources, available supply of seed from supplier, provence

of the seed source, and the individual budgetsforseed purchase. BFP

andCSRhave similarsoils(sandybottoms)whileCRsoilshavehigher

clay content. Granite Seed in Utah was the source for the BFP seed

supply. Wildlands Restoration in Tucson, Arizona, supplied the seed

for CSR and CR (refer to table 4 for listings of plant species).

	 Q: How successful was germination?

	 A: The results from the three sites indicate that germination success depends on a number of variables. The variables include, but

are not limited to, the preexisting field conditions, field preparation,

soil conditions, soil moisture content, natural rainfall, availability

of supplemental water, time of year of seeding, methods of seeding,

and provence of seed and seed quality. The climatic conditions were

similar for BFP and CSR as they are located west of the Galiuro

Mountain Range within the Lower San Pedro River Valley several

miles apart near Mammoth, Arizona. CR is in Aravaipa Canyon east

of the Galiuro Mountain Range and experiences different climatic

conditions. The average yearly rainfall for CSR and CR is 8 inches

(http://www.clrsearch.com/Mammoth_Demographics/AZ/WeatherTable 4—Native seed mix.

Black Farm Preserve (BFP)

Botanical Name Common Name Application Rate 

Hilaria jamesii Galleta 0.8# per acre

Setaria macrostachya Bristle Grass 1# per acre

Sporobolus airoides Alkali-Sacaton 1# per acre

Atriplex canescens Four Wing Salt-Bush 1# per acre

Aristida sps. Three-Awn .5# per acre

Boutelous curtipendula Sideoats Grama 6# per acre

Sporobolus crytandra Sand Drop Seed unknown

Camp Stool Ranch (CSR)

Botanical Name Common Name Application Rate

Trichachne californica Arizona Cottontop Mix @ 15# per acre

Eragrostis intermedia Plains Lovegrass

Setaria macrostachya Large-spike bristlegrass

Bouteloua barbata Sixweeks Grama

Bouteloua curtipendula Sideoats Grama

Bouteloua gracilis Blue Grama

Botriochloa barbinodis Cane Bluestem

Leptochloa dubia Green Sprangletop

Setaria macrostachya Bristle Grass

Sporobolus wrightii Wright’s Drop Seed

Cobra Ranch (CR)

Botanical Name Common Name Application Rate

Andropogon barbinoides Cane Beardgrass Mix @ 20# per acre

Bouteloua gracilis Blue Grama

Bouteloua rothrockii Rothrock Grama

Festuca microstachys Small Flowered Fescue

Leptochloa dubia Green Sprangletop

Plantago insularis Indian Wheat

Schizachyrium scoparium Little Bluestem

Setara macrotachys Plains Bristlegrass

Sporobolus crytandra Sand Drop Seed
USDA Forest Service Proceedings RMRS-P-67. 2013 307

Agricultural Field Reclamation Utilizing Native Grass Crop Production Curé

Forecast-Temperature-Precipitation?mammoth,az) while CR yearly

rainfall average is 15 inches (USDI BLM 2010). Rainfall affects the

ambientsoil temperature and the soil moisture content, both of which

influence seed germination. Although the three sites had variations

withgerminationsuccessitwouldappearthatCRhadinherentlyfewer

challenges due tomore favorable climatic conditions.Thefield conditions of BFP and CSR were similar in that they had been cultivated

within the past year even though the crops at each site were different.

CR had not been cultivated for a number of years and the soils were

compacted and contained a substantial weed seed load. The soil at

CR required tilling and irrigating for a season to reduce the weed

seed load and create a more favorable seed bed. The seed source for

BFP was Granite Seed in Utah. The seed supplied was harvested in

Utah and, therefore, did not have the same genetic characteristics as

the seed provided by Wildland Seed harvested in the Tucson area for

CSR and CR. The genetic variation will have an effect on the long

term success of the native grasses. This is evident with the necessity

for BFP to reseed in subsequent years as a result of poor germination

and seedling survival.

	 Q:Whatwasthe follow-up forwatering andmaintaining the fields?

	 A: BFP irrigated the fields for the first 3 years before sever and

transfer ofthe waterrightstook place.The purpose forthe conversion

of the agricultural fields to native grasses at BFP was to transfer the

water back to the river, the legal return of the water, and the water

rights back to the river. The weed problem is a continuing issue and

control with mowing on an as-needed basis is the method utilized.

CSR providesirrigation on a regular basis with an established schedule. A center pivot system that provides water to the fields is turned

on and off manually. Weed control is performed by mowing at the

appropriate times to prevent seed formation. CR provides irrigation

on an as-needed basis determined by soil moisture content and plant

health. The fields were flood irrigated initially but recent installation

of a wheel line system has allowed for changeover of some of the

fields. Weed control was performed by mowing at the appropriate

time and some hand weeding in select areas. The fields with native

grasses will not need to be completely re-seeded in 3 to 5 years as

do alfalfa fields or every year as with cotton.

	 Q: What is the current condition of the fields?

	 A: The BFP has weed populations with an abundance of native

grasses although many species of native grasses are declining. CSR

has weed populations with abundant grasses that provide substantial

forage and hay. The native grass populations are increasing and

competing with the weed population. Sixty-two 1-ton bales of native grass hay were harvested in the year 2010 from the 17.8 acres

as opposed to the 45 bales of Sudan sorgum from the same area in

previous years. The rancher indicates the cattle grazing on the native

grasses are healthier,weighmore atthe sale and produce better quality

meat.CRhassporadicweedpopulationswithnativegrasspopulations

increasing providing hay for livestock and erosion control within the

geographic area. Water infiltration of the soil has increased.

	 Q: What are the water savings?

	 A:Watersavings at BFPwere 100% as all the irrigation wells were

turned off and the water allowed to flow to the river. The water savings at CSR is estimated at 50% calculated from the amount of time

the pump runs compared to previous years with the high water-use

crops on the fields. CR water use is not well documented. The water

use on the fields to produce the previous crops was not recorded. A

piezometer wasinstalled in 2009 on the creek; prior to that no records

of water use or impact on the river were maintained.

	 Q: Is grass-fed beef produced?

	 A: BFP and CR do not produce grass-fed beef for market. CSR is

producing grass-fed beef and markets it at the local farmers market

and other related outlets. The current market price for grass-fed beef

is $6.50 per pound (Cunningham 2012) as opposed to the price for

grain finished or alfalfa-fed beef at $3.87 per pound (Kavilanz 2012).

Conclusions

	 Analysis of field conditions prior to the onset of reclamation is

critical to the approach taken for reclamation. Factors to be looked at

and included in the plan are the type ofsoil, historic vegetation, weed

seed load, existing or potential for irrigation, budget for the project

and goal for the crop to be produced. The plan for reclamation can be

formulated based on the previously mentioned factors and the goals

for the project. Consulting with the NRCS will assist in the development of a comprehensive plan.

	 Careful attention to soil preparation,seeding methods, and continued care are essential for crop success. Documentation from Cobra

Ranch indicates the areas with the highest success in establishing

native grasses (i.e., germination and health of seedlings) were the

sites in which weed seed load was reduced prior to planting, soils

were prepared with nutrients, and additions were made as necessary.

The method of seeding was based on the type of seed. Seeding occurredwhen appropriate seasonal parametersweremet,supplemental

irrigation was applied as needed, and weed control was performed as

needed. Provenance of seed appears to have contributed to success.

The seeded species appear to have established at higher frequency

in fields planted with local seed compared with seed originating in

Utah.

	 Camp StoolRanch demonstrated that use of native grassesreduced

water use (translated to energy use) and labor and the ongoing investment to maintain crops on the fields. The market for grass-fed beef

is strong and supports the premise for developing native grass crops.

The average retail price of ground beef is $3.87 (Kavilanz 2011),

while the market price for one pound of organic grass fed beef is

$6.50 (Cunningham 2012). CR has generated hay to use as mulch

for restoration purposes or as feed for cattle that are foraging in large

pastures where seed can be spread. The hay is valuable in more than

dollars and cents as it provides a seed source that contains the gene

poolindigenoustothe area, animportantfactorforrestorationprojects.

The neighbors are inquiring for hay for their livestock as well.

	 Analysis of the BFP fieldsindicatesthat it is possible to seed fields

and provide irrigation for a limited time with some success in crop

establishment. However, the long term success withoutsupplemental

irrigation is totally dependent on weather conditions and rainfall.

	 Each ofthe analyzed projects had different goals.The goalthatSRP

had for BFP wasto establish native grasses and sever and transfer the

water back to the river. The CSR goal is to establish a viable commercial agricultural crop. The crop in this case is grass-fed beef that

they are now able to provide with the forage and hay produced by the

native grass fields. The CR goal is to develop a seed bank, provide

forage crops, and initiate a soil improvement program while working

towards the ultimate goal of restoration of the giant sacaton habitat.

Each of the goals for the separate projects has been demonstrated as

being achievable. The key to success for the projects points to the

attention to detail and following the prescribed plan.

	 The naturalmonsoonal patterns play a key role in the establishment

of the grass crops. Additional irrigation is essential if the monsoon is

not an active season delivering adequate rainfall. Long-term water

needs for the crops will not be as great as the initial seeding and

establishment period. Water must be available on a regular basis

to have a successful germination and establishment of plants. Field
308 USDA Forest Service Proceedings RMRS-P-67. 2013

Curé Agricultural Field Reclamation Utilizing Native Grass Crop Production

reclamation to native grasses is a viable option for an agricultural

enterprise and holds multiple opportunities for the development of

a sustainable agricultural environment ensuring resiliency to the

changing climatic patterns through resource conservation. Resource

conservation was demonstrated by the three sites with the reduction

of water use and the reduction of extensive applications of fertilizers

high in nitrogen required by alfalfa and cotton crops.

Recommendations

	 Recommendations arepresentedbelowforfuture research, analysis,

and educational action.

	 1. Perform market-value studies for native grasses as forage, hay,

seed banks, and restoration projectsto substantiate the benefits of the

native grass crop.

	 2. Develop the economic information to substantiate native grasses

as a compelling agricultural crop.

	 3.Analyze the nutrient value of native grass hay compared to alfalfa

to be used as a marketing and educational tool.

	 4. Determine perton cost of native grass hay compared with alfalfa

(yield analysis) to assist with education about native grass farming.

	 5. Educate the public about the importance of utilizing native

grasses as a crop (figs. 1,2).

	 6. Extend a comparative analysis over a number of years by quantifying (1) labor for planting and care, (2) seed cost for the perennial

crop versus annual or short-term crops, (3) fuel costs for preparing,

planting, and field maintenance, and (4) nutrient and supplement

needs (particularly nitrogen/urea application) of native grass versus

other crops.

References

Anderson, G., Dick, E., Minor, J., and Prichard, A. 2006. Rethinking water

in the arid Southwest: The need for a new framework for managing water

in Arizona. Arizona’s Comprehensive Wildlife Conservation Strategy:

2005-2015. Arizona Game and Fish Department. 2 pp.

Baker, D. (n.d.) Bingham Cienega Natural Preserve The Nature Conservancy

Restoration Project. Saguaro Juniper Cooperation. Online: http://www.

saguaro-juniper.com/i_and_i/san_pedro/bingham_cienega.html. [March

10, 2012].

Bodner, Gita (personal communication, June 29, 2011, October 12, 2011).

Cunningham, J. 2012. Grass-fed beef. Coyote Creek Farm website: ttp://

coyotecreekfarm.org/beef/ [April 18, 2012].

Figure 1—Sample brochure.
USDA Forest Service Proceedings RMRS-P-67. 2013 309

Agricultural Field Reclamation Utilizing Native Grass Crop Production Curé

Figure 2—Sample restoration methodology.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Collins,T.,Bolin,B. 2007.Characterizing vulnerability to waterscarcity:The

caseof agroundwater-dependent,rapidlyurbanizingregion.Environmental

Hazards 7: 399–418.

Egen, Kristen (personal communication, November 23, 2010).

Haberstich,Mark(personalcommunication,December8,2011,January5,2012).

Hanson, Molly (personal communication, November 23, 2010, June 9, 2011,

June 29, 2011, October 12, 2011).

Katz, G., Haney J., Paradzick C., and Harris D. 2009. Mitigation, Restoration

and Endangered Species. In J.Stromberg, and B. Tellman (eds.). Ecology

and Conservation of the San Pedro River. Tucson, AZ: The University of

Arizona Press: 345-370.

Kavilanz, P. 2011. Beef prices soar. CNN Money website: http://money.

cnn.com/2011/03/31/markets/beef_price_increasing/index.htm. [April

18, 2012].

Mercer, Mike (personal communication, November 23, 2010).

Morehouse, B., Ferguson, D., Owen, G., [and others]. 2008. Science and

socio-ecologicalsustainability:ExamplesfromtheArizona-Sonora border.

Environmental Science and Policy 11(3): 272-84.

Price, J., Galbraith, C., Dixon, M., [and others]. 2005. Potential Impacts of

ClimateChange onEcologicalResources andBiodiversity in the San Pedro

Riparian National ConservationArea,Arizona.AReport to U.S. EPAfrom

the American Bird Conservancy. 543 p. Online: http://cfpub.epa.gov/si/

si_public_record_Report.cfm?dirEntryId=180083&CFID=15505330&C

FTOKEN=33891245&jsessionid=20308e19b20a47c456493el26563345

2703b [March 15, 2012].

U.S. Department of the Interior Bureau of Land Management (USDI BLM),

2010. Draft Aravaipa ecosystem management plan and environmental assessment. Online: http://www.blm.gov/pgdata/etc/medialib/blm/az/pdfs/

nepa/library/wilderness_plans/aemp.Par.32179.File.pdf/AravaipaEMP.

pdf. [April 18, 2012].

U.S. Fish and Wildlife Service. 1995. Endangered and threatened wildlife

and plants: Determining endangered status of the Southwestern Willow

Flycatcher.FederalRegister: status oftheSouthwesternWillowFlycatcher.

FederalRegister:10694-10715.Online:http://ecos.fws.gov/speciesProfile/

profile/speciesProfile.action?spcode=B094. [April 29, 2012].

Valencia, Ruth (personal communication, December 3, 2010, June 9, 2011).

Wolgast,Dan(personalcommunication,November23,2010,October12,2011).

Wong-Gonzalez,P.,Pineda,N.,Varady,R. 2008.Science and socio-ecological

resilience: Examplesfrom theArizona-SonoraBorder, Environmental Science & Policy 11(3): 272–284.
310 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Review of Black-Tailed Prairie Dog Reintroduction 

Strategies and Site Selection: Arizona Reintroduction 

Sarah L. Hale, and John L. Koprowski

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Holly Hicks

Arizona Game and Fish Department, Phoenix, Arizona

Abstract—The black-tailed prairie dog (Cynomys ludovicianus) was once widely distributed throughout the 

western United States; however, anthropogenic influences have reduced the species’ numbers to 2 percent 

of historical populations. Black-tailed prairie dogs are described as a keystone species in the grassland 

ecosystem, and provide many unique services, including burrows for other species (e.g. burrowing owls 

[Athene cunicularia] and rattlesnakes [Crotalus spp.]), nutrient rich soil that, in turn, provides rich vegetation 

for grazers, and food for many carnivores and birds of prey. Several efforts have been made to reestablish 

this species to its historical range. In southeastern Arizona, a recent reintroduction effort was built upon work 

of scientists that identified potential suitable areas with characteristics similar to those of existing prairie dog 

colonies in Mexico. Prairie dogs were first translocated to the sites in 2008, and individuals still remain on 

the landscape today. We compare this to other reestablishment efforts, and provide suggestions on ways to 

increase success of future reintroductions.

Introduction

	 Anthropogenic factorslike land conversion and habitat destruction

have become leading causes for decline in biodiversity worldwide

(Wilson 1988), and have created a need for conservation or restoration of many species. One commonly used method of restoration is

translocation. Translocation is the movement of living organisms

from one area with free release in another (IUCN 1987), and can be

used to establish, reestablish, or augment a population (Griffith and

others 1989). A translocation is considered successful when the action resultsin a self-sustaining population (Griffith and others 1989).

There are three classes of translocation: (1) Introduction: intentional

or accidental movement of an organism outside its native range; (2)

Re-introduction: the intentionalmovement of an organisminto native

range from which it has been extirpated by human activity or natural

catastrophes; and (3) Re-stocking: movement of numbers of plants

or animals of a species with the intention of building up the number

of individuals of that species in an original habitat (IUCN 1987). For

the purpose of this discussion, translocation will refer to the second

category, re-introduction.

	 The black-tailed prairie dog (Cynomys ludovicianus; BTPD) is

a burrowing rodent and is described as a keystone species in grassland ecosystems (Kotliar and others 1999, 2006; Miller and others

1994).This status suggests it provides a unique, significant service,

disproportionately to its abundance (Hoogland 2006), that no other

species can provide. BTPDs provide burrows for other species (e.g.

burrowingowls[Athene cunicularia] andrattlesnakes[Crotalus spp.]),

excavate nutrient rich soil that produces rich vegetation for grazers

(e.g. pronghorn [Antilocapra americana] and bison [Bison bison]),

and serve as food for many terrestrial carnivores and birds of prey

(Whicker and Detling 1988, Kotliar and others 1999, Underwood and

VanPelt2000).BTPDsalsomaintaingrasslandecosystemsbypreventing woody encroachment, contributing to landscape heterogeneity,

and creating fire breaksthrough the short vegetation on their colonies

(Kotliar and others 1999, Underwood and Van Pelt 2000). The loss

of a keystone species can have profound effects on an ecosystem,

including loss of biodiversity and community integrity (Kotliar and

others 1999). For example, at least nine species rely on prairie dogs,

and some populations decline as prairie dogs are eradicated (Kotliar

and others 1999). Otherspecies of prairie dogsincluding Gunnison’s

(C. gunnisoni: GUPD), white-tailed (C. leucurus: WTPD), Utah (C. 

parvidens: UTPD), and Mexican (C. mexicanus: MXPD) also may

serve similar roles (Davidson and Lightfoot 2007, Miller and others

1994, Miller and others 2000).

	 The BTPD was once widely distributed in NorthAmerica, but was

widely viewed as a pest, and control programs put into place over the

past century have reduced their numbers to 2 percent of historical

populations(Whicker and Detling 1988, Miller and others 1994). By

1960, the species was extirpated from Arizona (Underwood and Van

Pelt 2000), making Arizona the only state within their former range

to completely eliminate the BTPD. In 1972 an effort to reintroduce

the BTPD was made near Elgin, Arizona. It was unsuccessful due to

disagreementbetweenpartiesinvolvedaboutrelease sites andmethods

(Brownandothers1974),andprairiedogsbeingreleasedonthelandscape

without any site preparation (Brown, personal communication).

	 In2003, afterpetitionstolisttheBTPDandWTPDweredetermined

unwarranted by the U.S. Fish and Wildlife Service, a multi-state
USDA Forest Service Proceedings RMRS-P-67. 2013 311

Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: . . . Hale and others

plan was proposed to monitor and manage prairie dogs across their

ranges (McDonald and others 2011). In 2008, to improve grassland

health across the BTPD’s historical range, Arizona Game and Fish

Department (AGFD) implemented a plan to translocate BTPDs to

Las Cienegas National Conservation Area (hereafter, Las Cienegas;

fig. 1), in southeastern Arizona, which is located within the Madrean

Archipelago. We will review the methods used for site selection and

translocation of BTPDs to Las Cienegas, compare these methods to

those used in othertranslocations, and provide suggestionsto increase

the likelihood of successful future translocations.

Site Selection

	 A 2005 study, conducted by the University of Arizona, compared

characteristics of currently occupied BTPD colonies in Mexico with

characteristicsofpotentialtranslocationsites atLasCienegasinsoutheastern Arizona. Researchers evaluated grass cover, forb cover, bare

ground, visual obstruction, shrub density, tree density, gravel, sand,

silt/clay, and grassspeciesrichness(Coates 2005). Las Cienegas was

deemed a suitable site for translocation, and in 2008 the first colony

was prepared by AGFD (see colony preparation below).

	 Other translocation efforts have only evaluated slope, type of soil,

and type of vegetation on potential translocation sites(Long 2006) as

well asvisible andhistorical evidenceofformerprairiedogoccupancy

(Long 2006, Truett and others 2001). In areas being considered as

translocation sites, woody plants, shrubs, and tall grasses may have

encroached (Truett and others 2001) and modification to the environment must take place to prepare sites for translocation of BTPDs.

Colony Preparation

	 The bestsitesfortranslocationare thosewith vacant,intact burrows

(Long and others 2006); however, many translocations do not have

access to intact burrow systems and must prepare overgrown sites

for translocation. Methods of preparation include prescribed burns,

mechanicalshrub/woodyplantremoval,mowing,grazing, application

of herbicides, seeding, and installation of artificial burrows (Truett

and others 2001, U.S. Fish and Wildlife Service 2009). 

	 Between 2008 and 2011, AGFD removed mesquite, mowed grass

to <30 cm in height (the maximum height of vegetation preferred by

BTPDs; Hoogland 1995), and installed 25 artificial burrows on each

of four sites, 4 ha each, for translocation at Las Cienegas (fig. 2). A

backhoe was used to install each artificial burrow that consists of one

underground chamber, located 130-180 cm below ground accessible

through a length of 10 cm diameter flexible tubing. Each burrow entrance was covered with an acclimation cage to deter initial dispersal

(fig. 3), and four 20-25 cm deep starter burrows, one meter from the

entrance of each artificial burrow, were dug around each artificial

burrow entrance with an auger. Each colony site was located <5.6

km from another, within the dispersal range of BTPDs (up to 6 km;

Hoogland 2006).

Methods of Capture and Group Composition

Arizona Game and Fish Department pre-baited 15 x 15 x 60 cm,

double-door live traps (Tomahawk Live Trap Co., Wisconsin) and

observed individuals at source populations, in New Mexico and

Sonora, for 10 days prior to actual live-trapping. Observations serve

to determine if the population is healthy (i.e., does not have plague),

and to identify individual coteries, or family groups. After the initial

10 days,AGFD trappedBTPDs(ofthe arizonensissubspecies, which

was historically found in Arizona), dusted for fleas, and translocated

prairie dogs mostly together in observed coteries to increase success

and reduce dispersal (Shier 2006a,b).Acoterie istypically composed

of one adult breeding male, three to four related breeding females,

juveniles from the previous year, and yearlings (Hoogland 1995).

AGFDtranslocatedaminimumof60prairiedogstothreenewcolonies

at Las Cienegas, and will translocate individuals to the fourth colony

in October 2012.

	 In other translocation efforts with recipient sites in New Mexico

andSouthDakota,mixedfamily groups(randomlymixedprairie dogs

fromthe same colony) had similarsurvivalto same family groups, and

workersdiscontinuedtransportinganimalsinfamilygroupsbecause“it

was easier and more economical” (Long and others 2006). However,

a study of the same prairie dog colony in New Mexico indicated that

individualstranslocated in family groups had higherreproduction and

were five times more likely to survive post translocation than prairie

dogs that were not translocated in family groups (Shier 2006a). The

discrepancies in these two studies can be explained by the methods

of evaluating survivorship, and the size oftranslocated family groups.

Long and others(2006) estimated survivorship through visual counts

of prairie dogs 2 months after translocation, whereas Shier estimated

survivorship by live trapping all individuals and offspring 1 year after

translocation (Shier 2006b). Also, same-family groups translocated

to South Dakota were only partial family groups (average of five

individuals per group) whereas “same-family” groupstranslocated to

New Mexico were complete family groups(average of 11.3 individuals per group; Shier 2006b). This suggests that, wherever possible,

family groups should be determined prior to capture and transport,

and prairie dogs should be translocated as family groups to increase

the chance of success.

Figure 1—Approximate location of Las Cienegas National Conservation Area, Arizona, indicated by star.
312 USDA Forest Service Proceedings RMRS-P-67. 2013

Hale and others Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: . . . 

Figure 2—Colony locations at Las Cienegas National Conservation Area (Cieneguita, Road Canyon, Mud Springs) 

have received black-tailed prairie dogs (Cynomys ludovicianus). Gardner Canyon has been prepared but has not 

yet received prairie dogs. 

Figure 3—Artificial burrow consisting of an underground half-cylinder nest chamber connected to surface 

by 10-cm diameter flexible plastic tubing. Acclimation cage is placed above the burrow entrance to dampen 

dispersal and provide refuge from predators. Modified from Long et al. 2006.
USDA Forest Service Proceedings RMRS-P-67. 2013 313

Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: . . . Hale and others

Acclimation Cages

Hard release is the process of capturing, transporting, and releasing animals in a new location without any acclimation period (Clark

and others 2002, Franzreb 2004). Soft release is the same process of

animal capture and transport to a new location, but allows animals to

acclimate to their new location for a period of days or weeks before

release (Franzreb 2004).

	 Using the soft release method, AGFD installed wire acclimation

cages over burrow entrances to dampen dispersal and provide refuge

from predators (fig. 3). Prairie dogs were provided with food (Purina

herbivore chow, PhoenixZoo), carrots, andwater ad libitumwhile the

acclimation cageswere in place torestore energylostduringtransport,

burrow excavation, and stressfrom being in a new environment(U.S.

Fish andWildlife Service 2009).Acclimation cages were left in place

for 2 weeks; however, almost all prairie dogs excavated a tunnel out

of the cages before their removal (Sarah Hale, personal observation

of black-tailed prairie dogs at Las Cienegas, October 2011).

	 Prairie dogs not kept in acclimation cages have a higher probability

of dispersal. During dispersal, prairie dogs are more vulnerable to

predators, and therefore have high mortality rates (Hoogland 1995,

2006).Onegroupofprairiedogshad100percentdispersalfromstarter

burrows when acclimation cages were not used (Truett and others

2001). Another study found that, after acclimation cages were used

for 5 to 15 days, most animals continued to use starter burrows for

up to 1 year during which time they excavated new burrows nearby

(Truett and others 2001). These findings suggest that the use of acclimation cages increases the success of translocations by reducing

dispersal immediately following translocation.

Environmental Stochasticity 

Las Cienegas National Conservation Area is located at the southwestern periphery of the BTPD’s former range (fig. 4). This arid

system is much harsher than more northerly locations, and may cause

prairie dogs translocated to these systems to have poor survival and

reproduction compared to prairie dogsinmore temperate parts oftheir

range (Facka and others 2010). Between May and June 2011, one

BTPD population at Las Cienegas decreased by 54 percent, and only

produced one juvenile that subsequently died, but at least 9 females

showed signs of parturition (Hale, unpublished data). We speculate

that this was due to lower than average rainfall that created a lack

of available food resources. Furthermore, we speculate that lack of

resources caused increased infanticide, and the need to forage farther

awayfromthe safetyofburrows,making prairiedogsmorevulnerable

to predation; we documented three fatalities caused by predators on

colony peripheries where tall vegetation obscured nearby predators

(twocoyotes[Canis latrans];onered-tailedhawk[Buteo jamaicensis]).

Figure 4—Historical range of the black-tailed prairie dog (Cynomys ludovicianus: modified from Hall 1981). Approximate locations of source populations in New Mexico and Mexico are black circles. Approximate location of Las Cienegas National Conservation Area in Arizona is black square.
314 USDA Forest Service Proceedings RMRS-P-67. 2013

Hale and others Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: . . . 

Our findings are consistent with another study that observed prairie

dogs foraging on colony peripheries when resources were scarce,

and found they were more vulnerable to predation (Koford 1958)

than those foraging in colony centers. Drought lead to the collapse

and local extirpation of several BTPD populations translocated to

the Chihuahuan desert (Facka and others 2010). This suggests that

drought years can greatly hinder translocations of BTPDs to their

former range in arid regions.

	 After the precipitous population decline at one of the Las Cienegas

colonies,AGFDbegansupplementalfeedingstartinginJune2011.The

same feed provided to individualsin acclimation cages was placed at

or near burrow entrances in an attempt to keep prairie dogs closer to

burrows. No additional predation events were observed after supplemental feeding began, and only two more individuals disappeared

from the colony between June and October while the supplemental

feed was available (Hale, unpublished data). In 2012, supplemental

feeding began in March to increase survival and reproduction, and at

least70juvenileswereobservedatthe same colonythatproducedonly

one the previous year (Holly Hicks, Sarah Hale, personal observation

of black-tailed prairie dogs at Las Cienegas, April 2012). With the

apparent massive improvement to recruitment,supplemental feeding

will likely continue to be provided from March-July during dry years

until the colony appears to be resilient in drought periods.

Population Augmentation

Atranslocationis consideredsuccessfulwhenapopulationbecomes

self-sustaining (Griffithandothers1989).The threeoccupiedcolonies

atLasCienegas have not yet become self-sustaining (table 1), but, due

tothedrought andtheuniquenatureofthistranslocationeffort,itistoo

earlytodrawa conclusion.Because the threeLasCienegaspopulations

are not yet self-sustaining, populations have been augmented yearly,

usually in September or October, with new individuals, to increase

numbers and genetic diversity, from the same source populations

from which the new colonies are started (MacDonald Ranch, NM,

Ladder Ranch, NM, Sonora, Mexico). Augmentation has also been

recommended for populations of UTPDs when a significant decrease

in the spring count is observed (U.S. Fish and Wildlife Service 2009)

and was necessary for sustaining translocated populations of Gunnison’s prairie dogs in Sevellita National Wildlife Refuge in New

Mexico (Friggens and others 2009). Augmentation is not used in all

translocations, but may prove to be a valuable tool in arid climates

until populations can withstand drought and predation.

Conclusions

	 Arizona Game and Fish Department has returned a native species

to the state after a 50-year absence. With the help of the University

of Arizona, they were able to evaluate potential translocation sites

in more detail than other translocation efforts. Reliable methods of

site preparation and artificial burrow installation have been used;

methods of capture, group composition and acclimation, as well as

continued augmentation have all increased the likelihood of success

of populations translocated to Las Cienegas.

	 The aridclimateofLasCienegaswill continue toprovide challenges

in reestablishing the BTPD. Drought and predation have led to lower

survival and reproduction than expected, but supplemental feeding

at Las Cienegas seems to keep prairie dogs closer to their burrows,

reducesriskofpredation, andincreasesrecruitmentgreatly.Continued

supplementalfeeding during drought periods will be crucial to permit

the BTPD to become self-sustaining once again in the state. Future

translocationsinArizona, and other arid regions,should consider the

effects of drought, and provide supplemental food before population

declines occurin these sensitive populations.In the long term,seeding

sites and management for desired grass composition may be a tool

to replace supplemental feeding. Augmentation may also be a useful

method for increasing success of translocations, particularly in arid

climates.

Acknowledgments

	 We thank Bill Van Pelt and Tim Snow for their early efforts in

bringing the black-tailed prairie dog back to Arizona.

References

Brown, David E.; Todd, Richard L.; and Levy, Seymour H. 1974. Proposal

for the reintroduction of the black-tailed prairie dog into Arizona, Project

W-53-R-24. Arizona Small Game Investigations, Arizona Game and Fish

Department.

Clark, Joseph D.; Huber, Djuro; and Servheen, Christopher. 2002. Bear reintroductions: Lessons and challenges. Ursus 13: 335-345.

Coates,CarolA.2005.Suitabilityofpotentialhabitatforthe extirpatedArizona

black-tailed prairie dog. M.S.Thesis, University ofArizona,Tucson. 55 pp.

Davidson, Ana D.; and Lightfoot, David C. 2007. Interactive effects of keystone rodents on the structure of desert grassland arthropod communities.

Ecography 30(4): 515-525.

Table 1—Population abundances for black-tailed prairie dogs (Cynomys ludovicianus) at each colony on Las Cienegas between 2008 

and 2012. X indicates colony not yet in existence. Post-Augmentation indicates population size after new individuals were added to 

an existing colony. Population abundances prior to October 2011 are based on visual estimates, whereas after September/October 

2011 all individuals were trapped and counted. 

Oct 2009 Oct 2010 Sept/Oct 2011

Oct Pre- Post- Pre- Post- Pre- Post- April

2008 augmentation augmentation augmentation augmentation augmentation augmentation 2012

Road Canyon 74a 18 57 18 43 10 40 ≥23b

Mud Springs X 68a n/a 17 46 12 42 ≥25b

Cieneguita X X X 64a n/a 20 40 99

a

Colony received prairie dogs for the first time at this date.

b

Spring count not completed at this date for these colonies.
USDA Forest Service Proceedings RMRS-P-67. 2013 315

Review of Black-Tailed Prairie Dog Reintroduction Strategies and Site Selection: . . . Hale and others

Dullum, Jo Ann L. D.; Foresman, Kerry R.; and Matchett, Marc R. 2005.

Efficacy of translocations for restoring populations of black-tailed prairie

dogs. Wildlife Society Bulletin 33(3): 842-850.

Facka, Aaron N.; Roemer, Gary W.; Mathis, Verity L.; Kam, Michael; and

Geffen, Eli. 2010. Drought leads to collapse of black-tailed prairie dog

populations reintroduced to the Chihuahuan desert. Journal of Wildlife

Management 74(8): 1752-1762.

Franzreb, Kathleen E. 2004. The effect of using a “soft” release on translocation success of red-cockaded woodpeckers. In: R. Costa and S. J. Daniels,

Red-cockadedwoodpecker:Roadtorecovery.Washington:HancockHouse

Publishers: 301-306.

Friggens, Michael T.; Davidson, Ana D.; Collins, Scott L; Lightfoot, David

C.; and Koontz,Terri L. 2009.Reintroduction of Gunnison’s prairie dogsto

the Sevilleta National Wildlife Refuge: Lessons and challenges [abstract].

In: 94th ESAAnnual Meeting; 2009 Aug 2-7; Albuquerque, NM.

Griffith, Brad; Scott, J. Michael; Carpenter, James W.; and Reed, Christine.

1989. Translocation as a species conservation tool: Status and Strategy.

Science 245: 477-480.

Hale,SarahL.2011.Effectsofre-establishing black-tailed prairiedogcolonies

on southeasternArizona grasslands. Unpublished data on file at: University

of Arizona, Tucson, AZ.

Hall, Eugene R. 1981. The mammals of North America. 2nd ed, Vol 1. New

York: John Wiley & Sons, Inc. 600 pp.

Hoogland,John L. 1995. The black-tailed prairie dog: Social life of a burrowing mammal. Princeton: Princeton University Press. 557 pp.

Hoogland,John L. 2006. Conservation of the black-tailed prairie dog: Saving

NorthAmerica’s western grasslands.Washington DC:Island Press. 350 pp.

IUCN.1987.Positionstatementontranslocationsoflivingorganisms:Introductions,reintroductions, andre-stocking.InternationalUnionforConservation

of Nature and Natural Resources Council, Gland, Switzerland.

Koford, Carl B. 1958. Prairie dogs, whitefaces, and blue grama. Wildlife

Monographs, No. 3.

Kotliar, Natasha B.; Baker, Bruce W.; and Whicker, April D. 1999. A Critical

review of assumptions about the prairie dog as a keystone species. Environmental Management 24(2): 177-192.

Kotliar, Natasha B.; Miller, Brian J.; Reading, Richard P.; and Clark, Timothy

W. 2006. The prairie dog as a keystone species. In: Hoogland, J. L. Conservation of the black-tailed prairie dog: Saving North America’s western

grasslands. Washington DC: Island Press: 53-64.

Long, Dustin; Bly-Honness, Kristy; Truett,Joe C.; and Seery, David B. 2006.

Establishment of new prairie dog colonies by translocation. In: Hoogland,

J. L. Conservation of the black-tailed prairie dog: Saving North America’s

western grasslands. Washington DC: Island Press: 53-64.

McDonald, Lyman. L.; Stanley, Thomas R.; Otis, David L.; Biggins, Dean

E.; Stevens, Patricia D.; Koprowski, John L.; and Ballard, Warren. 2011.

Recommended methods for range-wide monitoring of prairie dogs in the

United States. U.S. Geological Survey Scientific Investigations Report

2011–5063. 36 pp.

Miller,Brian;Ceballos, Gerardo; andReading,Richard. 1994.The prairie dog

and biotic diversity. Conservation Biology 8(3): 677-687.

Miller,Brian;Reading,Richard;Hoogland,John;Clark,T.;Ceballos,Gerardo;

List, R.; Forrest, S.; Hanebury, L.; Manzano, P.; Pachecho, J; and Uresk,

D. 2000. The role of prairie dogs as a keystone species: Response to Stapp.

Conservation Biology 14(1): 318-321.

Shier, Debra M. 2006a. Effect of family support on the success of tranlsocated

black-tailed prairie dogs. Conservation Biology 20: 1780-1790.

Shier, Debra M. 2006b. Translocations are more successful when prairie dogs

are moved asfamilies. In: Hoogland,J. L. Conservation of the black-tailed

prairie dog: Saving North America’s western grasslands. Washington DC:

Island Press: 53-64.

Truett,Joe C.; Dullum,JoAnn L. D.; Matchett, Marc C.; Owens, Edward; and

Seery, David. 2001. Translocating prairie dogs: A review. Wildlife Society

Bulletin 29(3): 863-872.

Underwood, Jared G.; William E. Van Pelt. 2008. A proposal to reestablish

the black-tailed prairie dog (Cynomys ludovicianus) to southern Arizona.

NongameandEndangeredWildlifeProgramDraftTechnicalReport.Arizona

Game and Fish Department, Phoenix, Arizona.

U.S. Fish and Wildlife Service. 2009. Utah Prairie Dog (Cynomys parvidens)

Draft Revised Recovery Plan. U.S. Fish and Wildlife Service, Denver,

CO. 122 pp.

Whicker, April D.; and Detling, James K. 1988. Ecological consequences of

prairie dog disturbances. BioScience 38(11):778-785.

Wilson, Edward O. 1988. Biodiversity. Washington, DC: National Academy

Press. 521 pp.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
316 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Trajectory and Rate of Desert Vegetation Response 

Following Cattle Removal

Robert L. Minckley

Department of Biology, University of Rochester, Rochester, New York

Abstract—Cattle have grazed continuously over the past three centuries in the Sky Island region and most 

work has focused on how these grazers have affected riparian and grassland habitats. I examined the effects 

of grazing on a fuller spectrum of desert habitats that occur in the close proximity to the San Bernardino Valley of Mexico and the United States. Plots in each of five habitats were placed in two adjoining areas: where 

grazing had not occurred for 25+ years and where grazing had only ceased 4+ years before this study. Vegetation responded quickly to removal of cattle in mesic environments and formed a species-poor, structurally 

diverse community. Grazing in more xeric communities did not affect vegetation cover but increased plant 

species richness. The differences in grazed and ungrazed areas of xeric habitats may indicate these areas were 

little affected by grazing or that the response after grazing ends is slower than captured by the time course 

represented here. Exotic species were essentially limited to recently grazed areas of the riparian habitat. 

Introduction

	 Vegetation changed dramatically during the late 1800s over large

areas of the warm North American deserts, including the “Sky Islands” (sensuWarshall 1994), after a prolonged drought, coupledwith

large-scale cattle grazing period, followed by a period of unusually

high precipitation (Bryan 1925; Hastings and Turner 1965). Rivers

and streams that had coursed through uncut channels (1) incised the

landscape lowering water levels and (2) changed rivers and streams

from sources of perennial surface water to dry channels with flow

present only after sufficient rains (Bryan 1928; Cooke and Reeves

1975). Valleys that had been grasslands were replaced by shrubs and

subtrees (Bahre 1991; Bull 1999; Hastings 1959; Humphrey 1987).

The contribution of cattle grazing to the change in vegetation over

this period has been debated (Hastings and Turner 1965). Grazing

had been widespread and stocking rates often high over much of the

region since Spanish explorers first arrived in the 1600s (Wagoner

1952, 1960). However, the scant written description of this period

leaves it difficult to interpret what early effect the large cattle herds

had on vegetation.Alack of information on the early grazing impacts

coupled with observationsthat dramatic vegetation changes occurred

in the late-1800sin places cattle had never been suggeststhere islittle

support that grazing alone was the cause of abrupt vegetation change

in the late-1800s. Some combination of climatic and human-caused

factors likely coincided to bring about this change in vegetation

(Hasting and Turner 1965). Empirical results from the present-day

will provide further insights into this discussion.

	 Because of their importance to ranching and agriculture, the riparian and grassland habitats have received most attention regarding the

effects of cattle grazing on vegetation communities (Stromberg and

Tellman 2009, and referencestherein). However, these habitats cover

a minority of the area in the present-day sky island region. Fewer

studies have addressed how grazing affects desert scrub and other

more xeric habitats that cover the largest expanse of NorthAmerican

warm deserts.

	 This study examined differences and rates of change of plant species richness, cover and composition after cattle grazing ended in

five desert plant communities of the Chihuahuan desertscrub biome

(Brown 1994). In addition to habitat, grazing histories also differed

on adjoining areas; in one area cattle had been removed in 1979 after

it had been purchased as habitat for endangered species (the presentday San Bernardino National Wildlife Refuge, Department of the

Interior), and the other area had been ranched until 2000 when cattle

were removed and agriculture was stopped (the present day Rancho

San Bernardino, Fundación Cuenca de los Ojos). The difference in

history of land use allowed me to compare floras at sites that had

experienced recent cattle activity to floras at nearby sites that had no

cattle present for 25+ years. Plant sampling sites were in five highly

interdigitated habitat types in a small area (approximately 16 km2).

Close proximity of sites meant the same plant species could have

potentially occurred in each habitat and differences in plant species

richness and abundance due to rainfall, temperature, and other climatological conditions were minimized. Sampling designssuch asthese

has been called “space for time comparisons” and are informative

not only about how communities respond to disturbance but also the

rate and extent of succession after disturbance (Fukami and Wardle

2009). Specifically,I asked the following three questions:(1) do plant

species richness and cover increase in habitats with greater water

availability and is this pattern the same where recent grazing has

occurred as where it has not? (2) does the rate of vegetation change

correspond to water availability? and (3) how does water availability

influence distribution and species richness of non-native species?
USDA Forest Service Proceedings RMRS-P-67. 2013 317

Trajectory and Rate of Desert Vegetation Response Following Cattle Removal R. Minckley

Methods

	 The study was done in the San Bernardino Valley, which runs

north-south across the Mexico-United States border in northeastern

Sonora, Mexico, and southeastern Arizona, USA. Elevation is approximately 1070 m and climate is xeric temperate with an annual

average precipitation of 360 mm. Description of the climate and area

are detailed in Minckley (2008).

	 In fall 2003, eight sites were established in each of five habitat

types; desert scrub, mesquite scrub along minor (upland) drainages,

mesquite forest along major drainages, grassland, and riparian. The

same sites were sampled again in spring 2004. These habitats are not

discrete but do reflect a soil moisture gradient from highly saturated

in the riparian zone along a permanentstream to more xeric in upland

vegetation associations(creosote bush scrub,mesquite grassland, and

mesquite forest) where cacti and desert shrubs occur such as Acacia 

constricta (whitethorn acacia), Larrea tridentata (creosote bush), and

Flourensia cernua (tarbush). Grasses were common, but not dominant, in most of the grassland and were absent from the other upland

vegetation types. Mesquite (Prosopis juliflora) occurred in both the

grassland and mesquite forest, and creosote bush was the dominant

perennial in the creosote bush scrub. For brevity, hereafter I refer to

these vegetation types as follows; desert scrub, grassland, and riparian as written but mesquite scrub along minor (upland) drainages as

bosque dry, and mesquite forest along major drainages as bosque wet.

	 In each habitat, half (4 of 8) of the sampled sites were located on

ranchlandsinMexico that had been taken out of grazing 3 years before

this study began. Cattle on this ranch had been stocked year-round

when the cattle operation was active, and in the 2 years before cattle

were removed (1999-2000), stocking density was approximately

twice as high as recommended for long-term forage sustainability (J.

Austin, pers. comm.). This area is referred to hereafter as the grazed

area. The othersitesin the United States had not experienced grazing

since 1979 and are referred to hereafter as in the ungrazed area.

	 Vegetation was sampled using a multi-scaled sampling method

developed and described by Stohlgren and colleagues (Barnett and

Stohlgren 2003; Stohlgren et al. 1998). These plots are 20 m x 50 mand

subdivided into two non-overlapping 10 m2 and one 100 m2 subplots.

Thissize and rectangularshape of the plots provides a better estimate

ofvegetationheterogeneityinlandscapesthansmallerDaubenmireand

Parker transects that have been traditionally used by field ecologists

(Muller-Dombois and Ellenberg 1974). Stohlgren et al. (1998) found

that approximately 45-80% more plant species and a greater proportion of rare, habitat specialist species are sampled in these plots than

in the smaller plots. Where plots were located was determined using

random numbers. Once the plot origin was fixed, the long axis of the

plot was oriented to maximize number of species a plot included.

	 In each plot, the species composition, species frequency (= abundance) and cover of plants was measured. Plots were measured in all

five vegetation typesin the grazed and ungrazed areasfor a total of 40

plots (4 plots x 5 habitats x 2 grazing histories). Each plant species

was categorized as native or exotic based on information available

through the United States Department of Agriculture (plants.usda.

gov, accessed on 15April 2012) and a specieslist for Sonora, Mexico

(T. Van Devender, unpubl. data).

	 I first tested with a t-test if the plant species richness recorded in

plots differed among the spring and fall samples. Secondly, I tested

with a 2-way ANOVA for the effect of grazing on species richness

by treating habitat and grazing history as main effects with species

density as the variable. Third, I tested with a 2-way ANOVA for the

effect of grazing on percentage cover by plants by treating habitat and

grazing history as main effects with percentage cover asthe variable.

To gauge the level of heterogeneity among plots, habitats and grazing

histories, I compared all habitat and grazing history combinations

using Jaccard’s similarity index.

Results

	 There were 147 plant taxa recorded from the plots, of which 133

were identified to species and 14 were considered uncertain identifications. This latter group was not included in the species number

and cover estimates. There were more species in the spring than in

the fall when calculated as total species (spring = 105 species; fall =

89 species), or if calculated as average species richness per habitat

(fig. 1: spring = 18.7 + 1.06 s.e., fall = 12.4 + 1.06 s.e.; p < 0.0001).

Only the riparian habitat did not have more plantspeciesin the spring

than in the fall sample. The seasonal difference in species number

was particularly pronounced in the driest three habitats, desertscrub,

bosque dry, and grassland (fig. 1). In all of the analyses, we tested

both spring and fall data however, I limit the discussion to the spring

data because there were no differences in the results among seasons

and there were more species observed.

	 AcompletelistofthespecieswillbeavailableinthejournalChecklist

(www.checklist.org.br/) andmade available as aResearchSpecieslist

in Madrean Archipelago Biodiversity Assessment (MABA)/Southwest Environmental Information Network (SEINet) online database

(Madrean.org).

Species Richness and Cover

	 Grazed habitats had significantlymore plantspecies(mean = 21.75

+ 1.1 s.e.) than ungrazed habitats (mean = 15.65 + 1.1 s.e.; t-test, p

= 0.0005) and there were more species where grazing had occurred

recently than where cattle had not been present for approximately

23+ years, in all but the bosque dry habitat (fig. 2). Tukey’s test of all

pairwise comparisons showed there were significantly more species

in grazed areas of desert scrub, grassland, and riparian habitats than

in ungrazed areas of more mesic riparian and bosque wet habitats.

Figure 1—Seasonal differences in average number of plant species recorded from eight plots per habitat in five habitats. Each plot was sampled 

twice, once in spring 2004 and again in fall 2003. Significantly more 

species were recorded in the spring.
318 USDA Forest Service Proceedings RMRS-P-67. 2013

R. Minckley Trajectory and Rate of Desert Vegetation Response Following Cattle Removal

However, in most habitatsthere was no significant difference in plant

species richness when grazed and ungrazed areas were compared

except for the riparian habitat where there were significantly fewer

plant species in the ungrazed area than in the grazed area (fig. 2).

	 Mean percent cover did not change in response to grazing history.

Post-hoc Tukey’s test revealed only one comparison differed significantly; plots in ungrazed areas of grassland had less cover than in

plots of the ungrazed area of the riparian habitat (fig. 3). Cover was

not significantly different in any habitat when plots in grazed and

ungrazed areas were compared.

Species Similarity

	 Vegetation composition varied considerably among habitats, most

notably in the riparian habitat where cottonwood dominated the area

that was ungrazed, but had not become established downstream in

the 3 years after grazing had ended (figs. 4, 5). Approximately 30%

of the vegetation coverage in habitats was by four perennial plant

species (fig. 4), two that occurred in all habitats (mesquite and sacaton bunchgrass, Sacaton wrightii), one that occurred in all habitats

exceptriparian (creosote bush), and one limited to the riparian habitat

(cottonwood, Populus fremonti). Plant composition among plots was

very heterogeneous, as is best exemplified by the low mean species

similarities of the samples taken at different plots within the same

habitat and grazing history (mean = 37.9%, range from 0.28 bosque

wet grazed – 0.49 desert scrub ungrazed). 

Exotic Species

	 There were six introduced speciesfound in the plots; although only

the widespreadmustard, Eruca vescaria, occurred in desertscrub and

Figure 2—Mean plant species from plots in five habitats that had either 

experienced grazing ca. 5 years before this study or had not been grazed 

for 25+ years before the study. More plant species occurred in most 

habitats in the areas where cattle had been grazing recently. Letters above 

columns indicate significant differences. 

Figure 3—Mean plant cover from plots in five habitats that had either 

experienced recent grazing or had not been grazed for 25+ years. Cover 

in most habitats was similar in grazed and ungrazed areas except for the 

riparian zone. Letters above columns indicate significant differences. 

Figure 4—Differences in average cover of all vegetation, and of four 

perennial species in plots of five habitats that had either not been grazed 

by cattle for 25+ years or had experienced recent grazing.
USDA Forest Service Proceedings RMRS-P-67. 2013 319

Trajectory and Rate of Desert Vegetation Response Following Cattle Removal R. Minckley

bosque dry habitats and only tumbleweed, Salsola tragus, occurred

in the grassland habitat (table 1). All exotic species occurred in the

grazed riparian habitat, but only two of these species occurred in the

riparian habitat that was ungrazed. The amount of cover represented

by exotic species was less than 5% in all habitats except the riparian

habitat with the lowest coverage in the desertscrub habitat. Coverage

of introduced species in the riparian habitat was 7-fold greater in the

grazed area (25.4%) than in the ungrazed area (3.4%). 

Discussion

	 Despite considerable differencesin vegetation among plots within

habitats, in the five habitats, and in the 23+ year difference in grazing

history, the pattern of vegetation response showed water availability

affected how the community composition changed and how quickly

such changes occurred. The difference was most marked in the riparian habitat where a cottonwood-dominated gallery forest is now

Figure 5—The rapid response of riparian vegetation after the San Bernardino National Wildlife Refuge was established is evident from these photographs 

from 1979 (left) and 2002 (right) of the Rio San Bernardino (= Black Draw) at the United States-Mexico border. The longer and narrower white arrows 

in both images indicate north, and the heavy short arrow in the left hand image indicates the approximate source and direction that the image on the 

right was taken. The United States-Mexico border is distinct in the right hand photo by the continuous line extending from the top to the bottom of the 

aerial photograph and is less distinct in the left hand photos where it runs from the left to right and traverses the river at the last Populus fremontii on 

the Rio San Bernardino (-Black Draw). The aerial photograph from 1979 shows an unvegetated, exposed riverbed on both sides of the border with few 

P. fremontii. Along the river in the left photo is found an almost continuous forest that has grown since 1979 and is yet to establish along the river in 

Mexico, where grazing occurred until 3 years before the photo. 

Table 1—Distribution and identity of exotic species in spring, 2004. Four plots were sampled per habitat 

and grazing history type. Percent cover is the average for each exotic species in the four plots and 

percent cover of exotics is the percentage cover by exotic species of all vegetation (not percentage of 

entire plot). The greatest species richness and overall cover of exotic species occurred in the riparian 

ungrazed habitat. Exotic species were rare or absent in drier habitats.

Habitat, Num. plots % % Vegetation

grazing history Plant species (of 4 max) Cover cover by exotics

Bosque dry, grazed – – – –

Bosque dry, ungrazed Eruca vescaria 2 0.1 0.2

Bosque wet, grazed Eruca vescaria 1 0.1

Cynodon dactylon 1 0.5 0.9

Bosque wet, ungrazed Eruca vescaria 2 0.3

Cynodon dactylon 1 0.2

Salsola tragus 2 0.2 1.5

Grass, grazed Salsola tragus 1 0.2 0.5

Grass, ungrazed – – – –

Riparian, grazed Eruca vescaria 2 0.3

Cynodon dactylon 4 11.8

Melilotus indicus 3 2.6

Salsola tragus 2 0.6

Sorghum halepense 1 4.1

Tamarix ramosissima 1 0.03 25.4

Riparian, ungrazed Cynodon dactylon 2 1.5

Sorghum halepense 4 2.0 3.4

Scrub, grazed Eruca vescaria 2 0.3 0.6

Scrub, ungrazed Eruca vescaria 1 0.4 0.8
320 USDA Forest Service Proceedings RMRS-P-67. 2013

R. Minckley Trajectory and Rate of Desert Vegetation Response Following Cattle Removal

present where cattle had been removed in 1979 (fig. 5). Although in

this habitat the cover was the greatest of any habitat sampled (fig. 3),

species richness was the lowest (fig. 2). Xeric shrub and grassland

habitatsin the San Bernardino Valley are high in species number, and

rivaled only by grazed riparian habitat where cottonwood forest had

not become established. Shrub and grassland habitats in grazed and

ungrazed areas differed much less in the species found there (fig. 2)

and the percent cover(fig. 3)thanwas observed in the riparian habitat,

indicating low water availability does not limit species richness but

slows the rate vegetation responds once grazing is reduced. 

	 In this study area, grazing depressed recruitment of cottonwoods

in the riparian habitatsresulting in eventual extirpation of the species

where permanentwaterwas present.Cottonwoods presenttodaywere

largely absent in this area when cattle grazing was first discontinued

in 1979 (fig. 5). The absence of a cottonwood canopy in the grazed

riparian area enabled numerousshrubs and annualspeciesto colonize

and persist there, many of which were found otherwise in xeric habitats. Both the lack of sunlight and excess soil moisture exclude some

plant species from riparian habitats where continuous forest canopy

occurs. Upon cessation of grazing in riparian habitats, cottonwoods

and other fast growing mesic-adapted plants become established and

eventually displace other, lower-growing, species.

	 In comparison to riparian habitat, the effect of grazing on desert

scrub and grassland habitats is more difficult to interpret because it

is not possible to disentangle if (1) grazing had little effect on these

habitats, or (2) if grazing had a large effect and the rate vegetation

responded in these habitatswasslower.Though rarely considered, the

first hypothesisis not implausible given grazing density and duration

for more distant habitats is reduced because cattle remain within a

few kilometers ofsourcesfor water, especially when airtemperatures

are high. In the San Bernardino Valley, grassland, desert scrub, and

bosque dry habitats are often further from permanent water than

riparian and bosque wet habitats. Stock tanks in the grazed area hold

water ephemerally today (author, personal observation) and probably

behaved the same in the past. Therefore, the more xeric habitats may

have experiencedlowergrazingintensitythanriparianandbosquewet

habitats. If grazing did have little effect on desertscrub and grassland

communities in the San Bernardino Valley, these data suggest that

grazing results in greater plant species richness and little change in

composition.

	 Alternately, if grazing had a strong effect of vegetation in desert

scrub and grassland, this study is consistent with the conclusion

that after grazing ends the species richness declines gradually, and

the response by vegetation is slow. The 25-year span of vegetation

change represented in this study is not long enough to distinguish

which hypothesis (or others) is most reasonable. Further monitoring

of the sites studied here and others is needed.

	 Exotic species were most diverse and represented the greatest

coverage where soil moisture was greatest and sunlight was not impeded by a cottonwood canopy (riparian, grazed habitat). The lack

of exotic species in xeric habitats is consistent with observations of

other studies (Tellman 2002). Furthermore, there are clear management implications of this pattern given that the habitats where exotic

species are concentrated are the most localized of any habitat in the

region;management to repressthe establishment and spread of exotic

species may be sometimes feasible even if resources are limited.

	 Although only eight plots were sampled per habitat in this study,

and half of these were in areas that had been recently grazed or not

grazed for 25+years,the comparisonssuggest grazing influences vegetation in most, if not all, habitats and these effects may be positive or

negative. Where surface water occurs, vegetation respondsrapidly to

reduced grazing and generates a distinctive low-diversity community

dominated by trees. These same habitats are grazed most intensively

when cattle have accessto them, which resultsin greaterspeciesrichness and changes in vegetation composition that approached those

observed in adjacent xeric habitats. In xeric habitats, such as desert

scrub, the long-term effects are more difficult to establish based on

data in this study. It is possible these habitats are little changed other

than the increased species richness related to physical disturbance of

soils and better penetration or improved emergence by seeds. If they

did change substantially when grazed, the rate of recovery appears

slow. Either way, the differences in vegetation among habitats here

illustrate the complexity managers of biodiversity are confronted

with; is grazing maintained to promote local biodiversity or should

grazing be discontinued and fewer species maintained locally?

	 Although cattle removal leads generally toward lower plant species richness, particularly in the mesic habitats, cattle removal has

a disproportionate effect in the functional diversity of the whole

ecosystem (Petchey and Gaston 2002; Svenson and Enquist 2009).

Habitats with great vertical development (gallery forests of cottonwoods) not previously found, or barely present in the grazed area,

add ecological complexity to the natural setting. In the mesic sites

with low specific diversity, the capture of carbon in thick tree trunks

is greatly increased, and the litterfall and carbon addition to soils and

watercoursesisincreased manifold. These factors, mainly associated

to increased functional diversitymight increase the overall ecosystem

functioning and the beta and gamma diversity.

	 Finally, although this study was done in 1 year, the patterns match

those recorded during a vegetation survey in 2000 (Minckley and

Burquez, unpublished report), the year after cattle were first removed

from the grazed area of this study. This matching pattern supports

the conclusion that the effects reported in this study are still related

to grazing despite a 3-year lag after grazing ended in the ungrazed

area and a 25+-year lag after grazing ended in the ungrazed area.

Acknowledgments

Long-term support has been provided to R.L. Minckley by Joe and

Valer Austin (Fundación Cuenca de los Ojos) and Bill Radke (San

BernardinoNationalWildlifeRefuge,USFWS).Financialsupportwas

provided primarily by the Southwest Consortium for Environmental

Research and Policy with additional support from National Science

Foundation grant DEB0418871.

References

Bahre, C.J. 1991. A legacy of change: Historic impact on vegetation in the

Arizona Borderlands. University of Arizona Press, Tucson.

Bahre, C.J. 1995. Human impacts on the grasslands of southeastern Arizona.

In: McClaran, M. P., and T.R. Van Devender, eds. The desert grassland.

Tucson: The University of Arizona Press: 230-264.

Barnett, D.T.; Stohlgren, T.J. 2003. A nested intensity design for surveying

plant diversity. Biodiversity and Conservation 12: 255-278.

Brown, D.E., ed. 1994. Biotic communities: Southwestern United States and

northwestern Mexico. University of Utah Press, Salt Lake City.

Bryan, K. 1925. Date of channel trenching (arroyo cutting) in the arid Southwest. Science 62: 338-344.

Bryan, K. 1928. Changes in plant associations by changes in water table.

Ecology 9: 474-478.

Bull, W.B. 1991. Geomorphic responses to climatic change. Oxford University Press.

Bull, L.J.; Kirkby, M.J. 2002. Dryland rivers: Hydrology and geomorphology

of semi-arid channels. John Wiley & Sons.

Cooke, R.U.; Reeves, R.W. 1975. Arroyos and environmental change in the

American Southwest. Oxford University Press, London.
USDA Forest Service Proceedings RMRS-P-67. 2013 321

Trajectory and Rate of Desert Vegetation Response Following Cattle Removal R. Minckley

Fukami, T.; Wardle, D.A. 2009. Long term ecological dynamics: Reciprocal

insightsinto natural and anthropogenic gradients. Proceedings oftheRoyal

Society of London, B. 272, 2105-2115.

Hastings, J.R. 1959. Vegetation change and arroyo cutting in southeastern

Arizona. Journal of the Arizona Academy of Science. 1: 60-67.

Hastings, J.R.; Turner, R.M. 1965. The changing mile. An ecological study

of vegetation change with time in the lower mile of an arid and semiarid

region. University of Arizona Press, Tucson.

Humphrey, R.R. 1987. 90 years and 535 Miles: vegetation changes along the

Mexican border. University of New Mexico Press, Albuquerque.

Minckley, R.L. 2008. Faunal composition and species richness differences

of bees (Hymenoptera: Apiformes) from two North American regions.

Apidologie 39: 176-188. 

Muller-Dombois, D.; Ellenberg, H. 1974. Aims and Methods of Vegetation

Ecology. Wiley, New York.

Petchey, O.L.; Gaston, K.J. 2002. Functional Diversity (FD),speciesrichness,

and community composition. Ecology Letters 5: 402-411.

Stohlgren, T.J.; Bull, K.A.; and Otsuki, Y. 1998. Comparison of rangeland

sampling techniques in the Central Grasslands. Journal of Range Management 51: 164-172.

Stromberg. J.C.; Tellman, B., eds. 2009. Ecology and conservation of the San

Pedro River. University of Arizona Press, Tucson.

Svenson, N.G.: Enquist, B.J. 2009. Opposing assembly mechanisms in a

Neotropical dry forest: Implications for phylogenetic and functional community ecology. Ecology 90: 2161-2170.

Tellman, Barbara, ed. 2002. Invasive exotic species in the Sonoran Region.

UniversityofArizonaPressandtheArizona-SonoraDesertMuseum,Tucson.

Turner, R.M. 1994. Warm-temperate desertlands. In: Brown, D.E., ed. Biotic

communities: Southwestern United States and northwestern Mexico. University of Utah Press, Salt Lake City: 157-221.

Turner, R.M.; Bowers, J.E.; and Burgess, T.L. 1995. Sonoran desert plants:

An ecological atlas. University of Arizona Press, Tucson.

Wagoner, J.J. 1952. History of the cattle industry in southernArizona, 1540-

1940. Social Science Bulletin 20. University of Arizona, Tucson.

Wagoner, J.J. 1960. Overstocking ranges in southern Arizona during the

1870’s and 1880’s. Arizoniana 2: 23-27.

Warshall, P. 1994. The Madrean Sky Island Archipelago: A planetary overview. Gen. Tech. Rep. RM-GTR 264. Fort Collins, CO: U.S. Department

of Agriculture, Forest Service, Rocky Mountain Research Station.

Waters, M.R.; Haynes, C.V. 2001. Late Quaternary arroyo formation and

climate change in the American Southwest. Geology 29: 399-403.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
322 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-00. 2012 323

Fire
324 USDA Forest Service Proceedings RMRS-P-00. 2012
USDA Forest Service Proceedings RMRS-P-67. 2013 325

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane

G.; Collins, Loa C., comps. 2013. Merging science and management in

a rapidly changing world: Biodiversity and management of the Madrean

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 

Soil Erosion and Deposition Before and After Fire in Oak 

Savannas

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona

Hui Chen, Aaron T. Kauffman, and Cody L. Stropki

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Daniel G. Neary

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Abstract—Effects of low severity prescribed burning treatments and a wildfire on soil erosion and deposition in the oak savannas in the Southwestern Borderlands are reported. Measurements in the spring and 

fall, respectively, characterize soil movements following winter rains and high-intensity summer rainstorms. 

Annual values are also presented. Relationships between soil erosion and deposition and precipitation 

amounts, physiographic characteristics, and vegetation were analyzed to determine possible cause-and-effect 

implications. The information should be useful in developing strategies for re-introducing more natural fire 

regimes into the oak savannas.

Introduction

	 Low severity wildfires were common in the Southwest Borderlands before Euro-American settlement. However, the severities

and frequencies of the fires in the region have been altered since the

early 1900s largely because of fire suppression policies of management agencies and livestock grazing at this time removed significant

portions of the fire-carrying herbaceous fuels (Fulé and Covington

1995).Thesepractices andpolicieshave resultedinincreaseddensities

of tree overstories on many sites making them more susceptible to

wildfires, insects, and diseases. Unwanted woody plants have often

invaded productive rangelands. As a consequence of these conditions, management agencies and their collaborators are exploring

the possibility of re-introducing a “more natural” fire regime into

the ecosystems of the region including the oak savannas situated

between the higher-elevation and more dense oak woodlands and

lower-elevation grassland-shrub communities.

	 Estimates of average soil erosion and deposition before and after

cool- and warm-season prescribed burning treatments and a wildfire

in the oak savannas are the focus of this paper. Preliminary estimates

havebeenreportedearlier(Ffolliott andothers2005).The information

presented in this paper and the earlier publication should be useful

in accessing the possible effects of re-introducing a more natural fire

regime on soil movement in the oak savannas of the Southwestern

Borderlands region.

Cascabel Watersheds

	 Twelvewatersheds,ranging from20 to almost 60 ac in size, located

in the Peloncillo Mountains of southwestern New Mexico were the

study site. The total area of these watersheds, called the Cascabel

Watersheds, is 451.3 ac. They are situated between 5,380 and 5,590

ft in elevation. The nearest long-term precipitation station indicates

that annual precipitation averages 21.8 ± 1.2 in., with nearly one-half

falling in summer rainstorms. Streamflow originating on the watersheds is intermittent. High flows can be generated by high-intensity

summer rainstorms (Gottfried and others 2006). Most of the surface

soils are very gravelly or very cobbly sandy loams or sandy clay

loams (Robertson and others 2002). Additional vegetative, geologic,

physiologic, and hydrologic characteristics of the watersheds are

described elsewhere (Ffolliott and others 2008, Neary and Gottfried

2004, Youberg and Ferguson 2001, and others) and, therefore, are

not presented.

Prescribed Fire Treatments and Wildfire

	 The original objective of the research program on the Cascabel

Watersheds was to compare the effects of cool-season (November

through April) and warm-season (May through October) prescribed

burningtreatmentsonecosystemresourcesandhydrologicfunctioning

ofthe watersheds with similar evaluations on unburned watershedsto

assessthe impactsoftheburningtreatments.Followingthe calibration
326 USDA Forest Service Proceedings RMRS-P-67. 2013

Ffolliott and others Soil Erosion and Deposition Before and After Fire in Oak Savannas

period, four of the watersheds were burned during the cool-season

in early March 2008. Three of the four watersheds to be burned in

the warm-season were burned on May 20, 2008, with burning of the

fourth watershed scheduled for a later date. However, wind gusts of

up to 60 mph blew burning embers onto the remaining watershed to

be burned and the four control watersheds in the morning of May

21, 2008. The resulting Whitmire Wildfire crossed the watershed

boundaries and spread out to burn nearly 4,000 ac. The current research program, therefore, is evaluating the effects of the prescribed

burning treatments andthewildfire on the features originally sampled,

including soil erosion and deposition.

Fire Severities

	 A system relating fire severity to the soil-resource response to

burning (Hungerford 1996) was used to classify the severities of the

prescribed burning treatments and wildfire on the watersheds. Classifications at the sample plots on the watersheds (see below) were

then extrapolated to a watershed-basisto determine the proportion of

the watersheds that were unburned or had burned at low, moderate,

and high severities. It was determined that all of the watersheds had

burned at low severities (Stropki and others 2009).

Study Protocols 

Sampling Basis

	 Between 35 and 45 sample plots that had been established along

transects perpendicular to the main stream systems on each of the

Cascabel Watersheds were the basis for obtaining the measurements

of soil erosion and deposition. Intervals between these plots varied

depending on the size and configuration of the watershed. A total of

421 plots were located on the 12 watersheds. These plots have also

been used in other studies of ecosystem resources on the watersheds

(Gottfried and others 2007).

Measurements of Soil Erosion and Deposition

	 Three capped pins were placed around every third plot on the

watersheds to measure soil erosion and deposition. Measurements

of soil loss beneath a cap (soil erosion) or soil accumulation above a

cap (deposition) were made in the spring and fall to characterize soil

movementsfollowing periods ofwinterrains and summerrainstorms,

respectively. Occasionally, there was no change in the soil surface

beneath the cap in which case either the magnitudes of soil erosion

and deposition between the successive measurements equaled each

other or (what is less likely) neither erosion nor deposition occurred.

The capped pins were re-set to be flush with the soilsurface after each

measurement to facilitate subsequent measurements. Measurements

obtained at a plot were averaged to estimate soil movements at the

plot and the plot measurements on a watershed were then averaged

to describe soil erosion and deposition on a watershed-basis. A bulk

density value of 70.5 lbs/ft3 was used to convert the measurements

of soil erosion and deposition to corresponding measures in tons per

acre on a watershed-basis.

	 Initial measurements were made in the fall of 2004 with pre-fire

measurements continuinginthe springandfalluntiltheburningevents

occurred. Post-fire measurements were initiated in the spring of 2008

shortlyfollowingthe cool-seasonburnswiththesemeasurementsthen

made in the fall of 2008 and the spring and fall of 2009 and 2010.

Analysis

	 Measurements of soil erosion and deposition were analyzed separately because they are separate processes of soil movement. Results

of the Shapiro-Wilk test of normality indicated that the frequency

distributionsforsoil erosion and deposition were non-normal. Transformationsfailedtonormalizethedistributions.Therefore,occurrences

ofstatistical differencesin the measurements were determined by the

non-parametric Mann-Whitney test of significance (Zar 1999). This

test was applied because the estimates of the respective processes

were independent of each other. Plots with no measurable change in

soil movement were excluded from the analyses. Differences were

evaluated at the 0.10 level of significance.

Results and Discussion

	 Measurements of soil erosion and deposition are summarized by

bar graphs (see below) showing the magnitudes of the respective

processes before and after the burning events. Inferences relating to

differencesin the magnitudes ofthe two processes are not necessarily

valid, however, because the frequency distributions of the data used

in developingthebar graphswere non-normal.Significantdifferences

in the measurements of soil erosion and deposition were determined

through interpretations of the Mann-Whitney test.

Pre-Fire

	 There were no statistically significant differences in either soil

erosion or deposition among the individual watersheds throughout

the pre-fire period. Therefore, the respective data sets were pooled

for analysis.

Soil erosion—Soil erosion measurements obtained in the spring

following the winter rains were compared to the fall measurements

after the summer rainstormsto determine whetherseasonal or annual

differences occurred. Spring measurements were statistically similar

to the fall measurements with an average of 14.6 t/ac. Annual soil

erosion averaged 14.2 t/ac. Differences in soil erosion within the

seasons and years of measurement were insignificant.

Soil deposition—Soil deposition after the winter rains was also

comparedtothemeasurementstakenfollowingthe summerrainstorms

to determine if seasonal differences occurred before the burning

events. It was found that the deposition of soil in the spring differed

from the depositions in the fall, 4.6 and 7.8 t/ac, respectively, with

annual soil deposition averaging 6.3 t/ac. Differences in soil deposition within the seasons and years of measurement were inconsistent

and, therefore, considered insignificant.

	 There were no consistent relationships in the magnitudes of soil

erosion or deposition and rainfall patterns or the physiographic characteristics or vegetation surrounding the sample plots.

Post-Fire

	 Comparisons of the post-fire data sets indicated that there were no

significant differences in soil erosion or depositions on the respective watersheds experiencing the prescribed burning treatments or

wildfire. Therefore, the data sets were pooled for comparison with

pre-fire values.

Soil erosion—Soil erosion following the prescribed burning

treatments and wildfire was greater (19.1 t/ac) than measurements

of pre-fire soil erosion following winter rains but lower after the
USDA Forest Service Proceedings RMRS-P-67. 2013 327

Soil Erosion and Deposition Before and After Fire in Oak Savannas Ffolliott and others

summer rainstorms (10.5 t/ac) in relation to pre-fire measurements

(fig. 1). There was no significant difference in annual soil erosion as

a consequence of the burning events. The seasonal measurements of

soil erosion appeared to “balance” each other on an annual basis.

	 That the annual post-fire erosion of soil after the burning events

was within the range of the pre-fire value waslikely a result ofsimilar

erosive forces generated by rain drops impacting on the soil surface

throughout the pre- and post-fire periods. Furthermore, the absence

of widespread waterrepellentsoils afterthe burns(Stropki and others

2009), and, as a consequence, the likelihood of a little change in the

overland flows of water necessary to dislodge soil particles from a

site occurred. There also was little evidence of increased rill formations on the hillslopes of the watersheds following the three burning

events.

Soil deposition—Post-fire deposition of soil following the winter

rains was statistically similar to the pre-fire deposition (fig. 2). However, the deposition measured after high-intensity summer rainfalls

was significantly greater (11.7 t/ac) following the burning events for

a reason unknown to the authors. Soil deposition on an annual basis

was also greater following the burning events (9.4 t/ac) because of

the difference in the fall measurements.

	 Similar to the pre-fire findings,soil erosion and deposition following the prescribed burning treatments and wildfire were not related

to the rainfall events, physiographic characteristics, or vegetation.

Summary

	 The informationpresentedinthis andthe earlierpaperonsoilmovement on the Cascabel Watersheds (Ffolliott and others 2005) should

be useful to people interested in knowing the effects ofre-introducing

a more natural fire regime into the oak savannas of the Southwestern

Borderlands. However, knowledge of the effects of prescribed burning treatments of higher fire severities, in other seasons, and on the

array of ecosystem resources and factors affecting the hydrologic

functioning of watersheds in the oak savannas is necessary before

attempting to introduce more natural fire regimes through prescribed

burning treatments.

Acknowledgments

	 This study was supported by the Southwestern Borderlands Ecosystem Management Unit of the Rocky Mountain Research Station,

Phoenix,Arizona, the U.S. Forest Service National Fire Plan, and the

Arizona Agricultural Experiment Station, University of Arizona.

 References

Ffolliott, P. F.; Gottfried, G. J.; Stropki, C. L. 2008. Vegetative characteristics

andrelationshipsoftheSouthwesternBorderlands.Res.Pap.RMRS-RP-74.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 16 p.

Ffolliott, P. F.; Stropki, C. L.; Gottfried, G. J.; Neary, D. G. 2005. Initial estimate of soil erosion on the Cascabel Watersheds in the oak savannas of

theMalpaiBorderlandsregion. Hydrology andWaterResourcesinArizona

and the Southwest. 35:51-52.

Fulé, P. Z.; Covington, W. W. 1995. Changes in fire regimes and forest structures of unharvested Petran and Madrean pine forests. In: DeBano, L. F.;

Ffolliott,P.F.;Ortega-Rubio,A.;[andothers],tech. coords.Biodiversityand

management oftheMadreanArchipelago:The Sky Islands ofsouthwestern

United States and northwestern Mexico. Gen. Tech. Rep. RM-GTR-264.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Forest and Range Experiment Station: 408-415.

Gottfried, G. J.; Neary, D. G.; Ffolliott, P. F. 2007. An ecosystem approach

to determining the effects of prescribed fire on Southwestern Borderlands

oak savannas: a baseline study. In: Masters, R. E.; Galley, K. E. M., eds.

Fire in grassland and shrubland ecosystems: proceedings of the 23rd Tall

Timbers Fire Ecology Conference. Tallahassee, FL: Timbers Research

Station: 140-146.

Gottfried, G. J.; Neary, D. G.; Ffolliott, P. F.; Decker, D. D. 2006. Impacts of

a high-intensity summer rainstorm on two oak savannas watersheds in the

Southwestern Borderlands. Hydrology and Water Resources in Arizona

and the Southwest. 36:67-73.

Hungerford, R. D. 1996. Soils: Fire in ecosystem notes: unit II-I. Marana,AZ:

National Advanced Resources Technology Center.

Neary, D. G.; Gottfried, G.J. 2004. Geomorphology ofsmall watershedsin an

oak encinal in the Peloncillo Mountains. Hydrology and Water Resources

in Arizona and the Southwest. 34:65-71.

Robertson, G.; Damrel, D.; Hurja, J.; Leahy, S. 2002. Terrestrial ecosystem

survey of the Peloncillo watershed study area. Draft Rep. Albuquerque,

NM:U.S.DepartmentofAgriculture,ForestService,SouthwesternRegion.

Figure 1—Seasonal and annual soil erosion on the Cascabel Watersheds 

before and after the prescribed burning treatments and wildfire. Averages 

for the respective study periods are shown in the figure.

Figure 2—Seasonal and annual deposition of soil on the Cascabel 

Watersheds before and after the prescribed burning treatments and 

wildfire. Averages for the respective study periods are presented in 

the figure.
328 USDA Forest Service Proceedings RMRS-P-67. 2013

Ffolliott and others Soil Erosion and Deposition Before and After Fire in Oak Savannas

Stropki, C. L.; Ffolliott, P. F.; Gottfried, G. J. 2009. Water repellent soils following prescribed burning treatments and a wildfire in the oak savannas of

theMalpaiBorderlandsregion. Hydrology andWaterResourcesinArizona

and the Southwest. 39:5-8.

Youberg,A.; Ferguson, C.A.. 2001. Geology and geomorphology of 12 small

watersheds in the Peloncillo Mountains, central portion of the Malpai

Borderlands project area, Hidalgo County, New Mexico. Open-File Rep.

01-05. Phoenix, AZ: Arizona Geological Survey. 22 p.

Zar, J. H. 1999. Biostatistical analysis. Upper Saddle River, NJ: Prentice

Hall. 663 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 329

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Hillslope Treatment Effectiveness Monitoring on 

Horseshoe 2 and Monument Fires

Carly Gibson

USDA Forest Service, Stanislaus National Forest, Sonora, California

Ann Youberg

Arizona Geological Society, Tucson, Arizona

Marc Stamer

USDA Forest Service, San Bernardino National Forest, Fawnskin, California

Abstract—Between February and July, 2011, over 360,000 acres burned across the Coronado National 

Forest during one of the most active fire seasons in recorded history. Burned Area Emergency Response 

(BAER) Teams evaluated post-fire watershed conditions and prescribed treatments based on threats to 

known values at risk. Hillslope stabilization treatments were prescribed and implemented for areas of 

high soil burn severity on both the Horseshoe 2 and Monument Fires. These treatments consisted of 

seeding on the Horseshoe 2 Fire and application of agricultural straw mulch and seed on the Monument 

Fire. Initial monitoring results indicated one of three seeded species (Hordeum vulgare) emerged in both 

burned areas, slightly improving effective ground cover in both treatments. However, seeding treatments 

failed to meet monitoring success criteria for the Horseshoe 2 and Monument Fires. Hillslope erosion 

was reduced where mulch treatment was applied correctly and where slope gradients were moderate 

on the Monument Fire, and appeared to contribute to seeded species cover. In the Horseshoe 2 Fire, 

hillslope erosion was high on the treatments transects and was not reduced by seeding alone. A need 

for additional monitoring in spring 2012 exists and would improve the current understanding of the 

effectiveness of hillslope treatments. 

Introduction

	 The summer of 2011 saw record wildfires across the southwestern

United States. In southeastern Arizona, the Horseshoe 2 Fire burned

approximately 222,954 acres in the Chiricahua Mountains (http://

inciweb.org/incident/2225/), and the Monument Fire burned approximately 30,526 acresin theHuachucaMountains(www.inciweb.

org/incident/2324/) (fig. 1). Both fires burned watersheds that drain

onto developed private lands and rural ranches. InArizona, monsoon

rains immediately follow wildfire season and are often how fires are

ultimately extinguished. The quick, intense burst of rainfall from

relatively common (<2-5 yr frequency) storms can generate large

floods and debris flows in watersheds disturbed by wildfires. U.S.

Forest Service (USFS) Burned Area Emergency Response (BAER)

assessments were completed for both fires. Soil burn severity maps

show 12% of the Horseshoe 2 Fire burned at high severity and 30%

at moderate severity, while 7% of the Monument Fire burned at high

severity and 39% at moderate severity. Moderate to high soil burn

severity in both fires occurred on moderate to very steep slopesin the

upper watersheds. The BAER teams conducted hydrologic analyses

of post-burn conditions using a 5-year return-interval storm with an

intensity of 1/

2 inch/hour. Results indicated an estimated increase of

post-fire peak flows from 2-15 times in the Horseshoe 2 burned area

(USDA 2011a), and from 3-10 times in the Monument burned area

(USDA 2011b).

	 Tomitigatepredictedincreasesinpost-fire runoff andconsequential

risks posed to life, property, and soil productivity within and near the

burned areas, hillslope treatments were prescribed for selected areas

of moderate and high soil burn severity on USFS-managed lands.

Aerial seeding was applied to treatment areas in the Horseshoe 2

burned area from July 16 through July 18 and in the Monument Fire

on July 29. Seedmixturesincluded Hordeum vulgare (annual barley),

Bouteloua gracilis (blue grama) and Pascopyrum smithii (western

wheatgrass) for the Horseshoe 2 Fire, and H. vulgare, B. gracilis and

Elymus trachycaulus (slender wheatgrass) for the Monument Fire.

The seeding was applied, with variable seed coverage, by fixed-wing

aircraft. Agricultural straw mulch was applied over seeded units in

the Monument Fire from August 2 to 17, 2011; no agricultural straw

was used to stabilize hillslopes in the Horseshoe 2 Fire.

	 The objective of this study was to evaluate the initial effectiveness of hillslope treatments within both burned areas. To evaluate if

treatments were successful or not within the first year of application,

the study plots were monitored to determine if (1) seeded species

germinated and became established, (2) seeded species provide effective cover for soil stabilization, (3) straw mulch cover was uniform

throughout Monument Fire treatment units, (4) straw mulch treatment
330 USDA Forest Service Proceedings RMRS-P-67. 2013

Gibson and others Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires

successfully reduced soil erosion in the Monument Fire, and (5) wind

was a factor in moving the straw mulch in the Monument Fire.

Methods

Site Selection

	 Burned areas suitable for seeding and mulching treatments had

moderate tohighsoil andvegetationburnseveritywithslopegradients

between 40 and 60%.Appropriate areasfor treatment were identified

by the BAER teams (USDA 2011a,b,c). Within the treatment areas,

preliminary transect locations were selected on the basis of soil burn

severity (USDA 2011a,b) vegetation burn severity (USDA 2011c),

aspect (table 1), and accessibility. Potential treatment and control

transect locations were located in the field based on ArcGIS-derived

UTMcoordinates.Transectswere establishedinareasthatrepresented

average hillslope conditions while avoiding natural drainages which

may have contributed to pre-fire hillslope erosion that would not have

been mitigated by treatments The number and location of control

transects were limited as most of the potential treatment areas were

seeded (Horseshoe 2) or seeded and mulched (Monument).

Data Collection and Analysis

	 Two rain gauges were installed on the Monument Fire and one

was installed in the study area of the Horseshoe 2 Fire (northern half

of the fire) in early July with a second gauge installed in late July.

Rain fell on both fires before any of the hillslope treatments could

be applied. The first significant rains fell on the Monument Fire on

July 10 (table 2) and on the Horseshoe 2 Fire on July 11 (table 3).

Both storms produced debris flows and floods. The NOAAAtlas 14

classifies both storms as a 2-year/30-minute frequency. During July

and August, 10 storms on the Monument Fire (table 2) and 11 storms

on the Horseshoe 2 Fire (table 3) produced enough runoff to generate

either debris flows or floods with peak discharges sufficiently large

to threaten or cause damage to values at risk based on reports from

BAER team implementation teams and local residents.

	 Thirty-meter transects were established along contour to collect

rill network density and effective ground cover data (fig. 2). For each

transect, the width and depth of each rill was measured for the first

10 m, distance between rills was measured within the first 15 m, and

total number of rills was tallied for the entire length. Average values

for distance between rills and rill width and depth measurements for

each fire were calculated to compare treatments with controlsections.

Rill cross-sectional areas, ranges, averages, and standard deviations

were calculated by treatment for each fire.

	 Effective ground cover (EGC) data collected from 1 m square

quadrats included native vegetative cover, seeded species cover and

count, large woody debris, and litter (Brady and Weil 2000; DeBano

and others 1998; Pannkuk and Robichaud 2003). Agricultural straw

cover and clumps of agricultural straw within 1 m of treatment

transects were also measured in the Monument Fire. Square meter

quadrats were read every 3 meters for the length of each transect.

Cover frequency index (CFI) was calculated for each EGC variable

for treatment and control transects sampled in each burned area.

This metric combines frequency of occurrence and absolute percent

cover for each variable analyzed (Benkobi and Uresk 1996; USDA

2006). Ground cover was determined to be effective at reducing

erosion when all variables measured contributed to 70% or greater

cover (Pannkuk and Robichaud 2003; Robichaud 2005). Successful

treatmentimplementationandgerminationofseededspeciesincluded

the presence of all three seeded species, an average of greater than

20 seeded individuals in treatment quadrats, and a CFI of twice the

overall vegetation CFI for treatment transects (Johnson 2004).

	 Data were collected from September 22 through 27, 2011. Six

treatment and three control transects were established in the Horseshoe 2 burn area and seven treatment and two control transects were

established in the Monument burned area (table 1). One potential

treatment transect in the Monument Fire was abandoned due to safety

concerns.

Results

Rill Density 

	 Measurements were collected along transects to obtain rill density

and cross-sectional areas, although not all transects intersected rills.

On the Horseshoe 2 Fire, rill measurements were collected on four

of six of the treated transects and two of three of the control transects

(tables 4 and 5, fig. 3). One control transect intersected a single rill

beyond 15m; therefore, nomeasurementswere collectedOn average,

there were a greater number of rillsin the treatment transects, and rill

cross-sectional areas were 64% lower for treatment transectsthan for

control transects.

	 On the Monument Fire, rill measurements were collected on four

of seven treated transects and two of two control transects (tables 6

and 7, fig. 4). One treated transect intersected three rills beyond 15

meters;therefore,nomeasurementswere collectedforthistransect.On

average, there were a greater number of rills in the control transects.

However, rill cross-sectional areas were 37% lower for treatment

transects than for control transects.

Effective Ground Cover

H. vulgare was the only seeded species observed during data

collection. This grass species was present in 71% of Horseshoe 2

treatment quadrats, accounting for an average of 10.7% cover and 7.6

CFI (table 8), and in all Monument treatment quadrats, accounting

for an average of 7.8% cover and 7.6 CFI (table 9). H. vulgare cover

on the Horseshoe 2 Fire was highest where soil and vegetation burn

severity was moderate and lowest where soil and vegetation burn

Figure 1—Location of Horseshoe 2 Fire and Monument Fire in southeastern Arizona.
USDA Forest Service Proceedings RMRS-P-67. 2013 331

Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires Gibson and others

Table 2—Significant rainfall events from two ALERT gauges within the Monument Fire burned area. Dates of hillslope 

treatments are shown in right column.

Miller Canyon ALERT Gauge Ash Canyon ALERT Gauge

Treatments Date

Storm 

Total 

(mm)

Storm 

Duration 

(h:mm:ss)

Average 

Storm 

Intensity 

(mm/hr)

Storm 

Total (in)

Storm 

Duration 

(h:mm:ss)

Average 

Storm 

Intensity 

(mm/hr)

10-Jul 41.66 1:04:00 39.1

20-Jul 29.46 1:17:16 22.9 6.1 2:14:48 2.7

23-Jul 9.14 0:12:00 45.7 10.2 0:14:17 42.7

26-Jul 8.13 0:35:32 13.7

28-Jul 19.30 5:04:57 3.8 31.5 4:08:01 7.6

29-Jul 2.03 0:42:31 2.9 11.2 1:20:20 8.3 Seeding Applied

31-Jul 12.19 4:43:42 2.6 31.5 4:01:28 7.8

11-Aug 3.05 0:44:49 4.1 8.1 1:08:45 7.1 Straw Mulch Applied 

Aug 2-17 13-Aug 6.10 3:00:32 2.0 13.2 1:39:21 8.0

20-Aug 22.35 1:00:07 22.3 17.3 0:21:18 48.7

22-Aug 2.03 0:12:10 10.0 15.2 0:24:24 37.5

23-Aug 9.14 0:11:22 48.3

Table 1—Transect locations, soil burn severity, vegetation burn severity, 

aspect and transect sample type. (c = control, untreated)

Fire Transect

Burn Severity

Aspectc Sample Type

Soila Vegetationb

Monument

1 moderate high east treatment

2 moderate high east treatment

3 moderate high north treatment

4 high high east treatment

5 high high north treatment

6c moderate high north control

7c high high east control

8 moderate high east treatment

10 high high north treatment

Horseshoe 2

11 high high north treatment

12 high moderate south treatment

13c high high south control

14 moderate moderate west treatment

15c moderate moderate west control

16 mixed mixed west treatment

17 mixed mixed south treatment

18 high high north treatment

19c high high north control

a

Soil burn severity derived from BAER Assessment Team’s Final Soil Burn Severity GIS (USDA 

2011a & b).

b

Vegetation burn severity downloaded from USFS Remote Sensing Application Center, Salt 

Lake City, 09/17/2011.

c

Aspect generated from 30 meter DEM in ArcMap 9.3.1 with Spatial Analyst (ESRI 2011). 
332 USDA Forest Service Proceedings RMRS-P-67. 2013

Gibson and others Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires

Table 3—Significant rainfall events from two gauges within the Horseshoe 2 Fire burned area. Dates of hillslope 

treatments are shown in right column.

UA/B2 Chiri Gauge & NWS KC2CPZ1 Weather Station

Treatments Date Storm Total (mm) Storm Duration 

(h:mm:ss)

Average Storm Intensity 

(mm/hr)

11-Jul 54.4 1:38:31 33.1

12-Jul 8.6 0:35:52 14.4 Seeding Applied 

July 16-18 26-Jul 13.0 0:32:45 23.7

28-Jul 7.9 0:59:29 7.9

3-Aug 6.4 0:13:17 28.9

9-Aug 6.0 0:11:35 31.1

11-Aug 24.6 2:40:59 9.2

13-Aug 18.0 1:25:56 12.6

15-Aug 24.6 0:51:04 28.9

24-Aug 10.9 0:47:38 0.54

Figure 2—Transect sample design. Total number of rills were recorded for the length of transects and effective ground cover variables measured from quadrats.

Table 4— Distance between rills for the Horseshoe 2 Fire. Transect

Number of rills 

(0.00- 15.0 m)

Distance between rills (m)

Range Average

Treatment

14 0.1 to 3.85 0.83

0 NA NA

0 NA NA

5 0.12 to 1.55 0.57

7 0.5 to 5.75 2.00

18 0.08 to 2.05 0.52

Control

0 NA NA

3 0.15 to 6.3 2.29

0 NA NA

Table 5— Rill density and cross-sectional area measurements for the 

Horseshoe 2 Fire. Transect

Number of 

rills 

(0.00- 10 m)

Rill width (m) Rill depth (m)

Range Average Range Average

Treatment

9

0.15 to 

0.7 0.31

0.005 to 

0.04 0.02

0 NA NA NA NA

0 NA NA NA NA

5

0.07 to 

0.12 0.10

0.005 to 

0.03 0.02

4

0.05 to 

0.02 0.12

0.02 to 

0.03 0.02

11

0.09 to 

0.6 0.28

0.005 to 

0.06 0.02

Control

0 NA NA NA NA

3 0.3 to 1.3 0.67 0.03 0.03

0 NA NA NA NA
USDA Forest Service Proceedings RMRS-P-67. 2013 333

Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires Gibson and others

Figure 3—Horseshoe 2 average rill cross-sectional area with error bars 

for treatment and control samples. 

Table 6—Distance between rills for the Monument Fire. Transect

Number of 

rills 0- 15 m

Distance between rills (m)

Range Average

Treatment

15 0.04 - 6.3 0.68

0 NA NA

0 NA NA

22 0.04 - 2.92 0.55

0 28.35 NA

0 NA NA

0 NA NA

Control

19 0.09 - 2.95 0.69

12 0.06 - 2.5 0.83

Table 7—Rill density and cross-sectional area measurements for the 

Monument Fire. Transect

Number 

of rills 

0- 10 m

Rill width (m) Rill depth (m)

Range Average Range Average

Treatment

11 0.08 - 

0.48 0.26 0.015 - 0.06 0.03

0 NA NA NA NA

0 NA NA NA NA

8 0.05 - 

0.39 0.2025 0.005 - 0.035 0.013

0 NA NA NA NA

0 NA NA NA NA

0 NA NA NA NA

Control

11 0.07 - 

0.24 0.13 0.0025 - 0.03 0.01

10 0.11 - 

0.65 0.32 0.005 - 0.03 0.019

Figure 4—Monument Fire average rill cross-sectional area with error 

bars for treatment and control samples. 

severity was high. The highest and lowest H. vulgare cover on the

Monument Fire were recorded where soil burn severity was moderate

and vegetation burn severity was high. Four treatment transects on

the Horseshoe 2 Fire and one treatment transect on the Monument

Fire had an average of less than 20 individuals per quadrat. Average

percent cover was less than 20% on all treatment transects.

	 Other vegetative cover was comprised primarily of native species. No invasive species were observed during data collection. The

CFI of other vegetation was 10.0 for treatment and 4.8 for control

transects on the Horseshoe 2 Fire and 6.3 for treatment and 2.5 for

control transects on the Monument Fire. Non-vegetation effective

ground cover was comprised of litter and rock with the addition of

straw on the Monument Fire. Total non-vegetation effective ground

cover CFI was 22.8 for all treatment transects combined and 11.5 for

all control transects combined (table 9). The total number of straw

clumps within 1 m of a treatment transect ranged from zero to eight

on four of seven transects, indicating poor treatment application on

some of the treatment areas. Non-vegetative ground cover for

both fires had higher CFI for treatment and controls than overall

vegetation CFI.

Seeding Success Criteria Evaluation

	 Treatments failed to meet success criteria for seed treatment application for both fires (table 10). Success criteria were based on

monitoringmethodsoftheNuttallComplexontheCoronadoNational

Forest (Johnson 2004) and Santiago Fire on the Cleveland National

Forest (Wohlgemuth and others 2010).

Discussion

	 Both the Horseshoe 2 and Monument Fire burn areas experienced

significant precipitation prior to hillslope treatment implementation.

Followingtreatmentimplementation,the treatedareaswere exposedto

numerous otherstorms. These precipitation events varied in intensity

and location, and contributed to hillslope erosion before and after

treatment implementation.

	 After initial data collection in September of 2011, 33% of seeded

species (mostly H. vulgare) had established in treatment units in

both burned areas. Although Bouteloua gracilis (blue grama) and

Pascopyrum smithii(westernwheatgrass)were absentfromtreatment

transects, these species may yet emerge following 2011-2012 winter
334 USDA Forest Service Proceedings RMRS-P-67. 2013

Gibson and others Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires

rains. Germination of seeded species was somewhat successful as

only two of three seeded species were observed. Seeding slightly

improved EGC on hillslope treatment units on both fires where

seedsremained onsite following exposure to rainfall and slopes were

moderate (closer to 40%). Overall treatment CFI was higher than

controls and was attributed to both seeded and non-seeded species

cover. Lack of consistently high emergence or high CFI of seeded

species acrosstreatment transects was attributed to seed mobilization

during rainstorm events as high H. vulgare cover was observed on

roads and in riparian areas downslope and downstream from treatment units in both burned areas (C. Gibson, personal observation).

Agricultural straw mulch over seeding treatment on the Monument

fire is assumed to have contributed to the higher average number of

H. vulgare individuals because of higher number of individuals and

more even distribution than observed in the Horseshoe 2 treatment

quadrats (C. Gibson, personal observation). Site conditions such as

steeperslopes(closerto 60%) andmore concentrated runoffmay have

contributed to poor vigor and lower cover of H. vulgare in this burned

area when compared to post-treatment conditions in the Horseshoe 2

burned area (C. Gibson, personal observation).

	 Rill densities were high and distance between rills low on several

treatment transects. This may indicate that treatment implementation

was not entirely effective at stabilizing soil and reducing hillslope

erosion for either burned area. The timing of initial rill development,

however, occurred during thefirstsignificantstorms and priorto treatment. It is not possible to definitively say what effect treatments had

Table 10—Seeding treatment success matrix.

Fire Success criteria

Treatment 

transects 

meeting criteria 

(%)

Horseshoe 2

Average of 20 seeded individuals in 

quadrats 33

Seeded species CFI twice the 

overall vegetation CFI 33

Germination of all seeded species 0

Monument

20 seeded individuals in treatment 

quadrats 86

Seeded species CFI of twice the 

overall vegetation CFI 42

Germination of all seeded species 0

Table 9—Monument Fire effective ground cover variables cover frequency index by sample type.

EFG variable Treatment Control Treatment Control

Seeded species 

(treatment)

Hordeum vulgare 7.59 NA

6.46 2.54 Bouteloua gracilis 0 NA

Elymus trachycaulus 0 NA

Other vegetation 6.26 2.54

Straw 36.39 NA

Litter 2.96 1.15 22.78 11.54

Rock 32.34 50.35

Table 8—Horseshoe 2 Fire effective ground cover variables cover frequency index by 

sample type.

EFG variable Treatment Control Treatment Control

Seeded 

species 

(treatment)

Hordeum vulgare 7.62 NA

21.64 4.85 Bouteloua gracilis 0 NA

Pascopyrum smithii 0 NA

Other vegetation 10.02 4.85

Litter 4.85 27.14

40.58 50.75

Rock 32.49 23.26
USDA Forest Service Proceedings RMRS-P-67. 2013 335

Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires Gibson and others

on hillslope stabilization because rills were not measur ed priorto and

after the treatments. Seeded species that established on the hillslopes

appeared to provide some effective ground cover on upper and lower

slopes. On the Horseshoe 2 Fire, rock and litter contributed more to

total EGCthan all vegetation combined.Rock wasthe overall highest

contributor to EGC for treatment and control transects. Treatment

transects had higher overall vegetative cover than control transects,

but this is likely due to the high frequency and cover of other vegetation detected in frames of treatment transects and low total number

of control transects. Although H. vulgare increased EGC, uniform

establishment and healthy vigor was inconsistent among treatment

transects. This was particularly evident on steep slopes and where

soil burn severity was high as H. vulgare was seldom detected in

these locations and lacked robust stature that would contribute to

litter cover following senescence.

	 On the Monument Fire, the application of agriculturalstraw mulch

over the seed treatment contributed to EGC and slope stabilization.

When considering all treatment and control transects, hillslope seeding treatment failed to meet criteria for improving EGC and has not

sufficiently mitigated hillslope erosion. However, agricultural straw

mulch did improve overall EGC and appeared effective at reducing

hillslope erosion where slopes were gentle to somewhat moderate.

Overall non vegetative effective ground coverCFIfortreatments was

double that of controls, indicating agriculturalstraw mulch treatment

was successful.

	 The low total number of samples and the variety of slope characteristics encountered where transects were established contributed

to variable hillslope results. This is particularly evident by the range

of rill densities measured on treatment and control transects. It is

uncertain whether hillslope rill characteristics were the result of a

particularly intense localized storm or hillslope-treatment failure

since data were collected following several significant storms and

monsoonal moisture is highly variable. Despite a clear reduction in

rill dimensions, on average, there is no significant (statistical) difference between treatment and control. A higher sample size may show

the trend to be valid, but a more precise estimate of the difference is

necessary to assess treatment and cost effectiveness.

Conclusions and Recommendations

	 Publishedstudiesofthe effectivenessofpost-fire seedingtreatments

have occurred in southern California chaparral and in various conifer

ecosystems of the western United States (Beyers 2004; Robichaud

2005). Mulching (60% cover or greater) has been shown as the most

effective treatment for reducing erosion, especially when protection

is needed from the first storms that occur after fire (Beyers 2004).

Seeding is likely to provide effective control of erosion during the

first year only a third of the time (Beyers 2004). The effectiveness of

seeding in Arizona may be even less due to the intensity of rainfall

duringmonsoonalstorms.MoodyandMartin (2009)showedpost-fire

sedimentyieldsduringthefirst2yearsfollowingfire are stronglytiedto

rainfallintensity.Rainfallregimesbasedonthe2-year30-minute rainfall intensity place southeasternArizona into the 2 highest categories

(Arizona High – Horseshoe 2 Fire, andArizonaExtreme –Monument

Fire) for the entire western United States (Moody and Martin, 2009).

It is common that rains occur prior to treatment implementation as

seen on these fires because monsoon and wildfire season overlap in

Arizona. Thus, it is essential to understand if hillslope treatments are

effective for this area of the country.

	 InitialmonitoringresultsshowthatmulchingontheMonumentFire

appeared to be effective in reducing rill development and hillslope

erosion on gentle slopes (closer to 40%) but additional field data are

needed to verify this. No evidence of wind dispersal was observed,

however, poor application of straw (clumps) was observed. Implementation reportsindicated some ofthe straw bales were not properly

prepared for dispersal, which resulted in straw clumps.

	 Initial observations suggest that seeding does not appear to be

effective in this environment especially if the objective is slope

stabilization prior to first damaging storm event. To improve the

current knowledge of appropriate hillslope treatments and seeding

species for burned landscapes in southeastern Arizona, further data

collection fromtransects established in September of 2011 isstrongly

recommended for the second and third year post-fire. In addition to

data collection from the 2011 transects, establishment of additional

treatment and control transects is highly recommended to provide

sufficient information for comparison of treated and untreated areas

within the Horseshoe 2 and Monument Fires. Additional transects of

both treatment and controls will aide in identifying whether or not

results are statistically significant. Measurements specific to slope

degree and length will aide in identifying appropriate slope characteristics for future post-fire hillslope seeding and agricultural straw

mulch treatments. This will also provide an opportunity to determine

whether Bouteloua gracilis, Elymus trachycaulus or Pascopyrum 

smithii emerge in seeded sites. This is particularly important in the

case of P. smithii, which is a rhizomatous grass that is capable of

displacing native vegetation.

References

Benkobi,L;D.W.Ursek.1996.Seralstage classificationandmonitoringmodel

for big sagebrush/western wheatgrass/blue gramma habitat. In: Barrow,

J.R.; E.D. McArthur; R.E. Sosebee; R.J. Tausch, comps. 1996. Shrubland

ecosystem dynamics in a changing environment. 1996 May 23-25; Las

Cruses, NM. Procedings INT-GTR-338. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Research Station.

Beyers, Jan L. 2004. Postfire seeding for erosion control: effectiveness and

impactsonnativeplant communities.ConservationBiology.18(4):947-956.

Brady, N.;Weil,R. 2000. Elements ofthe nature and properties ofsoils. Upper

Saddle River, NJ: Prentice-Hall, Inc. 599 p.

DeBano, L.; Neary, D.; Ffolliott, P. 1998. Fire’s effects on ecosystems. New

York, NY: John Wiley & Sons. 333 p.

Johnson, A. 2004. Nuttall Complex 2004: treatment effectiveness monitoring report. Unpublished paper on file at: U.S. Department of Agriculture,

Coronado National Forest, Tucson, AZ.

Moody,J.A.; Martin, D.A. 2009. Forest fire effects on geomorphic processes.

In: Cerdà, A.; Robichaud, P.R., eds. Fire effects on soils and restoration

strategies. Enfield, NH: Science Publishers. Pgs. 41-79.

Moody, J.A., and Martin, D.A., 2009. Synthesis of sediment yields after

wildland fire in different rainfall regimes in the western United States.

International Journal of Wildland Fire. 18(1): 96-115.

Pannkuk, C.D; P.R. Robichaud. 2003. Effectiveness of needlecast at reducing erosion after forest fires. Water Resources Research. 39: 1333-1344.

Robichaud, P.R. 2005. Measurement of post-fire hillslope erosion to evaluate

andmodelrehabilitation treatment effectiveness andrecovery.International

Journal of Wildland Fire. 14: 475-485.

Robichaud, P.R.; Ashmun, L.E.; Sims, B.D. 2010. Post-fire treatment effectiveness for hillslope stabilization. Gen Tech. Rep. RMRS-GTR-240.

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station. 62 p.

Scott,D.F.;Curran,M.P.;Robichaud,P.R.;Wagenbrenner,J.W. 2009.Soil erosion afterforest fire.In:Cerdà,A.;Robichaud, P.R., eds. Fire effects on soils

and restoration strategies. Enfield, NH: Science Publishers. Pgs. 178-195.

USDA,ForestService.2006.CoverFrequencyMethodologyTechGuide.Available online from http://fsweb.wo.fs.fed.us/rge/inventory/cover-frequency.

php [accessed 21. September, 2011]
336 USDA Forest Service Proceedings RMRS-P-67. 2013

Gibson and others Hillslope Treatment Effectiveness Monitoring on Horseshoe 2 and Monument Fires

USDA, Forest Service. 2011a. Burned Area Report, Horseshoe 2 Burned

Area Emergency Recovery Assessment. September 2 3, 2011 FS-2500-

08, Interim Request #2: U.S. Department of Agriculture, Forest Service,

Coronado National Forest, Douglas Ranger District. 28p.

USDA, Forest Service. 2011b. Burned Area Report, Monument Fire Burned

AreaEmergencyRecoveryAssessment.August3,2011FS-2500-08,Interim

Request #1: U.S. Department of Agriculture, Forest Service, Coronado

National Forest, Sierra Vista Ranger District. 22p.

USDA, Forest Service. 2011c. Horseshoe 2 & Monument Fires RapidAssessment of Vegetation Condition after Wildfire. Published data on file at: U.S.

Department of Agriculture, Forest Service, Remote Sensing Applications

Center, Salt Lake City, UT.

Wohlgemuth, P., J. Beyers, P. Robichaud. 2010. The effectiveness of aerial

hydromulch as a post-fire erosion control treatment in southern California.

Unpublished paper on file at: U.S. Department of Agriculture, Pacific

Southwest Research Station, Riverside, CA.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 337

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Ecology and Management of Oak Woodlands and 

Savannas in the Southwestern Borderlands Region

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona 

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—Management of the Madrean oak woodlands and the less dense and ecologically different oak 

savannas must be based on sound ecological information. However, relatively little is known about the 

Madrean oak ecosystems in spite of the fact that they cover about 80,000 km2 in the southwestern United 

States and northern Mexico. Emory oak (Quercus emoryi), the dominant tree in most stands, is usually associated with other oak and juniper species. Trees are utilized for firewood, fence posts, and construction 

materials and oak acorns are gathered as food by local inhabitants. The woodlands and savannas provide 

important habitat for listed and sensitive wildlife species and forage for livestock grazing. Tree regeneration 

and water consumption are important considerations. Tree harvesting has been restricted because of heavy 

utilization in the past but coppice management could ease the supply situation. Recreational demands and 

fire management concerns in the woodlands are increasing as human populations grow in southern Arizona. 

This manuscript reviews the ecology and management of the overstory component of oak woodlands and 

savannas based on relevant literature from the southwestern United States and northern Mexico. 

Introduction

	 TheMadrean woodlands and less dense savannas are characterized

by mild winters and wet summers. Brown (1982) classified them as

belonging to the warm-temperate forests and woodlands biotic community and lists them as Madrean Evergreen Woodlands. There is a

longhistoryofhumanoccupationinthewoodlandsbutmanagementof

the Madrean oak ecosystems has become more complicated in recent

years because of the increasing human populations in the region and

societal demands. The woodlands and savannas are noted for their

beauty and biological diversity, containing representative plants and

animalsfrom the Rocky Mountainsto the north and the Sierra Madre

of Mexico to the south. The ecosystems provide important wildlife

habitat for federally listed and sensitive and more common speciesin

the United States. The region supports viable ranching communities

and is a recreational destination for people from urban areas. Trees

are utilized for fuelwood, fence posts, charcoal, and construction

materials (Ffolliott and Gottfried 1992) and oak acorns and pinyon

seeds are gathered as food by local inhabitants. Woodland health and

local water supplies are important considerations.

	 The exclusion of wildfires since the late 1800s because of overgrazing and fire suppression activities has resulted in increases in

woody vegetation and fuels and a decline in herbaceous resources.

Large wildfires such as the Horseshoe 2 and Monument Fires in

2011 have become common. Private and public land managers are

attempting to reverse thistrend by using prescribed andmanaged fires

(Gottfried and others 2007a). Management must be based on sound

ecological information. However, relatively little is known about the

Madrean oak ecosystems in spite of the fact that they cover about

80,000 km2 in the southwestern United States and northern Mexico.

This presentation focuses mainly on the ecology and management

of the tree overstory component of Madrean woodland and savanna

ecosystems based on research findings and relevant literature from

the southwestern United States and northern Mexico.

Description of the Woodlands and 

Savannas

	 Emoryoak(Quercus emoryi)isthedominanttree inmoststands and

isusuallyassociatedwithotheroaks,junipers andconiferspecies.The

biological center for the ecosystemsisin the Sierra Madre of Mexico

but they extend northward into the SouthwesternBorderlandsRegion

and centralArizona (Brown 1982). The Madrean oak ecosystems are

representative of arid and semi-arid dryland forests and woodlands

of the world (McPherson 1992). The oak woodlands and savannas

reach their best development in Arizona on the foothills of higher

mountains, including the Pinaleños, Peloncillos, Santa Catalinas,

Huachucas, and Chiricahuas (Ffolliott and Gottfried 2008a) and are

found from 1219 to 1981 m in elevation. They grade into oak-pine

and pine forests at the higher elevations and grasslands or desertshrub

at the lower extreme. Temperatures range from summer highs in or

above 32 oC to freezing in the winter.Average annual precipitation is
338 USDA Forest Service Proceedings RMRS-P-67. 2013

Gottfried and Ffolliott Ecology and Management of Oak Woodlands and Savannas in the Southwestern Borderlands Region

between 254 to 508 mm, depending on elevation, with approximately

half occurring during the summer monsoon period.

	 The woodlands and savannas are characterized by evergreen oaks

generally 6 to 15 m in height. Emory oak is often associated with

Arizona white oak (Q. arizonica), and Mexican blue oak (Q. oblongifolia). Gray oak (Q. grisea), a generally small tree, might replace

white oak in western New Mexico. Silverleaf oak (Q. hypoleucoides)

and netleaf oak (Q. rugosa) are found at higher elevations in the

Madrean oak ecosystems. A relatively large number of additional

oak species including Chihuahua oak (Q. chihuahuensis) and cusi

or encino roble (Q. albocincta) occur in Mexico (Brown 1982). On

many sites, Emory oak is associated with conifers such as alligator

juniper (Juniperus deppeana), redberry juniper (J. coahuilensis),

border pinyon (Pinus discolor) or Mexican pinyon (P. cembroides).

Chihuahua pine (P. leiophylla) often occurs in drainages. A number

of pine species are found with oak at higher elevationsthroughout the

border region and into the oak-pine communities of Mexico (Brown

1982). The density of shrubs varies by sites. Some shrub species are

velvetpodmimosa(Mimosa dysocarpa),redbarberry(Mahonia haematocarpa), andalderleafmountainmahogany(Cercocarpus montanus).

Grasses include several species of grama (Boutelous spp.), bullgrass

(Muhlenbergia emersleyi), and Texas bluestem (Schizachyrium cirratum). More complete lists of grasses, forbs, shrubs and half-shrubs

are listed in Brown (1982) and Ffolliott and Gottfried (2008a).

Differences Between Woodlands and Savannas

	 Ffolliott and others (2008b) examined the differences between the

woodlandsin the SanRafael Valley on the south side ofthe Huachuca

Mountains in Arizona and at the lower elevation, more open oak

savannas at Cascabel on the east side of the Peloncillo Mountains in

New Mexico. The oak savannas were more open and variable in the

spatial distribution of overstory trees than the oak woodlands. These

researchers tallied seven tree species at Cascabel but only four at the

San Rafael site. Emory oak accounted for 60% of the overstory trees

at Cascabel and junipers 15%. On the San Rafael site, Emory oak

was 89% of the overstory species composition and junipers were 1%.

Arizona white oak made up 12% of the overstory at Cascabel and 9%

at San Rafael. There were about 445 trees/ha at San Rafael compared

to 148 trees/ha at Cascabel. The number of small and large trees was

greater at San Rafael but the number of saplings was similar for the

two sites. Herbaceous plant species composition was similar on the

two areas with most species of grasses and forbs being perennial.

Autecology

	 Oak trees regenerate from seed or vegetatively from root or stump

sprouts. Borelli and others (1994) studied natural regeneration on

six areas of the Coronado National Forest of Arizona that had been

harvested for fuelwood and grazed by livestock. Emory oak,Arizona

white oak and other tree species occurred on the study sites. Fiftysix percent of the regeneration was stump sprouts, 25% were root

sprouts, and 19% were seedlings. Nearly 40% of the plantlets were

less than 0.3 m in height, 35% were between 0.3 and 1.0 m, and 25%

were greater than 1 m but less than 1.4 m. An average of 9.2 % of

the sample plots was stocked with regeneration varying from 1.2 to

15.4% depending on area. The average number of plantlets on the six

areas was 240 stems/ha with a range of 29.7 to 385.5 plantlets/ha.The

authors caution thatthe relatively lownumber ofseedlings could have

been influenced by a local drought that could have influenced acorn

production. Pase (1969)reported that conditionsfavorable for Emory

oak establishment only occurred once every 10 years in southeastern

Arizona. However, large numbers of acorns probably are consumed

by mammals and insects. The study by Borelli and others(1994) was

conducted in one year; however, surveys over a number of years are

necessary to get a true picture of regeneration in the oak woodlands.

	 Nyandiga and McPherson (1992) evaluated conditions needed for

germination of Emory oak andArizona white oak. Emory oak acorns

mature in about a year and then fall and germinate in July through

August. Arizona white oak acorns fall and germinate in November

and December. Germination generally occurred within the first 30

days for both species. These researchers found buried acorns had a

greater viability than acornsthat remained on the soilsurface or were

under litter. However, seed viability generally declined the longer an

acorn was buried. Acorn viability and germination was twice as high

undertree canopies asin open grassland sites.Adequate soil moisture

will enhancegerminationandemergence (Germaine andothers1996).

Germination percentages were 21 and 73% for Emory and white oak,

respectively. Emory oak acorn size is correlated positively with viability, germination, and seedling size (Germaine and others 1996).

	 Sanchini (1981) studied growth and mortality of Emory oak and

white oak in 1981 (cited in McPherson 1992; Touchan and Ffolliott

1999). He determined that mean annual diameter growth rate for

Emory oak was 0.35 cm/yr and forArizona white oak was 0.24 cm/yr.

Growth-ring data, while difficult to measure in oak species, indicated

that Emory oak reaches a maximum age at 200 years while white oak

achievesthisstatusin 250 years. The primary causes of death are fire,

ice storms, and drought.

	 Ffolliottandothers(2008b),ininvestigatingthedifferencesbetween

woodlands andsavannas confirmedthattreevolumesperhectarewere

higherin the denser woodlandsthan in the savannas.Volumes ofstem

wood varied from less than 2 to more than 100 m3/ha. The average

volumes for the two sites were about 15.86 m3/ha compared to 3.91

m3/ha. Annual volume growth rates in the dense stand at San Rafael

were 0.0077 ± 0.0010 m3/ha while they were 0.0049 ± 0.0015 m3/

ha at Cascabel. Growth was relatively rapid in the early and middle

stages of development but declined in older trees.

Fire

	 Fire was the main disturbance in the Madrean oak woodlands and

savannaspriortoEuropeansettlementinthe late1800s.Firesprobably

occurred every 10-20 years at the lower end of the oak woodlands.

These values were extrapolated from literature about forest fires in

standsthat are adjacent to semi-desert grasslands(McPherson 1992).

Fire histories are difficult to determine for the oak woodlands since

fire scars are not evident; however, conifers in the affected stands do

display fire scars. Fire frequencies and extent have declined in the

last century. The importance of fire for ecosystem health isjust being

recognized (Gottfried and others 2007a). Two prescribed fires and a

wildfire in the savanna at Cascabel reduced overstory tree density of

oaks and junipers by 21% (Ffolliott and others 2011) but resulted in

significant increasesin early-growing and late-growing grassesin the

understory (Ffolliott and others 2012). Basal sprouting at Cascabel

was observed on 37.4 % of the surviving oaks regardless of species.

A wildfire in the Catalina Mountains, north of Tucson, resulted in

11.1% mortality of Emory oak and 14.2% mortality of Mexican blue

oak. Emory oak produced more sprouts per tree after the fire than did

Mexican blue oak (Caprio and Zwolinski 1992).

	 An inventory in six siteswhereEmory oak dominated foundwoody

fuel loadings ranging from 0.36 to 7.82 tons/ha (Ottmar and others

2007). Measurements of litter and duff ranged from 3.26 to 10.72

tons/ha with soil surface cover of 40 to 82%. A study in pine-oak
USDA Forest Service Proceedings RMRS-P-67. 2013 339

Ecology and Management of Oak Woodlands and Savannas in the Southwestern Borderlands Region Gottfried and Ffolliott

woodlands in Arizona and northern Mexico found that sites with

high fire frequencies accumulated less fuels than sites with low fire

occurrences (Escobedo and others 2001). Sites in Mexico, which

had 9 to 13 fires since 1900, had total fuel loadings of 11.22 tons/ha

while sites in Arizona, that had from 0 to 5 fires during this period

had total fuel loadings of between 22.19 and 48.43 tons/ha. Higher

fuel loadingsincrease the potential for high-intensity stand-replacing

fires.

Silviculture

	 The oakwoodlands, and to a lesser extentsavannas,were harvested

heavily during the period of European settlement for mining and

construction timbers, fuel for the smelters and stamp mills, fence

posts, and for domestic uses by the miners and their families. The

overharvesting did not eliminate the oak woodlands but resulted in

a decrease in large trees and stand densities (Propper 1992). Interest

in oak fuelwood increased in the 1970s and 1980s because of fuel

shortages and increased oil prices. Federal lands provided 60% of the

fuelwood in southeastern Arizona. Managers were forced to restrict

harvesting during this period to guarantee sustainable fuelwood resources (Bennett 1992). Harvesting was limited to 3.62 m3 (1 cord)/

household/year in 1992.

	 Fuelwood harvesters were further restricted to leaving stumpsless

than 15-cm in height and slash accumulations of less than 46-cm in

height (Bennett 1992). Trees designated for harvesting were marked

by managers in what appeared to be either single-tree selection or

thinning from below depending on stand conditions and objectives.

On some Forest Service districts, fuelwood harvesting was limited

to trees that were smaller than 23 cm at the base. Larger trees were

maintained because of their importance to wildlife.

	 Vigorous sprouting of oak species after the removal of the tree

indicates that coppicing might form a basis to obtain satisfactory

regeneration since stump and root sprouts are more common than

seedlings(Borelliandothers1994).Furthermore,TouchanandFfolliott

(1999) determined that the fuelwood rotation for Emory oak sprouts

to reach a diameter at root collar of 15 to 20 cm can be reduced from

100 years to approximately 30 years by thinning sprouts on a stump.

Growth of residualsprouts depends on the number ofstemsretained.

Net growth/stump is lower if fewer stems are left. While there was

no difference if 1, 2, or 3 stems remained compared to the control,

which was not thinned, mean annual growth per sprout increased for

the single stem after its eighth anniversary.

	 Farah and others (2003) found that stump diameter was not a consistent factor that affected growth and volume of the thinned coppice

10 years after the thinning treatments were applied. Average growth

and volume was greater for the 1-stump sprout treatment than for the

other options at that time. The researchers recommended that stump

sprouts be thinned initially in years 6-8 after the top is removed.

Another subsequent thinning in year 15-20 should retain one sprout.

Three stump-sprouts can be retained if structural diversity to benefit

wildlife is a management goal.

Water Use

	 Water use by Emory oak has been studied with the heat pulse velocity method in the San Rafael Valley (Ffolliott and Gottfried 2000;

Gottfried and others 2007b; Shipek and others 2004). The study site

included an unharvested stand and an area harvested by the coppice

method that included sprouts and residual uncut trees (standards).

Average daily water use by mature trees averaged 17.5 liters/day,

while water use by sprouts averaged 4.0 liters/day. These averages

forstandards and sprouts were then extrapolated to a stand basis. The

harvested stand used more than 3,150 liters/ha/yr compared to 1,900

liters/ha/yr for the uncut stand (Ffolliott and Gottfried 2000). The

harvested stand contained 430 standards and 927 sprouts per hectare,

while the uncut area contained 452 mature trees and 27 sprouts per

hectare. Water use by the harvested site with the large number of

sprouts was equivalent to 80% and water use by the unharvested site

was equivalent to 45% of the annual precipitation.

	 Shipek and others (2004) measured water use by oak sprouts that

had been thinned to retain 1, 2, or 3 sprouts per stump or not thinned.

Stumps with one sprout used an average of 7.48 ± 0.31 liters/day

compared to the controls that used 35.3 ± 1.72 liters/day. The values

for stumps thinned to two or three stems were intermediate. Transpiration for stands with standards and sprouts thinned to 1-sprout per

stump and stands with unthinned stumps was equivalent to 34% and

80% of the annual precipitation, respectively.

	 Ffolliott (2004) developed an annual water balance for Emory

oak in southeastern Arizona. The calculations assumed that annual

precipitation averaged 450 mm. This amount was reduced by 45 mm

for interception by tree crowns; the remaining water was partitioned

mainly to infiltration into the soil (325-350 mm), and transpiration

(155-345 mm). Approximately 20-45 mm, about 4 to 10% of the

input, remained for streamflow. Thirty-four to 77% of the annual

precipitation was lost through transpiration.

Management Implicaitons 

	 Management ofthe Madrean oak woodlands and savannas must be

based on sound ecological information. Managers must understand

tree regeneration requirements to develop silvicultural prescriptions

that will maintain healthy oak woodlands and savanna ecosystems.

Vegetative reproduction by oaks and some junipers gives managers

optionsthat are not available to thoseworkingwith non-sprouting tree

species.Thinning oak sprouts willshorten the cycle for tree products.

Public and private land managers are attempting to reintroduce fire

into the region but these efforts also are often hindered by incomplete

ecological information about fire effects and fuel conditions. These

data are currently being collected on the Cascabel Watersheds and

elsewhere to support the fire programs. Research on tree and woodland water use and the impacts of silvicultural treatments such as

coppicing, should provide a basis for management prescriptions in

oak dominated watersheds.

References

Bennett,D.A.1992.FuelwoodextractioninsouthernArizona.In:Ffolliott,P.F.;

Gottfried, G.J.; [and others], tech. cords. Ecology and management of oak

and associated woodlands; 1992 April 27-30; Sierra Vista, AZ. Gen. Tech.

Rep. RM-218. Fort Collins, CO: U.S. Department of Agriculture. Forest

Service, Rocky Mountain Forest and Range Experiment Station: 96-97.

Borelli, S.; Ffolliott, P.F.; Gottfried, G.J. 1994. Naturalregeneration in encinal

woodlands of southeasternArizona. Southwestern Naturalist. 39:179-183.

Brown, D.E. 1982. 123.3 Madrean Evergreen woodland.In:Brown, D.E., ed.,

Biotic communities of theAmerican Southwest-United States and Mexico.

Desert Plants. 4:59-65.

Caprio, A.C.; Zwolinski, M.J. 1992. Fire effects on Emory and Mexican blue

oaksin southeasternArizona.In:Ffolliott,P.F.;Gottfried,G.J.;[and others],

tech. cords. Ecology and management of oak and associated woodlands;

1992April 27-30; Sierra Vista,AZ. Gen. Tech. Rep. RM-218. Fort Collins,

CO: U.S. Department of Agriculture. Forest Service, Rocky Mountain

Forest and Range Experiment Station: 150-154.
340 USDA Forest Service Proceedings RMRS-P-67. 2013

Gottfried and Ffolliott Ecology and Management of Oak Woodlands and Savannas in the Southwestern Borderlands Region

Escobedo, F.J.; Ffolliott, P.F.; Gottfried, G.J.; Garza, F. 2001. Fire frequency

effects on fuel loadings in pine-oak forests of the Madrean Province. Res.

Note RMRS-RN-10. Fort Collins, CO: U.S. Department of Agriculture.

Forest Service, Rocky Mountain Research Station. 7 p. 

Farah, M. H.; Ffolliott, P. F.; Gottfried, G. J. 2003. Growth and volume of

Emory oak coppice 10 years after thinning: A case study in Southeastern

Arizona. Western Journal of Applied Forestry. 18:77-80.

Ffolliott, P.F. 2004.Awater budget for Emory oak woodlands ofsoutheastern

Arizona:An expression of the initial approximation. Hydrology and Water

Resources in Arizona and the Southwest. 34:11-14.

Ffolliott,P.F.;Gottfried,G.J. 1992.Growth, yield, and utilization of oakwoodlands in the Southwestern United States. In: Ffolliott, P.F.; Gottfried, G.J.;

[and others], tech. cords. Ecology and management of oak and associated

woodlands; 1992 April 27-30; Sierra Vista, AZ. Gen. Tech. Rep. RM-218.

Fort Collins, CO: U.S. Department of Agriculture. Forest Service, Rocky

Mountain Forest and Range Experiment Station: 34-38.

Ffolliott,P.F.;Gottfried,G.J.2000.ChangesintranspirationbyEmoryoakfollowingtreeharvesting.In:Flug,M.;Frevert,D.,eds.,Watershedmanagement

and operations management 2000: proceedings of the conference. Reston,

VA: American Society of Civil Engineering. [CD ROM] Windows. 7 p.

Ffolliott, P.F.; Gottfried, G.J. 2008a. Plant communities and associations. In:

Ffolliott, P.F.; Davis, O.K., eds., Natural Environment of Arizona: from

deserts to mountains, University of Arizona Press, Tucson: 70-119.

Ffolliott, P.F.; Gottfried, G.J.; Stropki, C.L. 2008b. Vegetative characteristics

and relationships in the oak savannas of the Southwestern Borderlands.

Res.Pap.RMRS-RP-74.FortCollins,CO:U.S.Department ofAgriculture.

Forest Service, Rocky Mountain Research Station. 16 p.

Ffolliott, P.F.; Gottfried, G.J.; Chen, H.; Stropki, C.L.; Neary, D.G. 2012.

Fire effects on herbaceous plants and shrubs in the oak savannas of the

Southwestern Borderlands. Res. Pap. RMRS-RP-95. Fort Collins, CO:

U.S. Department ofAgriculture. Forest Service,Rocky MountainResearch

Station. 15 p.

Ffolliott, P.F.; Gottfried, G.J.; Stropki, C.L.; [and others]. 2011. Fire effects

on tree overstories in the oak savannas of the Southwestern Borderlands

Region. Res. Pap. RMRS-RP-86. Fort Collins, CO: U.S. Department of

Agriculture. Forest Service, Rocky Mountain Research Station. 13 p.

Germaine, H.L.; McPherson, G.R.; Rojahn, K.J.; Nicholas, A.M.; Weltzin,

J.F. 1997. Constraints on germination and emergence of Emory oak. In:

Symposium on oak woodlands: Ecology, management, and urban interface

issues;1996 March 19-22; San Luis Obispo, CA. Gen. Tech. Rep. PSWGTR-160. San Francisco, CA: U.S. Department of Agriculture. Forest

Service, Pacific Southwest Research Station: 225-230.

Gottfried,G.J.;Allen,L.S.;Warren,P.L.;McDonald,B.;Bemis,R.J.;Edminster,

C.B. 2007a. Private-public collaboration to reintroduce fire into changing

ecosystemsoftheSouthwesternBorderlandsRegion.FireEcology.5:85-99.

Gottfried, G.J.; Ffolliott, P.F.; Neary, D.G. 2007b. Hydrology ofsouthwestern

encinaloakecosystems:Areviewandmore.HydrologyandWaterResources

in Arizona and the Southwest. 37:19-30.

McPherson,G.R.1992.EcologyofoakwoodlandsinArizona.In:Ffolliott,P.F.;

Gottfried, G.J.; [and others], tech. cords. Ecology and management of oak

and associated woodlands; 1992 April 27-30; Sierra Vista, AZ. Gen. Tech.

Rep. RM-218. Fort Collins, CO: U.S. Department of Agriculture. Forest

Service, Rocky Mountain Forest and Range Experiment Station: 24-33.

Nyandiga,C.O.;McPherson, G.R. 1992. Germination oftwo warm-temperate

oaks Quercus emoryi and Quercus arizonica. Canadian Journal of Forest

Resources. 22:1395-1401.

Ottmar,R.D.;Vihnanek,R.E.;Wright,C.S.;Seymour,G.B.2007.Stereophoto

series for quantifying natural fuels. Vol. IX: Oak/juniper types in southern

Arizona and New Mexico. Gen. Tech. Rep. PNW-GTR-714. Portland,

OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest

Research Station. 41 p. 

Pase, C.P. 1969. Survival of Quercus turbinella and Q. emoryi in an Arizona

chaparral community. Southwestern Naturalist. 14:149-156.

Propper,J.G.1992.Cultural,recreational, andestheticvaluesofoakwoodlands.

In: Ffolliott, P. F.; Gottfried, G. J.; [and others], tech. cords. Ecology and

management of oak and associated woodlands.; 1992 April 27-30; Sierra

Vista, AZ. Gen. Tech. Rep. RM-218. Fort Collins, CO: U.S. Department

of Agriculture. Forest Service, Rocky Mountain Forest and Range Experiment Station: 98-102.

Sanchini, P.J. 1981. Population structure and fecundity patterns of Quercus 

emoryi and Q. arizonica in southeastern Arizona. Ph.D. dissertation, University of Colorado, Boulder. 149 p.

Shipek, D.C.; Ffolliott, P.F.; Gottfried, G.J.; DeBano, L.F. 2004.Transpiration

and multiple use management of thinned Emory oak coppice. Res. Pap.

RMRS-RP-48. Fort Collins, CO: U.S. Department of Agriculture. Forest

Service, Rocky Mountain Research Station. 8 p.

Touchan R.; Ffolliott, P.F. 1999. Thinning of Emory oak coppice: Effects

of growth, yield, and harvesting cycles. Southwestern Naturalist. 44:1-5.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 341

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Effects of Prescribed Fires and a Wildfire on Biological 

Resources of Oak Savannas in the Peloncillo Mountains, 

New Mexico

Gerald J. Gottfried

USDA Forest Service, Rocky Mountain Research Station, Phoenix, Arizona

Peter F. Ffolliott

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Daniel G. Neary

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Abstract—Private and public land managers are attempting to reintroduce fire into the ecosystems of the 

Peloncillo Mountains to reduce the density of woody species, increase the herbaceous plant cover, and 

improve the area’s ecological diversity. The Cascabel Watershed Study, which includes 12 small gauged 

watersheds, was started in 2000 to evaluate the impacts of cool-season (November-April) and warmseason (May-October) prescribed fires on biological and physical components of the oak (Quercus spp.) 

savannas that are common to the Southwestern Borderlands Region. The Whitmire Wildfire in May 2008 

modified the original experimental design. All three fire treatments resulted in low severity burns. The effects 

of the different fire treatments on tree overstories, herbaceous production, ground cover, selected wildlife 

species, and bird populations are presented. Measurements did not indicate many significant differences 

among treatments or with pre-fire conditions. For example, almost 78 percent of the overstory trees on the 

watersheds survived the fires. The main exception was a large increase in grass and total herbaceous plant 

production of early-growing species after the fires; annual grasses and forbs are not common at Cascabel. 

The fires also produced statistically significant increases in the production of late-growing grass species but 

the changes were not as large. 

Introduction

Fires caused by lightning or Native American people were the main 

disturbance in the Madrean Archipelago prior to the completion of the 

transcontinental railroad and the introduction of large herds of cattle 

into the region. The range was overstocked with cattle as ranchers, 

often belonging to large corporations, competed with their neighbors 

to control access to the land (Hadley 2005). The resulting overgrazing caused a decline in the herbaceous cover that previously carried 

natural fires throughout the landscape. Consequently, the density of 

trees and woody species and debris increased and the production of 

herbaceous species declined. Biodiversity and productivity of many 

sites within the Madrean Archipelago decreased. Aggressive fire 

suppression by land management agencies also contributed to the 

decline in fires.

Ranchers in the Upper San Simon, San Bernardino, and Animas 

Valleys became concerned about landscape changes that were resulting 

from the lack of fire. The Malpai Borderlands Group was formed in 

1992 by ranchers and similar minded individuals and organizations 

with the goal to “restore and maintain natural processes that create a 

healthy landscape” (McDonald 1995:485). Officials of the Coronado 

National Forest and Bureau of Land Management shared a similar 

view about the need to reintroduce prescribed and natural fires into 

the region. The Coronado National Forest and Malpai Borderlands 

Group worked together to formulate the Peloncillo Programmatic Fire 

Plan to reintroduce fire into the Peloncillo Mountains that straddle the 

Arizona-New Mexico border. The efforts resulted in four landscape 

prescribed fires between 1995 and 2007. The fires occurred in the 

warm season prior to the monsoon rains. However, there were questions about the effects of warm-season fires on several listed species, 

particularly the lesser long-nosed bat (Leptonyceris curasoae), which 

requires nectar and pollen from the Palmer agave (Agave palmeri), 

and the New Mexico ridge-nosed rattlesnake (Crotalus willardi obscurus). Some people recommended cool-season burns that might 

be less detrimental. However, there was little scientific information 

available to support either position on the burning season or to assist 

managers with general knowledge about the ecology and hydrology 

of the ecosystem. 

The U.S. Forest Service, Rocky Mountain Research Station established the Southwestern Borderlands Ecosystem Management Unit to 

provide public and private land managers with scientific information 

to support management decisions. The Cascabel Watershed Study was 

a research project specifically developed to answer questions about 
342 USDA Forest Service Proceedings RMRS-P-67. 2013

Gottfried and others Effects of Prescribed Fires and a Wildfire on Biological Resources of Oak Savannas . . .

the impacts of burning season on oak savanna ecosystem resources 

(Gottfried and others 2007, 2012). The Cascabel Study can be divided 

into research about the biological components of the oak savanna 

ecosystem and research about the physical environment, for example, 

hydrology, erosion, and sedimentation. This manuscript will report 

on the effects of fires on the biological resources.

Cascabel Watershed Study

The objective of the study was to determine the impacts of coolseason (November-April) and warm-season (May-October) prescribed 

burning on oak savanna hydrologic and biological resources (Gottfried 

and others 2007, 2012). The Cascabel Watersheds consist of 12 small 

watersheds located in the oak savannas on the southeast side of the 

Peloncillo Mountains. The watersheds range in size from 20 to 60 

acres covering about 451.3 acres and are located between 5380 and 

5590 feet in elevation. Geology, physiology, and hydrology of the 

study area have been described in articles cited by Ffolliott and others 

(2011b) and Gottfried and others (2012). The surface soils are very 

gravelly or very cobbly sand or sandy clay loams (Robertson and others 2002). Emory oak (Quercus emoryi) is the dominant tree species 

with alligator juniper (Juniperus deppeana) a common co-dominant 

species. Various species of grama grass (Bouteloua spp.) dominate the 

understory. Sets of two Parshall flumes were installed in each drainage as were sediment dams, basins, and channel cross sections. Two 

full weather stations were set up and these are supported by seven 

tipping bucket rain gauges. Measurements of fire effects, soil erosion 

and deposition, overstory trees, herbaceous vegetation, and wildlife 

use were collected in each watershed at permanent sampling points 

arranged on transects that were located perpendicular to the stream 

channel. There were 35 to 45 points on each watershed depending 

on its size and shape for a total of 421 sampling points. Sampling 

procedures varied depending on the resource being monitored.

Burning Treatments and Fire Effects

After a pretreatment sampling period, the watersheds were partitioned into three groups of four with one group receiving the coolseason prescribed burn, one receiving the warm-season prescribed 

burn, and one group reserved as controls. The cool-season watersheds 

were burned by crews from the Douglas Ranger District of the Coronado National Forest in March 2008 and three of the warm-season 

watersheds were burned on May 20, 2008. However, the fire escaped 

on May 21 and burned the remaining unburned watershed, the control 

watersheds, and about 4,000 acres in adjacent areas. It was named 

the Whitmire Fire. The study design was modified appropriately to 

account for the wildfire treatment on five of the watersheds.

 Fire severities were determined on the 421 points using a system 

developed by Hungerford (1996) that relates fire severity to the appearance of litter, duff, woody material, and soil condition. Classification 

of severity ratings at the points on each watershed were extrapolated 

to a watershed basis by Stropki and others (2009). They determined 

that 85% of the area receiving the cool-season burn had been exposed 

to a low severity fire, 5% had been exposed to a moderate severity fire 

and the remaining 10% of the area had been unburned. The spatial 

distribution of severities was similar on the watersheds that had been 

affected by the warm-season burn and the wildfire. However, Neary 

and others (2010) measured locations on the watersheds where highseverity fires had occurred, particularly where heavy accumulations 

of organic material had accumulated. Statistical analyses indicated 

no significant differences in burn severities among the watersheds 

receiving the cool-season or warm-season prescribed burns or the 

wildfire. Therefore, the data were combined to compare pre-treatment 

and post-treatment responses of the resources.

 Soil water repellency was measured using the drip test developed 

by Letey and others (2000). The occurrence and levels of soil water 

repellency was limited in extent and magnitude regardless of treatment. Approximately 90% of the area had no soil water repellency, 

5% had slight water repellency, and the remaining points had moderate 

to strong levels of repellency. 

Fire Effects On Biological Resources

Overstory Trees

The overstory tree cover was re-measured after the three fire treatments and the results were pooled because all watersheds had received 

low severity burns and no significant differences were detectable 

(Ffolliott and others 2011b). Approximately 78% of the trees that 

were present before the prescribed fires and wildfire initially survived. 

Eighty percent of the oak trees of all species and 75% of the juniper 

trees of all species survived. Scorch or top kill, was observed on 80% 

of the surviving oak and juniper trees. Thirty percent of the oak had 

one-third or less of the crown burned, 45% had between one-third and 

two-thirds affected and 25% had greater than two-thirds damaged. The 

relative proportions were similar for the junipers. Many of the severely 

burned trees might not survive into the future but basal sprouting could 

replace some of the dying trees. Sprouting was observed on 37% of 

the oaks and on 11% of the junipers. Large trees (≥ than 9 inches in 

diameter at the root collar) were more common on the watersheds 

than smaller trees and proportionally more large trees survived. The 

damage to large trees could be attributed to accumulations of slash 

and other organic material at their bases. The spatial distribution of 

trees at Cascabel was not impacted by the burning treatments.

Herbaceous Plant and Shrub Production

The production of early-growing and late-growing grasses was 

significantly greater after the three fires than before (Ffolliott and 

other 2012b). Depending on the event, there was a five to seven-fold 

increase in the production of early-growing grass species. Increases in 

late growing species were also significant but of a smaller magnitude. 

For example, the early growing grass production on the warm-season 

burned watersheds increased from 75 pounds/acre to 400 pounds/

acre. Fire effects on the minor forb component were inconsistent. The 

cool-season prescribed burn did not affect forb production, but the 

warm-season burn and the wildfire did result in increases. Late-season 

forbs were not affected by the fires. Total herbaceous production, 

which includes grasses and forbs, increased relative to pre-treatment 

condition because of the dominance of grasses in the understory. The 

increased production has been attributed to the increased soil water 

availability related to reductions in tree overstory and litter and duff 

layers and to increased soil nutrients. The growth of shrubs generally 

was not affected by the burning events, except for the late-season 

growth of shrubs on the watersheds impacted by the wildfire.

 The wildfire did produce an increase in warm season shrub production mainly because of tree sprouts. The persistence of these effects 

into the future is unknown.
USDA Forest Service Proceedings RMRS-P-67. 2013 343

Effects of Prescribed Fires and a Wildfire on Biological Resources of Oak Savannas . . . Gottfried and others

Palmer Agave

Palmer agave, which is protected by the State of Arizona, is important because it provides pollen and nectar to the Federally listed lesser 

long-nosed bats that migrate into the southwestern United States in 

the summer (Slauson and others 1999). The agave species occurs on 

rocky slopes in the oak savannas of the Southwestern Borderlands 

region. Concern about damage to this species was one reason for 

the Cascabel study. Information about the agave population on the 

watersheds was not collected prior to the burning events but a survey 

was conducted on the plots afterwards (Ffolliott and others 2010). 

The agave was found on 12.9% of the plots visited after the fires. The 

occurrence of agave could not be related to topography, soil surface 

characteristics, or to surrounding vegetation. No burned agave plants 

were found on the sample plots but an unknown number of burned 

or partially burned plants were observed outside the established 

plots. There was no pattern in the location of dead or injured agaves. 

Average agave production on the watersheds following the fires was 

about 0.25 pounds/acre with a range of a trace to 16.7 pounds/acre on 

individual watersheds. Ffolliott and others (2010) could not conclude 

if the Cascabel treatments had an effect on the Palmer agave population because of the lack of pretreatment data.

Wildlife

Mammals—The prescribed burns and the wildfire did not affect the 

use of the watersheds by two keystone species—Coues white-tailed 

deer (Odocoileus virginianus couesi) and desert cottontail (Sylvilagus 

audubonii) (Ffolliott and others 2012a). Fecal pellets were counted 

in the spring and autumn at the permanent sampling points. The area 

around each point was cleared after each survey. There were no differences in deer or cottontail use related to the three low-severity fires. 

However, counts in the spring after the winter season were higher 

than counts in the autumn after the monsoon rains. It appears that the 

deer use the savannas during the winter when weather conditions are 

relatively mild and move into the higher, cooler, and moister elevation 

woodlands and forests during the warm summer. The same pattern of 

higher spring use by the cottontail was observed. It was hypothesized 

that this could be related to high birth rates during the winter and 

subsequent predation during the summer.

Birds—Surveys of bird species and numbers were conducted around 

a subset of sampling points on each watershed during the spring 

and autumn before and after the fires (Ffolliott and others 2011a). 

Bird observations were made for 5-min at each point following the 

procedures described by Braun (2005). Some species were observed 

frequently while others were tallied occasionally. More bird numbers 

and species were tallied in the autumn after the monsoon rains than 

in the spring after the winter rains. This difference could be related 

to greater availability of food following the monsoon period. Some 

common sightings in both seasons included the Ash-throated Flycatcher 

(Mylarchus cinerascens), Bushtit (Psaltriparus minimus), Mexican 

Jay (Aphelocoma ultarmarina), Turkey Vulture (Cathartes aura), and 

Mourning Dove (Zenaida macroura). A more complete list of species is presented in Ffolliott and others (2011a). Calculations of bird 

species richness, species diversities, and evenness for both seasons 

suggest that the three fire treatments had little consistent effect on the 

ecological diversity at Cascabel. It was difficult to isolate the impact 

of the prescribed burns or the wildfire because of the large variability 

in tallies of species and numbers. 

Herpetofauna—No New Mexico ridge-nosed rattlesnakes were 

encountered on the Cascabel Watersheds. However, 8,951lizards 

belonging to 10 species were identified at Cascabel between 2004 

and 2010 (Goode 2011, Unpublished report). It is difficult to draw 

conclusions about the effects of fire on the lizard population because 

of the variability of rainfall, vegetative cover, and lizard numbers on 

the plots. A preliminary conclusion was that the fires did not affect 

the lizard population.

Conclusions

The Cascabel Watershed Study was established to determine the 

impacts of cool-season and warm-season prescribed fires on the ecology and hydrology of Madrean oak savannas. The effect of burning 

season on natural resources was an important concern among public 

and private land managers. The Whitmire Wildfire in 2008 modified 

the statistical design so that the current analyses evaluate the impacts 

of cool season and warm season prescribed fires and a wildfire on the 

biological and hydrologic resources. This paper discussed the impacts 

on the biological resources at Cascabel. The physical aspects of the 

study are reported elsewhere (Gottfried and others 2012).

The three fire treatments, regardless of prescription or timing, 

burned at low severity and produced similar impacts on the biological 

resources. The data from the treatments were combined and subsequent 

analyses compared pre-treatment with post-treatment conditions. The 

reduction in tree survival was minor and inconsequential in terms of 

management options (Ffolliott and others 2011b). The production 

of grasses and general herbaceous cover was the only resource that 

exhibited a significant change related to the fires and that was a positive increase in production (Ffolliott and other 2012b). The fires did 

not affect deer or cottontail use of the total area, probably because of 

a lack of significant changes to the tree cover. The lizard population 

probably was not affected by the fires but that needs further analyses, 

and no fire effects could be determined from the bird surveys. 

It is concluded that the low-severity fires at Cascabel did not have 

major impacts on most biological resources of the oak savannas. Fire 

managers might use the Cascabel Watershed results as an initial guide 

since there is little information available on fire effects in the region’s 

oak savannas, but realize that different pre-fire conditions or weather 

could produce other results (Ffolliott and others 2011b). Prescribed 

fires ignited under warmer and drier conditions or in denser stands 

of trees and brush could burn more severely and have more drastic 

impacts. Managers will have to consider all options as they develop 

and execute programmatic fire plans.

References

Braun, C.E., ed. 2005. Techniques for wildlife investigations and management. 

The Wildlife Society, Bethesda, MD.

Ffolliott, Peter F.; Chen, Hui; Gottfried, Gerald J. 2011a. Bird species and 

numbers of birds in oak savannas of the Southwestern Borderlands Region 

including effects of burning. Journal of the Arizona-Nevada Academy of 

Science. 42:75-83.

Ffolliott, Peter F.; Gottfried, Gerald J.; Woods, Steven R. 2010. Occurrence 

and production of agave on the Cascabel Watersheds following three burning events. Hydrology and Water resources in Arizona and the Southwest. 

40:7-10.

Ffolliott, Peter F.; Chen, Hui; Gottfried, Gerald J.; Stropki, Cody L. 2012a. 

Coues white-tailed deer and desert cottontail in the southwestern savannas: 

Their presence before and after burning events. Journal of the ArizonaNevada Academy of Science. 44(1): 1-5.

Ffolliott, Peter F.; Gottfried, Gerald J.; Chen, Hui; Stropki, Cody L.; Neary, 

Daniel G. 2012b. Fire effects on herbaceous plants and shrubs in the oak 

savannas of the Southwestern Borderlands. Res. Pap. RMRS-RP-95. 
344 USDA Forest Service Proceedings RMRS-P-67. 2013

Gottfried and others Effects of Prescribed Fires and a Wildfire on Biological Resources of Oak Savannas . . .

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 15 p.

Ffolliott, Peter F.; Gottfried, Gerald J.; Stropki, Cody L.; Chen, Hui; Neary, 

Daniel G. 2011b. Fire effects on tree overstories in the oak savannas of the 

Southwestern Borderlands Region. Res. Pap. RMRS-RP-86. Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

research Station.13 p.

Goode, Matt. 2011. Effects of fire and grazing on Malpai Borderlands herpetofauna: A multifaceted research program. Annual Report-2010. Unpublished 

report on file at: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station, Phoenix, AZ. 7 p.

Gottfried, Gerald J.; Neary, Daniel G.; Ffolliott, Peter F. 2007. An ecosystem approach to determining effects of prescribed fire on southwestern 

Borderlands oak savannas: a baseline study. In: Proceedings of the 23rd

Tall Timbers Fire Ecology Conference. Tall Timbers Research Station, 

Tallahassee, FL: 140-146.

Gottfried, Gerald; Neary, Daniel; Ffolliott, Peter; Koestner, Karen. 2012. 

Cascabel prescribed fire long-term watershed study: an opportunity to 

monitor climate change. Proceedings of a workshop held during the XXV 

IUGG General Assembly in Melbourne, Australia, June-July 2011. IAHS 

Pub. 353. 10 p.

Hadley, Diana. 2005. Landholding systems and resource management in the 

Sky Island Borderlands. In: Gottfried, G.J.; Gebow, B.S,; Eskew, L.G.; 

Edminster, C.B., comps. Connecting Mountain islands and Desert Seas: 

biodiversity and Management of the Madrean Archipelago II. Proceedings 

RMRS-P-36. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station:15-25. 

Hungerford, R.D. 1996. Soils: fire in ecosystem notes: Unit II-I. Marana, 

AZ; U.S. Department of Agriculture, Forest Service, National Advanced 

Resource Technology Center.

Letey, J.; Carrillo, M.L.K.; Pang, X.P. 2000. Approaches to characterize the 

degree of water repellency. Journal of Hydrology 231:61-65.

McDonald, Bill. 1995. The formation and history of the Malpai Borderlands 

Group. In: DeBano, L.F.; Ffolliott, P.F.; Ortega-Rubio, A.; [and others]. 

comps. Biodiversity and management of the Madrean Archipelago: The 

sky islands of the southwestern United States and northwestern Mexico. 

Gen. Tech. Rep. RM-GTR-50. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment 

Station: 483-486. 

Neary, Daniel G.; Gottfried, Gerald J.; Koestner, Karen A.; [and others]. 

2010. Burning temperatures and fire severity in cool and warm season 

prescribed fires and a wildfire in an oak savanna in the southwestern USA. 

Proceedings of the VI International Conference on Forest Fire Research, 

Coimbra. Portugal. 14 p.

Robertson, G.; Damrel, D.; Hurja, J.; Leahy, S. 2002. Terrestrial ecosystem 

survey of the Peloncillo watershed study area. Draft Rep. Albuquerque, 

NM: U.S. Department of Agriculture, Forest Service, Southwestern Region.

Slauson, L.; Dalton, V.; Dalton, D. 1999. Effects of prescribed burning on the 

Palmer agave and the lesser long-nosed bat. In: Gottfried, G. J.; Eskew, 

L.G.; Curtin, C.G.; Edminster, C.B., comps. Toward integrated research, 

land management and ecosystem production in the Malpai Borderlands: 

conference proceedings. Proceedings RMRS-P-10. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain research 

Station: 111-114. 

Stropki, Cody L.; Ffolliott, Peter F.; Gottfried, Gerald J. 2009. Water repellent soils following prescribed burning treatments and a wildfire in the 

oak savannas of the Malpai Borderlands Region. Hydrology and Water 

Resources in Arizona and the Southwest. 39:5-8.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 345

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Impacts of Wildfire on Wildlife in Arizona: A Synthesis

Shari L. Ketcham and John L. Koprowski

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Abstract—Due to a century of fire suppression practices, the Madrean Archipelago regions in Arizona have 

accumulated excessive fuel loads that increase wildfire sizes, intensities, and frequencies. Wildfire induced 

structural changes in forest ecosystems can either benefit or adversely impact wildlife species. Therefore, it 

is imperative to understand how wildlife species react to such ecosystem changes after wildfires in both the 

short-term and long-term time periods. We examined scientific literature to determine wildlife distribution, 

abundance, elevational migration, and behavioral changes (i.e. how wildlife use resources post-fire) in response 

to wildfire. Understanding the impacts of wildfire-induced habitat fragmentation and creation of edge effects 

on wildlife species will provide information about overall forest condition and forest management practices.

Introduction

The Madrean Archipelago stretches from the Mogollon Rim in 

northern Arizona to the Sierra Madre Occidental in northwestern 

Mexico (Ffolliott and others 1995; Warshall 1995). The archipelago 

consists of 40 Sky Islands, which are a series of isolated mountain 

ranges that contain several vegetative zones that extend upwards from 

flat, low-lying arid regions (Ffolliott and others 1995; McLaughlin 

1995). Sky Islands have geographically isolated species since the last 

glaciopluvial event and therefore harbor elevated species diversity 

and richness (Lomolino and others 1989; Warshall 1995). The 19 Sky 

Island complexes in Arizona provide refuge for the great diversity of 

mammals, reptiles, and ants (Warshall 1995). However, biodiversity 

of endemic montane species is negatively impacted by grazing, soil 

erosion, introduced exotic species, habitat fragmentation and wildfire. 

Fire suppression practices for over a century have caused an increase 

in fuel loads that allow wildfires to increase in frequency, intensity, and 

size (Covington and Moore 1994; Sackett and others 1994; Swetnam 

1990). These large, intense fires are rapidly changing ecosystems and 

these cover type conversions may have adverse impacts on flora and 

fauna. The occurrence of such large scale, high intensity wildfires is 

predicted to continue to increase in the region exacerbated by climate 

change (Westerling and others 2006). Herein, we briefly review the 

historical pattern of fire and the effects of wildfire on wildlife in 

Arizona.

Wildfire Impacts

Historical Pattern

Before 1900, low-severity ground fires were common and burned 

every 2-15 years (Brown and Smith 2000; Kiltie 1989; Swetnam and 

Baisan 1996b; Thomas and McAlpine 2010). Since fire suppression 

practices altered the natural fire regime in Arizona, catastrophic standreplacing wildfires are increasing in frequency and have burned over 

1.5 million ha within the last decade (fig. 1; Southwest Coordination 

Center 2012). Flora and fauna species may not be adapted to these 

increasingly large and intense wildfires (Swetnam and Baisan 1996b). 

Fires can create a mosaic of burn severities across the landscape due 

to varying fuel load accumulations and other factors, particularly, in 

montane forests where pine-oak, mixed-conifer, and ponderosa pine 

(Pinus ponderosa) dominate the upper vegetative zones. Depending on 

the size of burn severity patches, these mosaics can fragment habitats, 

create edge effects, and change vegetative structure and composition. 

Therefore, understanding the impacts of these mosaic patterns of burn 

severities on wildlife species is ecologically important. 

Wildlife

Common wildfire impacts on wildlife include direct mortality, 

injury, increased predation from lack of cover, or starvation from lack 

of food availability (fig. 2; DeBano and others 1998; Ream 1981). 

Generalist species, such as coyotes (Canis latrans) and great horned 

owls (Bubo virginianus), are typically not significantly impacted by 

wildfires since they may exhibit prey switching if other food resources 

are limited. Specialist species are more likely to be adversely impacted 

by wildfire than generalist species since specialists typically concentrate on a single resource. The ability for wildlife to survive wildfires 

depends on food availability, cover, mobility, behavior, and structural 

diversity (DeBano and others 1998; Lyon and others 1978; Patton 

1992). Structural diversity is the uniformity, severity, size, intensity, 

season, and duration of the fire that creates edges between adjacent 

vegetative types (edge effect), creates snags that provide cover or 

cavities for use by wildlife, and creates environmental heterogeneity through the mosaic effect (DeBano and others 1998; Lyon and 

others 2000; Wright and Bailey 1982). Wildlife can be impacted by 

structural diversity if a wildfire burns uniformly at a moderate to high 

intensity rate across a large area due to lack of vegetation and cover 

throughout that landscape. Some highly mobile wildlife species will 

be able to travel long distances to emigrate from those areas; however, 

other wildlife species may try to emigrate but cannot get out of the 
346 USDA Forest Service Proceedings RMRS-P-67. 2013

Ketcham and Koprowski Impacts of Wildfire on Wildlife in Arizona: A Synthesis

and others 1998; Howard and others 1959). Some wildlife species will 

not leave nests, burrows, cavities, or dens during fire, which subjects 

them to smoke inhalation and possibly death by suffocation. However, 

many fossorial mammals can survive wildfires since most burrow 

systems are extensive, subterranean tunnels that protect against heat 

intensity (Erwin and Stasiak 1979; Lyon and others 2000; Sutherland 

and Dickman 1999; Vernes 2000). 

Wildfires that burn during the spring typically impact more wildlife 

species than wildfires that burn during any other season. Because 

spring is the time when breeding occurs for a majority of species, 

fires that erupt during this critical period can destroy nests, dens, 

burrows, or cavities and/or make them more open to predation by 

opening tree canopies or understory cover (Ward 1968). Adults are 

sometimes unable to emigrate from nests, dens, burrows, or cavities 

during fire due to the lack of mobility of offspring (Koprowski and 

others 2006; Lyon and others 2000). During fire, young animals are 

more prone to injury or mortality than adults; however, wildlife with 

high reproduction rates allow faster population recovery post-fire 

(Lyon and others 2000).

Granivores and Insectivores—Wildfire impacts granivore and 

insectivore species through loss of resources; however, some of these 

species respond positively to wildfires due to adaptation to open 

space or added food and cover over time (Lyon and others 2000). 

Birds in southeastern Arizona such as mourning doves (Zenaida 

macroura), vesper sparrows (Pooecetes gramineus), savannah sparrows (Passerculus sandwichensis), lark sparrows (Chondestes grammacus), horned larks (Eremophila alpestris; Bock and Bock 1992), 

and hairy woodpeckers (Picoides villosus) in northern Arizona forests 

(Covert-Bratland and others 2006) are typically found in burned 

areas due to an increase in seed production and insect infestations 

post-fire (Bock and Bock 1992, Covert-Bratland and others 2006). 

Woodpeckers emigrate into high-severity burns where most trees 

have been killed due to an abundance of insects and cavities (Lyon 

and others 2000), but home range sizes increase over time since the 

fire, which indicates that initially habitat quality increases but then 

is reduced over time (Covert-Bratland and others 2006). Other birds 

in southeastern Arizona such as Eastern meadowlarks (Sturnella 

magna), Cassin’s sparrows (Aimophila cassinii), Botteri’s sparrows 

(A. botteri), and grasshopper sparrows (Ammodramus savannarum), 

avoid burned areas for 2-3 years post-fire due to lack of grass and 

shrub cover (Bock and Bock 1992). Most birds will typically escape 

fire due to their extreme vagility (Erwin and Stasiak 1979; Gelluso 

and others 1986; Hakala and others 1971; Lyon and others 2000; 

Peres 1999); however, most ground-dwelling birds are likely to be 

adversely impacted by fire (Lyon and others 2000). 

 Small mammals also respond differentially to wildfire. Lack of 

food and/or predation are typical causes for small mammal decline 

within the first 2 months post-fire (McMurry and others 1996; Simons 

1991). At Fort Huachuca Military Reservation, presence of species 

did not change post-fire for Merriam’s kangaroo rat (Dipodomys 

merriami), silky pocket mice (Perognathus flavus), American deer 

mice (Peromyscus maniculatus), hispid pocket mice (C. hispidus), 

Northern grasshopper mice (Onychomys leucogaster), and Southern 

grasshopper mice (O. torridus), which indicates that these species did 

not emigrate from burned areas immediately post-fire (Litt and Steidl 

2011). However, presence decreased on burned areas for Northern 

pygmy mice (Baiomys taylori), fulvous harvest mice (Reithrodontomys 

fulvescens), and Arizona cotton rat (Sigmodon arizonae), which may 

indicate direct mortality, predation from lack of cover, starvation from 

lack of food, or emigration to unburned areas immediately post-fire 

(Litt and Steidl 2011; Steidl and Litt 2009). Arizona cotton rat, fulvous 

harvest mice, and Merriam’s kangaroo rat increased in abundance in 

Figure 1—Hectares burned in Arizona 2001-2011.

Figure 2—Impacts of fire on wildlife species. Arrows mean influences 

that can impact wildlife species.

moderate to high-severity patch. Ultimately, those species that cannot 

find their way out can potentially die of starvation or predation. 

Most wildlife deaths during wildfires are due to smoke inhalation, 

direct burn, and behavior such as the inability or reluctance to evacuate 

(Bock and Lynch 1970; Buech and others 1977; Bulan and Barrett 

1971; Chew and others 1959; Harrison and Murad 1972; Lyon and 

others 2000). Direct burns are a result of a wildfire that moves extremely fast and burns at an exceptionally high intensity, usually >63 

°C, which is lethal to both small and large wildlife species (DeBano 
USDA Forest Service Proceedings RMRS-P-67. 2013 347

Impacts of Wildfire on Wildlife in Arizona: A Synthesis Ketcham and Koprowski

burned areas while silky pocket mice, Northern grasshopper mice, and 

desert pocket mice (Chaetodipus penicillatus) decreased in abundance 

in burned areas (Litt and Steidl 2011; Steidl and Litt 2009). Abundances 

of yellow nosed cotton rats (S. ochrognathus), hispid pocket mice, 

and Northern pygmy mice did not change post-fire (Litt and Steidl 

2011). In the Mazatzal Mountains, Ord’s kangaroo rat (D. ordii), Merriam’s kangaroo rat, cactus mice (Peromyscus eremicus), and pocket 

gophers (Thomomys spp.) use burned areas more than unburned areas, 

whereas Bailey’s pocket mice (C. baileyi) use unburned areas more 

than burned areas (Monroe and others 2004). Merriam’s kangaroo rats 

respond positively to burned areas due to being a generalist species 

and favoring open areas, which adversely impacts Bailey’s pocket 

mice due to lack of cover (Price 1978; Rosenzweig and others 1975; 

Simons 1991). 

Wildfire may adversely impact tree squirrels due to reduction of 

nests, cavities, and food resources (Kirkpatrick and Mosby 1981) and 

through creation of edge effects and fragmented habitat, but most 

squirrels temporarily emigrate during wildfire and are successful in 

avoiding wildfire (Bendell 1974). However, endangered Mt. Graham 

red squirrels (MGRS; Tamiasciurus hudsonicus grahamensis) in 

the Pinaleño Mountains are adversely impacted by large, intense 

stand-replacing wildfires since these types of catastrophic fires did 

not occur historically (Koprowski and others 2006). After the Nuttall 

fire in 2004, surveys were conducted to determine effects of wildfire 

on MGRS (Sanderson and Koprowski 2009). No MGRS carcasses 

were found but seven resident squirrels were not relocated post-fire 

(Koprowski and others 2006). However, a charred and dead Abert’s 

squirrel (Sciurus aberti) was found (Greer, personal communication). 

Mt. Graham red squirrels require a midden, a long-term central cache 

site, for winter survival. These midden sites are typically located in 

dense, live portions of forest where greater amounts of seedfall occur, 

which may be severely impacted by catastrophic wildfires that opens 

forests and kills trees (Koprowski and others 2006; Wood and others 

2007). The presence of small localized fire does not immediately 

trigger abandonment, as a MGRS was observed to continue to use 

a smoldering nest tree that had received a lightning strike (Merrick 

and others 2010). After surviving the direct effects of wildfire, many 

squirrel species respond positively to fire with equivalent or superior 

survival, decreased home range size in areas of reduced fire intensity 

and in areas where the fire did not reach the canopy, or use of burned 

areas more than unburned areas (Blount and Koprowski 2012; Doumas 

and Koprowski 2012; Gwinn 2011; Leonard and others 2010; Pasch 

and Koprowski 2011). Abert’s squirrels introduced to Mt. Graham 

used areas within the perimeter of severe burns more than endangered 

MGRS (Gwinn 2011). Mexican fox squirrels (S. nayaritensis) used 

areas that experienced low-severity burn more than unburned areas 

and those that experienced higher burn severity, which indicates that 

this species is adapted to the low-severity fires that were historically 

experienced in the Chiricahua Mountains of southeastern Arizona 

(Doumas and Koprowski 2012).

Lizards are typically impacted by wildfire due to changes in vegetative structure and composition (Means and Campbell 1981; Russell 

and others 1999). In the southern Mazatzal Mountains near Four 

Peaks, lizard abundances in burned areas were greatest in chaparral 

and forest due to an increase in insect infestation post-fire (Cunningham and others 2002). In burned chaparral, Sonoran spotted whiptail 

(Cnemidophorus sonorae), Gila spotted whiptail (C. flagellicaudus), 

eastern fence lizard (Sceloporus undulatus), and ornate tree lizards 

(Urosaurus ornatus) were most abundant post-fire (Cunningham and 

others 2002). In unburned chaparral, collared lizards (Crotaphytus 

collaris) and short horned lizards (Phrynosoma douglassi) were 

most abundant (Cunningham and others 2002). Other lizards such as 

western whiptail (C. tigris), plateau striped whiptail (C. velox), and 

little striped whiptail (C. inornatus) use both unburned and burned 

chaparral (Cunningham and others 2002). However, western whiptail 

lizards decrease in abundance over time while other lizards increase 

in abundance in burned chaparral (Cunningham and others 2002). In 

burned forest, plateau striped whiptail, ornate tree lizard, Sonoran spotted whiptail, western whiptail, Gila spotted whiptail, collared lizards, 

western banded gecko (Coleonyx variegatus), and Great Plains skink 

(Eumeces obsoletus) were most abundant post-fire (Cunningham and 

others 2002). In unburned forest, short horned lizards and Madrean 

alligator lizards (Elgaria kingi) were most abundant (Cunningham 

and others 2002). Eastern fence lizards and little striped whiptail use 

both unburned and burned forest (Cunningham and others 2002).

Herbivores—Herbivores are primarily impacted by wildfire in the 

short-term due to loss of vegetation structure and composition. For 

this reason, most herbivores select unburned areas initially post-fire. 

However, once vegetative regrowth occurs, most herbivores will 

immigrate to burned areas. In the San Francisco Peaks, elk (Cervus 

canadensis) frequently use high-severity burn areas once aspen 

resprout occurred (Bailey and Whitham 2002). High-severity burn 

areas regenerate aspen ramets faster and with greater biomass than 

unburned, low-severity, and moderate-severity sites (Bailey and 

Whitham 2002). Elk continually graze high-severity burn areas for 

up to 3 years post-fire, which decreases aspen biomass, but aspen 

biomass increases by threefold in moderate-severity burn areas since 

elk herbivory was not prevalent (Bailey and Whitham 2002). Desert 

bighorn sheep (Ovis canadensis mexicana) in the Santa Catalina 

Mountains also use burned areas that open dense understory (Cain 

III and others 2005), since vegetative removal increases their visibility and ability to avoid predators (Etchberger and others 1989; 

Krausman and others 1996, 2001; Wakelyn 1987). Unburned areas 

with excessive, dense understory will be abandoned by sheep because 

these areas become unsuitable for sheep persistence (Krausman and 

others 1996). Burned areas had higher visibility measurements than 

unburned areas because unburned areas had an increase of vegetation 

over a 32-year period of fire suppression (Krausman and others 1996). 

Collared peccaries (Pecari tajacu) on the Three Bar Wildlife Area 

avoided burned desert scrub and chaparral areas for 1-2 years post-fire 

due to lack of cover and food resources such as cactus, acorns, and 

legumes (O’Brien and others 2005). Lower elevations in unburned 

chaparral and desert scrub were used more frequently by peccaries 

post-fire because these areas provided more thermal cover (O’Brien 

and others 2005). However, after 2 years, burned chaparral vegetation 

regenerates and provides as much cover as unburned chaparral (O’Brien 

and others 2005). Conversely, burned desert scrub vegetation did not 

completely recover, which indicates that these areas did not provide 

adequate cover for peccaries, even after 4 years post-fire (O’Brien 

and others 2005). White-throated woodrats (Neotoma albigula) in 

the Mazatzal Mountains were found to use unburned areas more than 

burned areas post-fire (Monroe and others 2004). Fire likely kills 

many woodrats since they usually do not emigrate from nests during 

fire (Quinn 1979; Simons 1991; Tevis 1956); however, woodrats that 

do emigrate during fire are found more frequently in unburned areas 

(Monroe and others 2004).

Carnivores and Omnivores—Wildfire impacts on carnivores and 

omnivores are rarely studied because most are generalists that are 

vagile and exhibit prey switching when main food resources are 

limited. Female black bears (Ursus americanus) in the Mazatzal 

Mountains used unburned patches 90% of the time (Cunningham 

and others 2003). Before the fire, female black bears with cubs used 

higher elevations to avoid males from predating their offspring; 
348 USDA Forest Service Proceedings RMRS-P-67. 2013

Ketcham and Koprowski Impacts of Wildfire on Wildlife in Arizona: A Synthesis

however, females with cubs switched use from higher elevations 

to lower elevations post-fire likely due to lack of food resources 

and cover (Cunningham and Ballard 2004). Black bear survival 

post-fire is high; however, juvenile and female populations and cub 

recruitment declined (Cunningham and Ballard 2004). Male black 

bears primarily use burned areas and have larger home ranges postfire (Cunningham and Ballard 2004), which indicate that resources 

were lacking in burned areas (Cunningham and others 2003). In the 

southern Mazatzal Mountains, gray foxes (Urocyon cinereoargenteus) 

use unburned areas more frequently than burned areas, while coyotes 

use both unburned and burned areas (Cunningham and others 2006). 

Gray foxes primarily use unburned areas because soft mast made up 

the majority of their diet (Cunningham and others 2006). Since over 

90% of vegetation was lost post-fire, this influences gray foxes to 

decline in abundance likely due to starvation (Cunningham and others 

2006). Coyote abundance did not seem to be impacted by wildfire, 

likely due to being a generalist species and an opportunistic predator 

(Cunningham and others 2006). In the Coconino National Forest and 

Coronado National Forest, Mexican spotted owls (Strix occidentalis) 

were not adversely impacted in the short term by presence or severity 

of wildfire, even though these forests have experienced catastrophic 

stand-replacing wildfires in the past (Jenness 2000). This outcome 

is likely due to owls being extremely vagile, generalist species.

Discussion

Wildfire can either positively or adversely impact wildlife species 

depending upon how intensely the fire burns, size of the burn severity 

patches, and the mosaic pattern left behind. Unfortunately, most studies do not incorporate these factors into their research design. Most 

forest dwelling wildlife species in the southwest are likely adapted to 

frequent, low-severity ground fires (Brown 1982; Humphrey 1974; 

Marshall 1963; Wright and Bailey 1982); however, increased fuel 

load accumulations from a century of fire suppression practices have 

created large, high-intensity crown fires and most wildlife species have 

not yet adapted to these conditions (Bendell 1974; DeBano and others 

1998; Singer and Schullery 1989). Wildfire impacts wildlife species by 

reduction or loss of food and cover and overall changes in structural 

diversity (Lyon and others 1978; Lyon and others 2000; Patton 1992). 

Vagile, generalist species are probably only impacted by wildfire in 

the short term, since they will exhibit prey switching when resources 

are limited thus being able to find other food resources throughout 

a mosaic of burn severity types. Specialist and small, non-fossorial 

species are likely to be impacted the greatest by wildfire via direct 

mortality and loss of food resources and cover (DeBano and others 

1998; Erwin and Stasiak 1979; Lyon and others 2000; Sutherland and 

Dickman 1999; Vernes 2000). Wildfires tend to create a more open 

forest understory and a mosaic of burn severities across the landscape 

that can fragment habitats, create edge effects, and change vegetative structure and composition. Some wildlife species tend to thrive 

in these areas, while others are hindered because they require cover 

to protect them from predation. Most mammal and avian predators 

increase their use of burned areas, which further exacerbates mortality 

of wildlife species post-fire (Lawrence 1966, Lyon and others 2000). 

Management Implications

Fire is successful at reducing fuel loads, an important objective where 

fuel has accumulated to unprecedented levels, but may have adverse 

impacts on some wildlife species. Most studies do not distinguish use 

in burned areas. We suggest that research design must include use 

versus availability models to produce more significant results. We are 

also suggesting that burn severities must also be distinguished. This 

will allow managers to better understand how various burn severities can impact wildlife. Managers must understand life histories of 

individual wildlife species to understand how they may be impacted 

by wildfire and manage accordingly; however, effects of fire on many 

wildlife species are relatively unknown. Recent wildfires in Arizona 

emphasize our lack of knowledge about the short-term and long-term 

effects of wildfire on wildlife, while providing a unique opportunity 

to undertake such studies.

References

Bailey, J. K.; Whitham, T. G. 2002. Interactions among fire, aspen, and elk 

affect insect diversity: reversal of a community response. Ecology. 83(6): 

1701–1712.

Bendell, J. F. 1974. Effects of fire on birds and mammals. New York: Academic Press. 553 p.

Blount, S. J.; Koprowski, J. L. 2012. Response of the Mount Graham red 

squirrel (Tamiasciurus hudsonicus grahamensis) to postfire conditions. 

Southwestern Naturalist. 57(1): 8–15.

Bock, C. E.; Lynch, J. F. 1970. Breeding bird populations of burned and 

unburned conifer forests in the Sierra Nevada. Condor. 72(2): 182–189.

Bock, C. E.; Bock, J. H. 1992. Response of birds to wildfire in native versus 

exotic Arizona grassland. Southwestern Naturalist. 37(1): 73–81.

Brown, James K.; DeByle, Norbert V. 1982. Developing prescribed burning 

prescriptions for aspen in the Intermountain West. Proceedings of the 

symposium: Fire—its field effects; 1982 October 19-21; Jackson, WY. 

Missoula, MT: Intermountain Fire Council; Pierre, SD: South Dakota 

Division of Forestry, Rocky Mountain Fire Council: 29-49.

Brown, J. K.; Smith, J. K. 2000. Wildland fire in ecosystems: effects of fire on 

flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Research Station. 257 p.

Buech, R. R., Siderits, K.; Radtke, R. E.; Sheldon, H. L.; and Elsing, D. 

1977. Small mammal populations after a wildfire in Northeast Minnesota. 

Res. Pap. NC–151. St. Paul, MN: U.S. Department of Agriculture, Forest 

Service, North Central Forest Experiment Station. 12 p.

Bulan, C. A.; Barrett, G. W. 1971. The effects of two acute stresses on the 

arthropod component of an experimental grassland ecosystem. Ecology. 

52(4): 597–605.

Cain III, J. W.; Johnson, H. E.; and Krausman, P. R. 2005. Wildfire and desert 

bighorn sheep habitat, Santa Catalina Mountains, Arizona. Southwestern 

Naturalist. 50(4): 506–513.

Chew, R. M., Butterworth, B. B.; and Grechman, R. 1959. The effects of fire on 

the small mammal populations of chaparral. Journal of Mammalogy. 40: 253.

Covert-Bratland, K. A., Block, W. M.; and Theimer, T. C. 2006. Hairy woodpecker winter ecology in ponderosa pine forests representing different 

ages since wildfire. Journal of Wildlife Management. 70(5): 1379 –1392.

Covington, W. W.; Moore, M. M. 1994. Southwestern ponderosa forest structure: 

Changes since Euro-American settlement. Journal of Forestry. 92(1): 39–47.

Cunningham, S. C.; Babb, R. D.; Jones, T. R.; Taubert, B. D.; and Vega, R. 

2002. Reaction of lizard populations to a catastrophic wildfire in a central 

Arizona mountain range. Biological Conservation. 107(2): 193–201.

Cunningham, S. C.; Ballard, W. B.; Monroe, L. M.; [and others]. 2003. Black 

bear habitat use in burned and unburned areas, central Arizona. Wildlife 

Society Bulletin. 31(3): 786–792.

Cunningham, S. C.; Ballard, W. B. 2004. Effects of wildfire on black bear 

demographics in central Arizona. Wildlife Society Bulletin. 32(3): 928–937.

Cunningham, S. C.; Kirkendall, L.; and Ballard, W. 2006. Gray fox and 

coyote abundance and diet responses after a wildfire in central Arizona. 

66(2): 169–180.

Debano, L. F.; Neary, D. G.; and Ffolliott, P. F. 1998. Fire’s effects on ecosystems. New York: John Wiley and Son, Inc. 352 p.

Doumas, S.; Koprowski, J. L. 2012. In Press. Return of fire as a restoration 

tool: long-term effects of burn severity on habitat use by Mexican fox 

squirrels. Restoration Ecology. 
USDA Forest Service Proceedings RMRS-P-67. 2013 349

Impacts of Wildfire on Wildlife in Arizona: A Synthesis Ketcham and Koprowski

Erwin, W. J.; Stasiak, R. H. 1979. Vertebrate mortality during the burning of 

a reestablished prairie in Nebraska. American Midland Naturalist. 101(1): 

247–249.

Etchberger, R. C.; Krausman, P. R.; and Mazaika, R. 1989. Mountain sheep 

habitat characteristics in the Pusch Ridge Wilderness, Arizona. Journal of 

Wildlife Management. 53(4): 902–907.

Ffolliott, P. F.; DeBano, L. F.; and Ortega-Rubio, A. 1995. Relationship of 

research to management in the Madrean Archipelago region. In: DeBano, 

L. H.; Ffolliott, P. H.; Ortega-Rubio, A.; [and others]; Biodiversity and 

management of the Madrean Archipelago: The sky islands of southwestern 

United States and northwestern Mexico; 1994 Sept. 19-23; Tucson, AZ. 

Gen. Tech. Rep. RM-GTR-264. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 31-35.

Geluso, K. N.; Schroder, G. D.; and Bragg, T. B. 1986. Fire-avoidance behavior of meadow voles (Microtus pennsylvanicus). American Midland 

Naturalist. 116(1): 202–205. 

Greer, V. L. 2009. [Personal communication to J. L. Koprowski].

Gwinn, N. 2011. Differential response to fire by an exotic and an endemic 

species complicates endangered species conservation. Thesis, University 

of Arizona, Tucson, Arizona, USA.

Hakala, J. B., Seemel, R. K.; Richey, R. A.; and Kurtz, J. E. 1971. Fire effects 

and rehabilitation methods —Swanson-Russian Rivers fires; 1971 April 

13-14; Fairbanks, AK. Proceedings of a symposium at the University of 

Alaska College. Portland, OR: U.S. Department of Agriculture, Forest 

Service, Pacific Northwest Forest and Range Experiment Station. 275 p.

Harrison, R. E.; Murad, J. L. 1972. Effects of annual prescribed burning on 

nematode populations from a Louisiana pine forest. Journal of Nematology. 4: 225–226.

Howard, W. E.; Fenner, R. L.; and Childs; H. E. 1959. Wildlife survival in 

brush burns. Journal of Range Management. 12(5): 230–234. 

Humphrey, R. R. 1974. Fire in the deserts and desert grassland of North 

America. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and ecosystems. 

New York, NY: Academic Press: 365-400.

Jenness, J. S. 2000. The effects of fire on Mexican spotted owls in Arizona and 

New Mexico. Thesis, Northern Arizona University, Flagstaff, Arizona, USA.

Kiltie, R. A. 1989. Wildfire and the evolution of dorsal melanism in fox squirrels, Sciurus niger. Journal of Mammalogy. 70(4): 726–739.

Kirkpatrick, R. L.; Mosby, H. S. 1981. Effect of prescribed burning on tree 

squirrels. In: Wood, G. W. Prescribed fire and wildlife in southern forests. 

Clemson, SC: Belle Baruch Forest Service Institute: 99–101.

Koprowski, J. L., Leonard, K. M.; Zugmeyer, C. A.; and Jolley, J. L. 2006. 

Direct effects of fire on endangered Mount Graham red squirrels. The 

Southwestern Naturalist. 51(1): 59–63.

Krausman, P. R., Long, G.; and Tarango, L. 1996. Desert bighorn sheep and 

fire, Santa Catalina Mountains, Arizona; 1996 March 11-15; Tucson, AZ. 

Proceedings Gen. Tech. Rep. RM-289. Ft. Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 162-168.

Krausman, P. R.; Dunn, W. C.; Harris, L. K.; Shaw, W. W.; and Boyce, W. M. 

2001. Can mountain sheep and humans coexist? Proceedings of the Second 

International Wildlife Management Congress. 2: 224–227.

Lawrence, G. E. 1966. Ecology of vertebrate animals in relation to chaparral 

fire in the Sierra Nevada foothills. Ecology. 47(2): 278–291.

Leonard, K. M.; Koprowski, J. L. 2010. Effects of fire on endangered Mount 

Graham red squirrels (Tamiasciurus hudsonicus grahamensis): response 

of individuals with known fates. Southwestern Naturalist. 55(2): 217–224.

Litt, A. R.; Steidl, R. J. 2011. Interactive effects of fire and nonnative plants 

on small mammals in grasslands. Wildlife Monographs. 176: 1–31.

Lomolino, M. V., Brown, J. H.; and Davis, R. 1989. Island biogeography 

of montane forest mammals in the American southwest. Ecology. 70(1): 

180–194.

Lyon, L. Jack; Crawford, Hewlette S.; Czuhai, Eugene; [and others]. 1978. 

Effects of fire on fauna: a state of knowledge review. Gen. Tech. Rep. WO6. Washington, DC: U.S. Department of Agriculture, Forest Service. 22 p.

Lyon, L. J.; Telfer, E. S.; and Schreiner, D. S. 2000. Direct effects of fire and 

animal responses. In: Smith, J. K. Wildland fire in ecosystems: effects of 

fire on fauna. Gen. Tech. Rep. RMRS-GTR-42. Ogden, UT : U.S. Department of Agriculture, Forest Service: 17–23.

Marshall, J. T., Jr. 1963. Fire and birds in the mountains of southern Arizona. 

Tall Timbers Fire Ecology Conference. 2: 135–141.

McLaughlin, S. P. 1995. An overview of the flora of the sky islands, southeastern 

Arizona: diversity, affinities, and insularity. In: DeBano, L. F., Gottfried, 

G. J., Hamre, R. H., and Edminster, C. B. Biodiversity and management 

of the Madrean Archipelago: the sky islands of southwestern United States 

and northwestern Mexico: A Symposium Proceedings. Gen. Tech. Rep. 

RM-GTR-264. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service: 60–70.

McMurry, S. T.; Lochmiller, R. L.; Boggs, J. F.; [and others]. 1996. Demography and condition of populations of white-footed mice (Peromyscus 

leucopus) in late and early successional habitats. Journal of Mammalogy. 

77(2): 335–345.

Means, D. B.; Campbell, H. W. 1981. Effects of prescribed burning on amphibians and reptiles. In: Wood, G. W. Prescribed fire and wildlife in southern 

forests: proceedings of a symposium; 1981 Apr 6-8; Myrtle Beach, SC. 

Georgetown, SC: The Belle W. Baruch Forest Science Institute of Clemson 

University: 89-96.

Merrick, M. J., Gwinn, R. N.; Minor, R. L.; [and others]. 2010. Endangered 

Mount Graham red squirrel (Tamiasciurus hudsonicus grahamensis) uses 

nest following lightning strike. Southwestern Naturalist. 55(1): 123–124.

Monroe, L. M., Cunningham, S. C.; and Kirkendall, L. B. 2004. Small mammal 

community responses to a wildfire on a central Arizona sky island. Journal 

of the Arizona Nevada Academy of Science. 37(2): 56–61.

O’Brien, C. S., Boyd, H. M.; Krausman, P. R.; [and others]. 2005. Influence of 

wildfire and coyote presence on habitat use by collared peccaries. Wildlife 

Society Bulletin. 33(3): 865–875.

Pasch, B., and Koprowski, J. L. 2011. Impacts of fire suppression on space 

use by Mexican fox squirrels. Journal of Mammalogy. 92(1): 227–234.

Patton, D. R. 1992. Wildlife habitat relationships in forested ecosystems. 

Portland: Timber Press. 442 p.

Peres, C. A. 1999. Ground fires as agents of mortality in a central Amazonian 

forest. Journal of Tropical Ecology. 15(4): 535–541.

Price, M. V. 1978. The role of microhabitat in structuring desert rodent communities. Ecology. 59(5): 910–921.

Quinn, R. D. 1979. Effects of fire on small mammals in the chaparral. 

California-Nevada Wildlife Transactions. 1979: 125–133.

Ream, C. H. 1981. The effects of fire and other disturbances on small mammals 

and their predators: an annotated bibliography. Gen. Tech. Rep. INT-106. 

Ogden, UT: U.S. Department of Agriculture, Forest Service: 1–64.

Rosenzweig, M. L., Smigel, B.; and Kraft, A. 1975. Patterns of food, space 

and diversity. Monographiae Biologicae. 28: 241–268.

Russell, K. R.; Van Lear, D. H.; and Guynn, D. C., Jr. 1999. Prescribed fire 

effects on herpetofauna: review and management implications. Wildlife 

Society Bulletin. 27(2): 374-384.

Sackett, S. S.; Haase, S.; and Harington, M. G. 1994. Restoration of southwestern ponderosa pine ecosystems with fire. In: Covington, W.W.; DeBano, L.F. Sustainable Ecological Systems: Implementing an Ecological 

Approach to Land Management. Gen. Tech. Rep. RM-247. Flagstaff, AZ: 

U.S. Department of Agriculture, Forest Service: 115–121.

Sanderson, H. R.; Koprowski, J. L. 2009. The last refuge of the Mt. Graham 

red squirrel: ecology of endangerment. Tucson: University of Arizona 

Press. 360 p.

Simons, L. H. 1991. Rodent dynamics in relation to fire in the Sonoran desert. 

Journal of Mammalogy. 72(3): 518–524.

Singer, F. J.; Schullery, P. 1989. Yellowstone wildlife: populations in process. 

Western Wildlands. 15: 18–22.

Southwest Coordination Center. 2012. Historical fires and acres. [Accessed 25 

April 2012]. <http://gacc.nifc.gov/swcc/predictive/intelligence/ytd_historical_data/historical/wf/average_per_year_suppression.pdf>. 

Steidl, R. J.; Litt, A. R. 2009. Do plant invasions change the effects of fire on 

animals? Fire Ecology. 5(1): 56–66.

Sutherland, E. F.; Dickman, C. R. 1999. Mechanisms of recovery after fire 

by rodents in the Australian environment, a review. Wildlife Research. 

26(4): 405–419.
350 USDA Forest Service Proceedings RMRS-P-67. 2013

Ketcham and Koprowski Impacts of Wildfire on Wildlife in Arizona: A Synthesis

Swetnam, T. W. 1990. Fire history and climate in the southwestern United 

States. In: J. S. Krammes, J. S., Effects of fire in management of southwestern natural resources: A Symposium Proceedings. Gen. Tech. Rep. 

RM-GTR-191. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service: 6–17.

Swetnam, T.W.; Baisan, C.H. 1996b. Fire histories of montane forests in 

the Madrean Borderlands. In: Ffolliott, P.F.; DeBano, L.F.; Maker, M.B., 

Jr.; [and others], eds. Effects of Fire on Madrean Province Ecosystems: 

A Symposium Proceedings. Gen. Tech. Rep. RM-GTR-289. Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service: 15–36.

Tevis, L. 1956. Effect of a slash burn on forest mice. Journal of Wildlife 

Management. 20(4): 405–409.

Thomas, P. A.; McAlpine, R. S. 2010. Fire in the forest. New York: Cambridge 

University Press. 238 p.

Vernes, K. 2000. Immediate effects of fire on survivorship of the northern 

bettong (Bettongia tropica): an endangered Australian marsupial. Biological 

Conservation. 96(3): 305–309.

Wakelyn, L. A. 1987. Changing habitat conditions on bighorn sheep ranges in 

Colorado. Journal of Wildlife Management. 51(4): 904–912.

Ward, P. 1968. Fire in relation to waterfowl habitat of the delta marshes. In: 

Proceedings, 8th annual Tall Timbers fire ecology conference. Proceedings 

of the Tall Timbers Fire Ecological Conference. 8: 254–267.

Warshall, P. 1995. The Madrean sky island archipelago: a planetary overview. 

In: DeBano, L. F.; Gottfried, G. J.; Hamre, R. H.; and Edminster, C. B., 

eds. Biodiversity and management of the Madrean Archipelago: the sky 

islands of southwestern United States and northwestern Mexico: A Symposium Proceedings. Gen. Tech. Rep. RM-GTR-264. Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service: 6–18.

Wood, D. J. A.; Drake, S.; Rushton, S. P.; [and others]. 2007. Fine-scale analysis 

of Mount Graham red squirrel habitat following disturbance. Journal of 

Wildlife Management. 71(7): 2357–2364.

Wright, H. A.; Bailey, A. W. 1982. Fire ecology-United States and Southern 

Canada. New York: John Wiley and Sons. 528 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 351

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Burned Saguaro: Will They Live or Die? 

Marcia G. Narog and Bonni M. Corcoran

	 USDA, Forest Service, Pacific Southwest Research Station, Riverside, California

Ruth C. Wilson

	 Department of Biology, California State University, San Bernardino, California

Abstract—Thousands of acres of giant saguaro (Carnegiea gigantea) habitat in the Sonoran Desert have 

been scorched by fire in recent decades. Resource managers struggle to maintain scenic landscapes featuring majestic saguaro faced with the challenges of fire, non-native species invasion, and recreational needs 

of millions of annual visitors. Successfully managing this iconic plant community requires understanding its 

fire resilience or intolerance. We compared survival of 162 saguaro 10 years after two wildfires on the Mesa 

Ranger District, Tonto National Forest, Arizona. Six 350-m point-quarter transects were randomly placed 

among burned and unburned areas. Four individual saguaro were measured at each of 8 points spaced 5 m 

apart along each transect. Ten years post-fire, high saguaro mortality and stunted growth were observed in 

both burns. Average saguaro mortality was 32% with apical height growth of 0.9 m after the Vista View fire 

compared to 51% and 0.6 m after the higher severity River fire. Unburned areas had 7% saguaro mortality and apical growth of 1.13 m. High compared to low fire severity burned areas nearly doubled saguaro 

mortality and stunted the growth of surviving saguaro by 30%. Our results suggest that maintaining strict 

fire prevention and suppression measures in high value saguaro habitat may be necessary to ensure that 

expansive vistas will continue to include this iconic cactus.

Introduction

Annually, about six million visitors travel to the Sonoran Desert 

on the Tonto National Forest, Arizona, to view a tourism favorite, 

the giant saguaro cactus (Carnegiea gigantea). Sadly, thousands of 

acres of this cactus’s thorn-shrub vegetation community have been 

scorched by fire during the last 4 decades (Narog and others 1995). 

Because of its slow growth and great size at maturity, the saguaro 

and its habitat will take a century or more to return to their previous 

grandeur after fire. Presently, maintaining and restoring this ecosystem 

poses a multidimensional problem. Resource managers of the saguaro 

thorn-shrub habitat face challenges managing this ecosystem such as 

recurrent fire, uncertain saguaro retention and regeneration, invasion 

of non-native species (Cave and Patten 1984; D’Antonio and Vitousek 

1992; Tellman 2002) and the recreational needs for millions of annual 

visitors. 

Natural saguaro attrition and recruitment can be highly variable. 

Turner (1990) followed two different saguaro populations over 29 

years that had 26 and 80% mortality; recruitment was correspondingly 11% vs. 76%. He found that episodic recruitment corresponded 

to heavy precipitation years. Humphrey (1984), and more recently 

Brooks and Pyke (2002), noted how effects of human activities, such 

as grazing, shortened fire return intervals, and increases in invasive 

plants, have reduced biodiversity in all North American deserts. High 

fire mortality and difficulties predicting future fire and climatic events 

further complicate saguaro conservation. Comprehensive studies of 

the ecology and natural history of saguaro have been done (Pierson 

and Turner 1998; Robichaux 1999; Steenbergh and Lowe 1971, 

1977, 1983; Turner and others 1995). However, long-term post-fire 

quantitative work using measures of fire injury, mortality and recovery 

for saguaro and other Sonoran Desert plants are poorly documented 

(Wilson and others 1995, 1996, 1998). Reasons why some fire-injured 

saguaro persist over time and survive while others perish still need to 

be determined. Recent research has included studies on the impacts 

of fire and invasive species on soil ecology (Steers and Allen 2011) 

and the use of GIS relational database management systems for fire 

history analysis (Swantek and others 1997). Broadening the knowledge 

and scope of environmental changes with an expanded database and 

increasing the resource tools available for measuring change in this 

thorn-shrub desert habitat will facilitate the understanding of fire 

impacts on saguaro and its ecosystem’s response to burning.

We’ve been studying the long-term effects of fire on saguaro 

populations following two fires that burned thousands of hectares on 

the Mesa District, Tonto National Forest, Arizona (Narog and others 

1995). During May 1993, the Vista View fire burned 1,200 ha (3,000 

ac), including scenic vistas of saguaro along Highway 87. Nearby, 

during July 1995, about 4,047 ha (10,000 ac) of saguaro habitat burned 

in the high intensity River fire. Post-fire saguaro injury and mortality 

were apparent in both areas. However, why some burned saguaro 

lived and others died was not always clear (fig. 1). In this paper we 

present data on the growth and mortality of individual saguaro from 

the first 10 years after these two wildfires. 

Methods

The 1993 Vista View and 1995 River wildfires charred saguaro 

habitat in an area known as The Rolls, near Four Peaks Road on the 
352 USDA Forest Service Proceedings RMRS-P-67. 2013

Narog and others Burned Saguaro: Will They Live or Die? 

Tonto National Forest, Arizona (Wilson and others 1995). Saguaro 

measurement data were collected along six 350-m point-quarter line 

transects randomly placed in and near the two adjacent wildfires 

(one control and five within fire perimeters; fig. 2a). Transects were 

oriented north to south, and both burned and unburned saguaro plants 

were represented in the study population. Eight points, located at 50 

m intervals, were sampled along each transect. At each point, the 

nearest saguaro in each quarter was chosen for measurement (fig. 2b). 

Saguaro dimensions, fire exposure, and mortality were documented 

over time for 162 individual saguaro. The saguaro is highly variable 

in the number of branching arms each plant has and the contributions 

that the arms give to the total biomass of each cactus. Therefore, to 

estimate how growth differed among the burned and unburned saguaro 

individuals, we simply used maximum height gain or loss for comparisons. Transects had different numbers of saguaro due to overlap 

from previous 50 m sample points, variable topography, irregular 

burn perimeters, and roads. Fire severity was defined after personal 

communications with fire fighters who fought the fires and the Tonto 

National Forest Resource Officer as well as personal observations 

made of fuel reduction on our post-fire study.

Individual cacti were remeasured at each successive visit to the 

study area. Unburned transects were measured in 1994, 1997, 2000, 

2003 and 2005. Transects in the Vista View fire perimeter were visited 

in 1994, 1998, and 2003 (1, 5, and 10 years post-fire). The River fire 

transects were measured in 1997, 2000, and 2005 (2, 5, and 10 years 

post-fire). 

Results

Data from unburned saguaro sampled from 1994 through 2005 

are presented in table 1. Tables 2 and 3 show data from each saguaro 

exposed to fire over 10 years later. Transect 2 (fig. 2b) was unburned 

and had no recorded fire history. Some saguaro on Transect 3 escaped 

burning, but others were subjected to varying fire severity by the 

River fire. The River fire was less intense at its perimeter, which 

included part of Transect 3. More intense fire behavior was observed 

by fire personnel in Transect 4.5 (Kerr, personal communication). 

Observational assessment of fire intensity correlated well with the 

amount of fuel reduction found during our post-fire measurements. 

Transect 6 showed the greatest fire severity among the three transects 

burned in the Vista View fire (fig. 2a). Burn scar measurements on 

individual saguaro (Narog and Wilson unpublished data), in addition 

to the presence of abundant shrub and cactus skeletons, illustrated the 

lower intensity of the Vista View fire compared to most of the River 

fire.

Saguaro mortality and growth during the first 10 years after fire 

differed among the unburned, low fire severity, and high fire severity 

areas. We observed 7% mortality after 10 years of saguaro growing 

in unburned areas (table 4). Saguaro mortality was 5 to 10 times 

greater in burned than in unburned areas. Saguaro mortality for the 

three transects on the lower severity Vista View fire averaged 32%. 

Saguaro mortality varied on the River fire from 22% along Transect 

3 (low severity area) to 77% for Transect 4.5 (high severity area) 

(table 4). Smaller saguaro had greater mortality than larger; most 

individuals less than 4 m in height did not survive either fire. 

Height measurements of individual saguaro ranged from less than 

1 m to over 10 m. Saguaro height growth averaged 1.1 m per plant 

over 10 years in the unburned area (table 1) compared to 0.9 m for 

saguaro burned in the Vista View fire (table 2) and 0.6 m for the River 

Figure 1—Some post-fire saguaro survived and grew while others nearby 

died. Photo was taken 10 years after the 1995 River Fire, Tonto National 

Forest, Arizona. 

Figure 2—(a) Location of study and fire perimeters on the Mesa District, Tonto National Forest near Phoenix, Arizona. (b) Saguaro were 

sampled every 50 m along randomly placed 350-m transects in burned 

and unburned areas.
USDA Forest Service Proceedings RMRS-P-67. 2013 353

Burned Saguaro: Will They Live or Die? Narog and others

Table 1—Ten year point-quarter data for unburned Transect 2 and the unburned portion of 

Transect 3 showing saguaro live and dead heights (m) measured between 1994 and 2005, 

Tonto National Forest, Arizona (N = 30).

1994 1997 2000 2003 2005

Saguaro Live Dead Live Dead Live Dead Live Dead Live Dead

2 6.5 -- 7.0 -- 7.1 -- 7.2 -- 7.4 --

23 6.5 -- 5.8 -- 6.2 -- 6.4 -- 6.4 --

24 2.4 -- 2.9 -- 3.5 -- 3.8 -- 4.2 --

25 3.2 -- 3.4 -- 4.2 -- 4.5 -- 4.8 --

26 3.8 -- 4.0 -- 4.4 -- 4.6 -- 4.8 --

27 6.5 -- 7.5 -- 8.7 -- 9.0 -- 9.2 --

28 5.4 -- 6.1 -- 6.3 -- 6.7 -- 7.3 --

29 8.5 -- 9.0 -- 9.5 -- 10.1 -- 10.4 --

212 7.0 -- 8.0 -- 8.2 -- 8.5 -- 8.6 --

213 4.8 -- 3.5 -- 3.5 -- 3.5 -- 3.7 --

216 7.0 -- 7.7 -- 7.8 -- 8.1 -- 8.3 --

217 6.5 -- 7.3 -- 7.5 -- 8.0 -- 8.2 --

220 5.0 -- 5.6 -- 5.8 -- 6.0 -- 6.0 --

222 4.5 -- 4.8 -- 5.1 -- 5.1 -- 5.4 --

223 4.0 -- 4.7 -- 5.2 -- 6.2 -- 6.0 --

224 0.8 -- 1.1 -- 1.5 -- 1.8 -- 2.2 --

225 7.0 -- 7.7 -- 7.6 -- 7.8 -- 8.1 --

227 6.0 -- 6.1 -- 6.5 -- 6.4 -- 3.7 --

228 3.8 -- 4.1 -- 4.8 -- 5.2 -- 5.8 --

229 2.4 -- 3.3 -- 4.2 -- 4.6 -- 5.0 --

230 5.0 -- 5.7 -- 6.0 -- 6.2 -- 6.6 --

232 5.5 -- 6.3 -- 6.1 -- 6.4 -- 6.7 --

221 3.6 -- 3.6 -- -- 3.2 -- 3.3 -- 3.1

21 -- 2.5 -- 2.5 -- -- -- -- -- --

333 5.5 -- 6.0 -- 6.2 -- -- -- 7.4 --

335 2.0 -- 2.3 -- 2.9 -- -- -- 3.6 --

336 2.1 -- 3.4 -- 3.4 -- -- -- 4.2 --

342 4.5 -- 4.7 -- 5.1 -- -- -- 5.5 --

350 3.7 -- 4.2 -- 4.9 -- -- -- 6.1 --

358 7.7 -- 7.3 -- 7.7 -- -- -- 8.1 --

Sum 141.2 2.5 153.1 2.5 159.9 3.2 136.1 3.3 173.7 3.1

Average 4.9 5.3 5.7 6.2 6.2

Std dev 1.9 1.9 1.9 1.9 1.9

fire (table 3). By 2005, only six saguaro on Transect 4.5 survived from 

the 1995 River fire. Based on remnants of burned saguaro found along 

Transect 4.5 immediately after the River fire, we determined that 

saguaro had an initial loss in height. They measured approximately 

92 m in dead height compared with the 25 m of height of surviving 

saguaro. By 2005, the six remaining live saguaro had accumulated 6 

m of additional height. After 10 years, about 80 m of remnant saguaro 

skeletons were no longer evident and had decomposed. Only 12 m 

of dead saguaro remained.

In the Vista View fire, saguaro final total live height (286 m) was 

less than initial cumulative live saguaro height (343 m). Wind toppled 

or blew tops off some damaged saguaro in the Vista View area. The 

cumulative height of dead saguaro at 1 and 5 years post-fire was 

similar, about 20 m. 

Discussion 

Our long-term post-fire study has followed individual saguaro for a 

decade. Some surviving saguaro cacti and associated plants still appear 

to be suffering adverse effects from fire, even after 10 years. Unlike 

other studies tracking saguaro longevity and growth, we focused on 

fire-injured saguaro and associated vegetation in our observations. 

The desert thorn-shrub habitat had no recent documentation of fire 

effects on saguaro survival or mortality. Huge areas on the Tonto 

National Forest had recently burned when we initiated our study 

(Narog and others 1995). Subsequently, increasing public appreciation 

for desert vistas and early results from our study led to improved fire 

management. A change was made in policies at the national forest, 

state, and local levels that recognized the fire sensitivity of saguaro 

and its associated ecosystem (Hunt 1997; Narog and others 1999; 

Wilson and others 1995). 

Short term saguaro mortality is only part of the story. After 10 years 

we discovered that saguaro were still dying from delayed effects of 

fire injury. Both short- and long-term saguaro mortality was higher 

in burned than in unburned areas of our study. Natural attrition in 

unburned sites was minimal; therefore, environmental anomalies and 

variable weather conditions would not account for the higher loss 

of saguaro or their reduced growth in burned areas. While saguaro 

showed long-term adverse effects from both low and high severity 

fires, they survived low fire severity better than high severity. Not 

all fire-injured saguaro died immediately after the fire (Narog and 

Wilson 2004): some persisted for longer periods of time, and others 
354 USDA Forest Service Proceedings RMRS-P-67. 2013

Narog and others Burned Saguaro: Will They Live or Die? Table 2—Ten year point-quarter data for Vista View fire burned Transects 1, 6 and 7 showing live and dead saguaro heights (m) measured between 1994 and 2003, Tonto National Forest, Arizona. 1994 1998 2003 Saguaro Live Dead Live Dead Live Dead T1* T6* T7* T1 T6 T7 T1 T6 T7 T1 T6 T7 T1 T6 T7 T1 T6 T7 T1 T6 T7 64 12 44 1.9 2.6 6.0 -- -- -- 2.6 2.7 6.2 -- -- -- 3.7 3.2 6.4 -- -- -- 11 32 83 6.5 4.0 3.3 -- -- -- 6.5 5.8 3.3 -- -- -- 3.2 4.6 3.8 3.5 -- -- 14 34 73 7.5 1.3 2.4 -- -- -- 6.5 1.6 2.5 -- -- -- 8.0 2.0 2.8 -- -- -- 44 41 11 5.5 6.3 3.1 -- -- -- 6.2 6.1 3.5 -- -- -- 6.4 3.6 4.0 -- -- -- 71 43 12 3.6 5.0 1.6 -- -- -- 4.5 6.0 1.7 -- -- -- 6.8 5.2 2.1 -- -- -- 13 52 13 8.5 5.0 5.8 -- -- -- 7.0 5.2 7.3 -- -- -- 8.0 6.4 8.0 -- -- -- 41 53 14 5.5 9.0 4.1 -- -- -- 6.2 9.5 4.8 -- -- -- 7.4 6.4 3.4 -- -- -- 51 61 22 8.5 1.4 8.0 -- -- -- 8.5 1.4 7.5 -- -- -- 8.7 1.4 6.2 -- -- -- 61 64 24 1.2 2.0 2.3 -- -- -- 1.6 2.2 2.3 -- -- -- 2.0 2.3 2.6 -- -- -- 12 71 34 8.0 2.3 4.0 -- -- -- 7.2 2.5 4.2 -- -- -- 8.3 3.1 5.7 -- -- -- 23 72 41 3.5 3.0 7.0 -- -- -- 3.9 3.3 7.2 -- -- -- 4.3 3.6 4.8 -- -- -- 24 73 51 4.5 3.8 4.2 -- -- -- 5.1 4.0 4.3 -- -- -- 5.7 4.6 5.4 -- -- -- 31 81 62 4.5 3.5 5.0 -- -- -- 4.7 3.5 5.3 -- -- -- 4.9 5.5 5.7 -- -- -- 33 22 63 4.5 5.5 5.9 -- -- -- 5.1 6.9 5.8 -- -- -- 6.6 -- 4.2 -- 6.9 -- 42 44 71 6.0 8.5 2.5 -- -- -- 7.4 8.3 2.9 -- -- -- 7.9 -- 3.5 -- 8.1 -- 52 51 72 6.5 8.5 8.0 -- -- -- 7.1 5.9 8.0 -- -- -- 7.8 -- 3.8 -- 5.1 -- 53 54 74 4.5 2.5 2.9 -- -- -- 6.6 -- 3.0 -- 0.2 -- 5.1 -- 3.5 -- 1.2 -- 54 11 81 6.5 -- 4.0 -- 2.5 -- 7.2 -- 4.2 -- 2.3 -- 7.5 -- 4.8 -- 3.9 -- 73 13 84 6.0 -- 3.0 -- 1.3 -- 6.2 -- 3.1 -- 1.1 -- 7.2 -- 3.0 -- 9.4 -- 81 14 31 7.0 -- 5.3 -- 0.5 -- 5.2 -- 5.5 -- 0.5 -- 8.9 -- -- -- 2.5 6.3 83 21 61 6.0 -- 8.5 -- 2.0 -- 5.3 -- 9.0 -- 1.9 -- 6.8 -- -- -- 5.4 9.6 84 23 33 4.3 -- 6.5 -- 0.8 -- 3.6 -- 6.4 -- 0.4 -- 5.4 -- -- -- 2.5 7.6 82 31 52 4.0 -- 7.8 -- 2.0 -- 6.3 -- 7.8 -- 2.0 -- 5.6 -- -- -- 2.0 7.0 34 82 53 3.1 -- 13.0 -- 0.5 -- 3.6 -- 13.0 -- 0.6 -- 4.1 -- -- -- 12.5 -- 62 84 64 4.3 -- 5.3 -- 0.2 -- 4.5 -- 7.0 -- 0.1 -- -- -- -- -- 0.7 7.0 22 83 21 2.5 -- 4.8 -- 0.3 -- 2.5 -- -- -- -- 0.7 -- -- -- 0.9 -- -- 62 74 54 0.5 -- -- -- 1.5 5.0 -- -- -- 0.7 -- 5.5 -- -- -- -- -- 3.3 21 -- -- -- -- -- 2.6 -- -- -- -- -- 3.0 -- -- -- -- -- 0.4 -- -- 32 -- -- -- -- -- 1.5 -- -- -- -- -- 1.0 -- -- -- -- -- -- -- -- Sum 134.9 74.1 134.2 4.1 11.6 5.0 141.1 74.8 135.8 4.7 9.0 6.2 150.3 51.9 83.7 4.8 60.2 40.8 Average 5.0 4.4 5.2 2.1 1.2 5.0 5.4 4.7 5.4 1.6 1.0 3.1 6.3 4.0 4.4 1.6 5.0 6.8 Std dev 2.1 2.5 2.5 0.8 0.8 0.0 1.7 2.4 2.6 1.3 0.8 3.4 1.9 1.6 1.5 1.7 3.6 2.1 *Transect 1: N = 29; Transect 6: N = 27; Transect 7: N = 27. Total N = 83.
USDA Forest Service Proceedings RMRS-P-67. 2013 355

Burned Saguaro: Will They Live or Die? Narog and others

Table 3—Ten year point-quarter data for River Fire burned Transects 3 and 4.5 showing live and dead saguaro heights (m) measured between 

1997 and 2005, Tonto National Forest, Arizona (N = 26).

1997 2000 2005

Saguaro Live Dead Live Dead Live Dead

T3 T4.5 T3 T4.5 T3 T4.5 T3 T4.5 T3 T4.5 T3 T4.5 T3 T4.5

34 2 4.1 5.8 -- -- 4.6 6.5 -- -- 5.3 6.7 -- --

38 3 1.8 4.5 -- -- 2.2 5.2 -- -- 2.8 5.6 -- --

39 5 6.8 4.0 -- -- 7.2 4.4 -- -- 7.9 5.1 -- --

43 21 4.0 2.3 -- -- 4.4 3 -- -- 4.7 3.5 -- --

44 28 6.2 4.5 -- -- 6.3 5.2 -- -- 6.5 6.0 -- --

45 32 7.4 3.9 -- -- 7.5 4.2 -- -- 7.8 4.5 -- --

46 1 6.3 3.2 -- -- 6.8 3.4 -- -- 7.1- -- -- 0.8

47 11 4.4 6.2 -- -- 5.1 6.5 -- -- 5.8 -- -- 2.0

48 7 6.0 3.7 -- -- 5.2 -- -- 1.7 6.6 -- -- 1.5

49 10 4.4 3.9 -- -- 5.1 -- -- 1.1 5.8 -- -- 1.2

52 17 3.8 4.6 -- -- 4.1 -- -- -- 4.8 -- -- 1.2

54 22 9.8 4.2 -- -- 10 -- -- 1.5 10.2 -- -- 1.5

55 9 8.0 3.3 -- -- 8.4 -- -- 1.5 8.8 -- -- --

56 13 5.2 8.3 -- -- 5.3 -- -- 4.1 5.9 -- -- --

57 16 2.3 8.3 -- -- 2.4 -- -- 3.5 2.8 -- -- --

59 24 3.4 7.5 -- -- 3.5 -- -- 7.5 3.8 -- -- --

62 4 5.5 -- -- 2.4 5.9 -- -- 2.3 6.5 -- -- 2.3

64 8 5.5 -- -- 1.7 5.9 -- -- 1.6 6.4 -- -- 1.6

63 12 5.0 -- -- 0.6 -- -- 1.5 0.6 -- -- 1.5 --

51 25 -- -- 0.6 2.9 -- -- 0.6 2.7 -- -- -- --

60 26 -- -- 04 4.1 -- -- 0.4 2.9 -- -- -- --

40 15 -- -- 1.5 3.9 -- -- -- -- -- -- -- --

41 20 -- -- 0.3 0 -- -- -- -- -- -- -- --

23 -- -- -- 2.4 -- -- -- -- -- -- -- --

27 -- -- -- 1.9 -- -- -- -- -- -- -- --

29 -- -- -- 4.6 -- -- -- -- -- -- -- --

Sum 99.9 78.2 6.4 24.5 99.9 38.4 2.5 31.0 109.5 31.4 1.5 12.1

Average 5.3 4.9 1.6 2.5 5.6 4.8 0.8 2.6 6.1 5.2 1.5

Std dev 2.0 1.8 1.7 1.4 2.0 0.2 0.6 2 1.9 1.1 0.5

Table 4—Percent mortality among all saguaro plots unburned 

and burned after the River and Vista View wildfires, Tonto 

National Forest, Arizona. Note: Six unburned saguaro are 

from Transect (T) 3. 

Transect Live Dead % Dead

Unburned

T2 & T3 28 2 7

Burned

River T3 18 5 22

River T4.5 6 20 77

Vista View T1 24 5 17

Vista View T6 13 14 52

Vista View T7 19 8 30

Sum 80 52

Average 39

ultimately perished. Our data show that 5 years after fire was not sufficient time to evaluate the impact of fire on the growth increments 

in fire-damaged plants or the ultimate fate of individuals from the 

burned saguaro populations. We hope to reassess the saguaro in this 

study at 20 years post-fire to determine whether further mortality has 

occurred and to see if height growth trends have changed since our 

last measurements.

Our results showed a loss and decomposition of nearly 80% of a 

healthy pre-burn saguaro population on one transect during the first 

10 years following a wildfire. Narog and Wilson (2005) documented 

persistent loss of associated vegetation that can serve as nurse plants 

in all the burned areas on our study sites 10 years after the fires. 

Turner and others (1966) found poor saguaro seedling establishment 

and survival when appropriate environmental conditions and nurse 

plants are lacking. Proactive restoration efforts for burned saguaro 

ecosystems may be needed particularly after high severity or shortinterval repeated fire. Maintaining strict fire prevention and suppression 

measures in high value saguaro habitat may be necessary to ensure 

that expansive vistas continue to include these iconic cacti.

Acknowledgments

We thank the Mesa Ranger District, Tonto National Forest, USDA, 

Forest Service Region 3 for partial funding and resources used for this 

project. We appreciate the great help from our field crew members 

Christina Escobar, Warren Hanna, Julie Lam, Valerie Oriol, Mark 

Parlow, Christie Sclafani, and Catherine Yang in field preparation, 

data collection, and data processing. Helpful review comments on an 

earlier draft of this paper were provided by Timothy Paysen, Melody 

Lardner, and Jan Beyers.

References

Brooks, M. L.; Pyke, D. A. 2002. Invasive plants and fire in the deserts of 

North America. In: Galley, K.E.; Wilson, T. P., eds., Proceedings of the 

invasive species workshop: The role of fire in the control and spread of 

invasive species. Fire Conference 2000: the First National Congress on Fire 

Ecology, Prevention, and Management. Misc. Pub. No. 11. Tallahassee, FL: 

Tall Timbers Research Station: 1-14. 

Cave, G. H.; Patten, D. T. 1984. Short-term vegetation responses to fire in 

the upper Sonoran Desert. Journal of Range Management 37(6):491-496. 
356 USDA Forest Service Proceedings RMRS-P-67. 2013

Narog and others Burned Saguaro: Will They Live or Die? 

D’Antonio, C. M.; Vitousek, P. M. 1992. Biological invasions by exotic 

grasses, the grass/fire cycle, and global change. Annual Review of Ecology 

and Systematics 23:63-87. 

Humphrey, R. R. 1974. Fire in the deserts and desert grasslands of North 

America. In: T.T. Kozlowki; C.E. Ahlgren, eds. Fire and Ecosystems. New 

York: Academic Press: 366-400. 

Hunt, Scott E. 1997. Fire management in the Sonoran Desert on state and private 

lands. Presentation 4 at the conference on Effects of Urbanization in the 

Sonoran Desert; October 8-10, 1997. Available online at: http://ag.arizona.

edu/oals/urbanization/fire.html [accessed 29 April 2012].

Narog, M. G.; Koonce, A. L.; Wilson, R. C.; Corcoran, B. M. 1995. Burning 

in Arizona’s giant cactus community. In: Weise, D. R.; Martin, R. E., tech. 

coords. The Biswell Symposium: Fire Issues and Solutions in Urban Interface and Wildland Ecosystems. Gen. Tech. Rep. PSW-GTR-158. Berkeley, 

CA: USDA Forest Service, Pacific Southwest Research Station: 175-176.

Narog, M. G.; Wilson, R. C. 2005. Post-fire saguaro community: impacts of 

associated vegetation still apparent 10 years later. Gottfried, G. J.; Gebow, 

B. S.; Eskew, L. G.; Edminster, C. B., comps. Connecting mountain islands 

and desert seas; biodiversity and management of Madrean Archipelago II. 

RMRS-P-36. Fort Collins, CO: USDA Forest Service, Rocky Mountain 

Research Station: 421-425.

Narog, M. G.; Corcoran, B. M.; Wilson, R. C.; Kerr, K. 1999. Fire management in the saguaro shrub. In: Sugihara, N. G.; Morales, M. E.; Morales, 

T. J., eds. Proceedings of the Symposium on Fire Management: Emerging 

Policies and New Paradigms. Association for Fire Ecology. Misc. Publ. 

No. 2:147-155.

Narog, M. G.: Wilson, R. C. 2004. Delayed mortality: Saguaro are still dying 

10 years after wildfire. P2.9. 2nd International Wildland Fire Ecology and 

Fire Management Congress; November 16-20, 2003. Available online at: 

www.ams.confex.com/ams/pdfpapers/66142.pdf [accessed 27 April 2012].

Pierson E. A.; Turner. R. M. 1998. An 85-year study of saguaro (Carnegiea 

gigantea) demography. Ecology 79: 2676-2693. 

Robichaux, R. H. (ed.). 1999. Ecology of Sonoran Desert Plants and Plant 

Communities. The University of Arizona Press, Arizona-Sonoran Desert 

Museum, AZ. 303 p.

Steenbergh, W. F.; Lowe, C. H. 1971. Ecology of the saguaro: I, the role of 

freezing weather in a warm-desert plant population. Research in the Parks, 

National Park Service Symposium Series 1:49-92.

Steenbergh, W. F.; Lowe, C. H. 1977. Ecology of the saguaro: II, reproduction, germination, establishment, growth and survival of the young plant. 

Science Monograph Series No 8. Washington, DC: U.S. Department of the 

Interior, National Park Service. 

Steenbergh, W. F.; Lowe, C. H. 1983. Ecology of the saguaro: III, growth and 

demography. Science Monograph Series No 17. Washington, DC: U.S. 

Department of the Interior, National Park Service.

Steers, R. J.; E. B. Allen. 2011. Fire effects on perennial vegetation in the 

western Colorado Desert, USA. Fire Ecology 7(3): 59-74.

Swantek, P. J.; Halvorson; W. H.; Schwalbe, C. R. 1997. The Use of GIS and 

the Internet for analyzing fire history of the Sonoran Desert: a regional 

approach in Arizona. Available online at: proceedings.esri.com/library/

userconf/proc97/proc97/to200/pap196/ [accessed 27 April 2012].

Tellman, Barbara. 2002. Invasive Exotic Species in the Sonoran Region. The 

University of Arizona Press, Arizona-Sonoran Desert Museum, AZ. 420 p.

Turner, R. M. 1990. Long-term vegetation change at a fully protected Sonoran 

Desert site. Ecology 71:464-477. 

Turner, R. M.; Bowers, J. E.; Burgess, T. L. 1995. Sonoran Desert Plants, an 

Ecological Atlas. The University of Arizona Press, Tucson, AZ; 146-149.

Turner, R.M.; Stanley, M.; Alcorn, G.O.; Booth, J. A. 1966. The influence 

of shade, soil, and water on saguaro seedling establishment. Botanical 

Gazette 127(2/3):95-102. 

Wilson R. C.; Narog, M. G.; Corcoran, B. M. 1996. Postfire saguaro injury in 

Arizona’s Sonoran Desert. In: Proceedings of conference on Effects of Fire 

on Madrean Province Ecosystems; March 11-15; Tucson, AZ. Gen. Tech. 

Rep. RM-GTR-289. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Forest and Range Experiment Station: 247-252.

Wilson R. C.; Narog, M. G.; Corcoran, B. M. 1998. Unburned fuels in an 

Arizona upland saguaro-shrub community. In: Reynolds, R.; Reynolds, 

J., eds. Proceedings of the Desert Research Symposium: Finding faults 

in the Mojave. Redlands, CA: San Bernardino County Museum SBCMA 

Quarterly 45(1, 2):109.

Wilson, R. C.; Narog, M. G.; Koonce, A. L.; Corcoran, B. M. 1995. Post fire 

regeneration in Arizona’s giant saguaro shrub community. In: Proceedings of 

Conference on Biodiversity and Management of the Madrean Archipelago: 

the Sky Islands of Southwestern United States and Northwestern Mexico; 

1994 September 19-23; Tucson, AZ. Gen. Tech. Rep. RM-GTR-264. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Forest and Range Experiment Station; 424-431.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 357

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Post-Wildfire Erosion in the Chiricahua Mountains

Ann Youberg

Arizona Geological Survey, Tucson, Arizona

Daniel G. Neary, Karen A. Koestner, and Peter E. Koestner

USDA Forest Service, Rocky Mountain Research Station, Flagstaff, Arizona

Abstract—The Horseshoe 2 Fire burned 90,226 ha (222,954 ac) of the Chiricahua Mountains in the 

Coronado National Forest of southeast Arizona from May 8 to June 25, 2011. This mountain range in the 

Madrean Archipelago was burned by widespread fires prior to 1890, numerous small fires after 1890, and, 

more recently, the 11,129 ha (27,500 ac) Rattlesnake Fire in 1994. The latter fire resulted in significant erosion during post-fire monsoon storms that eroded deep gullies (~10 m) in Ward Canyon (upper West Turkey 

Creek). Post-fire runoff deposited large amounts of sediment in lower channels, particularly in Rucker Creek 

and in Rucker Lake. The Horseshoe 2 Fire burned approximately 70% of the mountain range with a mosaic 

of fire severities. Post-fire erosion from floods and debris flows produced additional erosion and sedimentation that is currently being evaluated through fieldwork and repeat aerial photography. This paper discusses 

ongoing work regarding post-fire erosion and the impacts on channel systems in the Chiricahua Mountains.

Introduction

	 Record-setting wildfires burned across Arizona in 2011. The

human-caused Horseshoe 2 Fire burned 11,110 ha (222,954 acres)

of the Chiricahua Mountains between May 8th and June 25th, 2011,

on the Coronado National Forest (CNF) (http://inciweb.org/incident/2225/). This fire burned approximately 70% of the mountain

range (fig. 1) and currently stands as the 4th largest wildfire in state

history (Southwest Coordination Center: http://gacc.nifc.gov/swcc/

predictive/intelligence/intelligence.htm).Monsoonrainsoftenfollow

immediately after the wildfire season and are often how fires are ultimately extinguished. Short-duration, intense bursts of rainfall from

relatively common (<2-5 year frequency) storms can generate large

floods and debris flowsin watersheds disturbed by wildfires(DeBano

and others 1998; Jenkins, 2011; Neary and others 2005). U.S. Forest

Service (USFS) Burned Area Emergency Response (BAER) assessments were completed forthe burned area.The soil burn severity map

shows 12% of the Horseshoe 2 Fire burned at high severity and 30%

at moderate severity (fig. 1).TheBAERteam conducted a hydrologic

analysis of post-burn conditions using a 5-year return interval storm

with an intensity of 12.5 mm/hr (0.5 in/hr), and a geologic analysis

using burn severity, basin morphometrics, and soil characteristics to

assessthepotentialforpost-firedebrisflows.These analysesindicated

that post-fire peak discharges might increase from 2-15 times pre-fire

conditions, and several watersheds had a high potential for debris

flows (USDA Forest Service 2011).

	 Moody and Martin (2009) compared post-fire sediment yields

acrossthewesternUnitedStateswith30-minute rainfallintensityfrom

10 rainfall regimes. They found that generally channels contributed

~75% of the post-fire sediment yield while hillslopes contributed

~25%, although most of the runoff came from hillslopes(Moody and

Martin 2009). Understanding the sources of sediment from burned

areas is important for assessing post-fire geomorphic responses of

basins and the effectiveness of post-fire mitigation measures, both of

which informresourcemanagement decisions(Robichaud and others

2000). To understand sediment sources, baseline conditions must

be established. Due to the short window of opportunity in Arizona

between fire containment and the first monsoon storms, these data

can be difficult to acquire. Thisstudy established post-fire, pre-storm

conditions using aerial photographs flown immediately after the fire.

Channel and hillslope conditionsweremonitoredthrough the summer

to document basin responses to significant rainfall. The objectives of

this study are to monitor, document, and quantify post-fire erosion

from channels and hillslopes, on a broad scale, in the Chiricahua

Mountains following an approach used after the 2010 Schultz Fire in

northern Arizona (Youberg and others 2011). The goals of this study

are to document burned basin responses (floods vs debris flows) to

significant rainfall, and the locations of post-fire erosion (channels vs

hillslopes)to understand how post-fire erosion and deposition change

over time, and how those changes influence post-fire flooding and

sedimentation or recovery. The objectives of this paper are to report

on the current status of the project and to discuss some of the basin

responses from rainfall during the 2011 monsoon.

Background

	 Following the 2010 Schultz Fire near Flagstaff, Arizona, debrisflow and flood deposits were identified and mapped to assess basin

response to rainfall. Limited channel measurements were collected,

including an evaluation of available sediment in one small sub-basin

following the 2010 monsoon (Carroll 2011), and a single series of
358 USDA Forest Service Proceedings RMRS-P-67. 2013

Youberg, Neary, Koestner, and Koestner Post-Wildfire Erosion in the Chiricahua Mountains

cross-section surveys used to estimate peak flows from a productive

runoff event across the burn area (Koestner and others, in press). No

hillslope measurements were taken. Thus, to assess post-fire erosion

following the Schultz Fire, aerial photography (scale 1:12 k) was

flown after monsoon to capture initial post-fire erosion. The aerial

imagery provided both a visualrecord of erosion and ameansto create

high-resolution (15 cm - 1 m) digital elevation models (DEMs) from

the digital data using photogrammetric analysis. This method is also

being utilized on the Horseshoe 2 Fire to monitor and assess post-fire

erosion via a collaborative effort between the CNF, the U.S. Forest

Service Southwestern Regional Office (RO), the Rocky Mountain

Research Station, and the Arizona Geological Survey.

	 Extensive erosion following the Horseshoe 2 Fire was anticipated,

given severe erosion previously observed after the 1994 Rattlesnake

Fire. Tree-ring data show that the Chiricahua Mountains were burned

by widespread fires prior to 1890 and numeroussmall fires after 1890

(Swetnam 2005). Due to fire suppression and livestock grazing, the

fire regime changed during the 20th century (Swetnam 2005). The

1994 Rattlesnake Fire, which burned ~11,110 ha (27,500 ac) along

the crest of the range (MTBS 2012), is considered one of the earliest large fires of the current fire regime. Approximately 21% of the

total area was burned at high severity and 12% at moderate severity

(CNF files). The monsoon rains following the 1994 Rattlesnake Fire

caused significant post-fire erosion. Rucker Lake, at the south end

of the range, had high sediment input prior to the fire and required

frequent dredging to keep the lake open (Lefevre and Neary 1999).

During a post-fire monsoon storm multiple debris flows initiated in

upper Rucker Canyon, and coarse sediment filled Rucker Lake. This

event was unprecedented, delivering approximately three times the

pre-fire sediment load to Rucker Lake (Lefevre and Neary 1999).

Some ofthe most impressive erosion occurred in upperWardCanyon

on the steep western flank of Chiricahua Peak, where a 10-m gully

formed. By 2004, the gully bottom had filled in approximately 4-5

m and widened 2-3 m through mass wasting processes (P. Pearthree,

personal communication).

Study Area

	 The study area was selected using several criteria including burn

severity, accessibility, proximity to communities, and the potential

for significant post-fire erosion. The study area encompasses several

canyons in the northern part of the burn area including Pinery, North

Pinery, Pine andEastWhitetailCanyons, andEastTurkeyCreek (fig. 1).

Pinery Canyon and East Turkey Creek are dissected by the 42 Road,

the only major east-west road through the mountain range, and East

TurkeyCreekcontainsthe small communityofParadise.NorthPinery

and East Whitetail both have extensive areas of high burn severity

(>~70%), and East Whitetail also contains a small community. Pine

Canyon was partially burned in the Rattlesnake Fire with significant

post-fire erosion. A church camp is also located within Pine Canyon

and extensive patches on the steep northern slopes above the camp

and in the upper canyon were burned at high severity.

Figure 1—Location map of the Horseshoe 2 Fire with burn severity. Lower left inset panel shows the location of the Horseshoe 2 Fire in southeastern 

Arizona. The map of the fire shows soil burn severity map, location of rain gages, and study area outlined in box. The right image shows the study area.
USDA Forest Service Proceedings RMRS-P-67. 2013 359

Post-Wildfire Erosion in the Chiricahua Mountains Youberg, Neary, Koestner, and Koestner

Methods

Aerial Photography

	 The CNF with assistance from the RO GIS/Photogrammetry

Program had aerial photography flown in the northern portion of the

fire with the goal of capturing post-fire, pre-storm conditions of the

study watersheds. The air photos were flown on July 1, 8, 14 and 15

at a scale of 1:12000. In addition to these photos, the RO collected

aerial photographs in August and September as part of their ~5-year

cyclicalresource aerial photography program.These resource photos

are digital with a resolution of 25 cm (10 in).

Precipitation Data

	 Thefirstraingage (CHIRI_B2)withinthe burnedareawasinstalled

on the east side of Buena Vista Peak, near the 42D road, on July 1 by

the University of Arizona/Biosphere 2. A temporary weather station

with a recording gage (KC2CPZ-1) was installed on July 26 on top

of Buena Vista Peak at the Barfoot Lookout location by the National

Weather Service (NWS). A second temporary weather station (KC2CPZ -2) outside our study area was installed by the NWS within

ChiricahuaNationalMonument onAugust 3.Two permanentALERT

gages were installed by the Arizona Department of Water Resources

(ADWR) on August 26. One is co-located on top of Buena Vista

Peak and will remain after the temporary weatherstation isremoved.

The second (King of Lead) is at the headwaters of East Whitetail

Canyon, also outside of our study area near the King of Lead mine.

In October, the USGS installed a stream warning system and a rain

gage in Chiricahua National Monument.

Field Monitoring and Other Data

	 Visits to key study areas were conducted as soon as possible following significantstorm events during the monsoon of 2011. Smaller

stormsweremonitoredremotelyandbasinresponsesdeterminedfrom

reports,videos andphotographstakenbyBAERImplementationteam

members,localresidents anddistrictpersonnel.Fieldvisits coveredas

much of the study area as possible, focusing on those areas that had

received significant rainfall, to document and photograph hillslope

erosion and channel condition. Channel deposits were classified,

when possible, as either flood flow or debris flow. In Pinery Canyon

tributary, channel material was deposited on the 42 road, providing

very good exposures of deposits for flow-type classification. In East

Turkey Creek, however, the channels above the road have fairly deep

basins. Often during field visits these deposits were either entirely

buried leaving no cross-sectional exposures to characterize deposits

or,more likely, the road crewhad already cleared the road and cleaned

out the basinsleaving no undisturbed depositsto evaluate. Numerous

photos and reports from these locations during the largest storms

provide some information regarding basin responses.

Results

	 The first few rains that fell on the study watersheds were of moderate intensity and duration. They moved some sediment and ash

in and through the channels, resulting mainly in nuisance flooding.

Hillslope rillingwas noted in the study area following a June 29 storm.

The first major rainfall occurred on July 11 (fig. 2). The CHIRI rain

gage recorded 54.4 mm (2.14 in) in 1 hour, 38 minutes, with a peak

30-minute intensity of 35.6 mm (1.4 in) (Steve DeLong, unpublished

data). The online NOAAAtlas 14 (http://dipper.nws.noaa.gov/hdsc/

pfds/) classifies this 30-minute intensity as a 2- to 5-year frequency

storm. On the east side of the range, where the rain gage sits, floods

were observed in the East Turkey Creek drainage. On the west side

of the range, debris flows occurred in several tributaries of Pinery

Canyon, depositing debris on the 42 Road, and in tributaries to Pine

Canyon above the church camp. Numerous storms throughout the

summer produced large enough floodsto generate documentation via

reports, photos and videos. Additional debris-flow producing storms

occurred on July 28, with a debris flow noted in a tributary to East

Whitetail Canyon, and onAugust 15 with evidence of debris flowsin

Pine Canyon and East Turkey Creek (fig. 2). This storm significantly

impacted the church camp with debris flows depositing just upstream

and flooding throughout the camp. No debris flows were observed

in North Pinery Canyon. Some landslides had been observed in this

canyonduringtheBAERassessment(Youberg2011)butnoadditional

movement was noted over the monitoring period.

	 The aerial photographs flown in July to capture post-fire, pre-storm

watershed conditions were actually flown over 4 days. The east side

of the area was flown on July 1 and 8, prior to any significant rainfall,

and the west side was flown on July 14 and 15 following the first

debris-flow producing storm. There is a slight overlap in two flight

lines along the ridge crest that capturesthe post-fire, pre-storm condition and the watershed condition after the first major storm. This is

particularly evident at the top of Pine Canyon in an area that had been

burned by the Rattlesnake Fire. The pre-fire imagery shows limited

vegetation with large areas of grass cover. The soil burn severity map

showslow to moderate burn severity in this area, while the vegetation

burn-severityshowshighburnseverity.The removalofvegetationand

the intensity of the rainfall were sufficient to cause dense rilling and

to re-incise gullies that had begun to fill in after the post-Rattlesnake

Fire erosion.

Discussion and Ongoing Work

	 During the summer of 2011 basin responses to rainfall were documented for five canyons burned by the Horseshoe 2 Fire. Debris flows

occurred in Pinery and Pine Canyons on July 11, in East Whitetail

Figure 2—Rainfall storm totals with dates of debris-flow producing storms. 

The Univeristy of Arizona’s rain gage, CHIRI_B2, was installed on July 1st

and is located along the 42D Road. The NWS weather station, KC2CPZ-1, 

is located on top of Buena Vista Pevak and was installed on July 26th. 
360 USDA Forest Service Proceedings RMRS-P-67. 2013

Youberg, Neary, Koestner, and Koestner Post-Wildfire Erosion in the Chiricahua Mountains

on July 26, and in East Turkey Creek and Pine Canyon on August

15. Channels affected by debris flows are being significantly altered

as pulses of sediment are transported downstream with subsequent

flows. As a result, channel cross-section areas are changing with

each flow event. All other storms throughout the summer produced

flood flows. Hillslope rilling, however, began with the first few minor

storms in areas of high soil burn severity. Subsequent, more intense

rains contributed to additional rilling and erosion.

	 A review of the channel conditions in Rucker, Ward and upper

Pine Canyonsshow channels and hillslopesthat had not recovered by

2011.Intheupper canyons,priortothe summerstorms,oversteepened

channel banks of gullies had laid back and the bottoms had started to

fill in (fig. 3). Following the summer storms, however, evidence of

re-incised gullies and new hillslope rills and gullies are present in all

ofthese canyons(fig.3).InlowerRuckerCanyonthepost-Rattlesnake

gravel/cobble channel bed had re-established a low-flow channel in

some areas but not in others. As new sediment moves through the

system these low-flow channels will once again fill with coarse sediment. Long-term disturbances such as these should be expected in

canyons burned at moderate to high burn severity by the Horseshoe

2 Fire.

	 It was difficult to assess rainfall intensity associated with different

flow events throughout the study area. There were only two gauges

for the first half of the summer and they were essentially co-located.

Rainfall intensity can sometimes be assessed for larger areas by

comparing gage data with radar data. This has been done for the 2010

Schultz Fire and is being done for the 2011 Monument Fire in the

HuachucaMountains(Youberg,unpublisheddata).Unfortunately,the

Chiricahua Mountains are far from the surrounding radar stations,

consequently, the beam is too far above the top of the mountain and

often does not reliably capture rainfall on the mountains. Thus, no

definitive conclusions can be drawn regarding rainfall intensity and

return frequency for any ofthe study basins exceptforthe headwaters

of Pinery Canyon and a few tributaries to East Turkey Creek, which

contain the CHIRI_B2 and KC2CPZ-1 gages. The largest storm

recorded during the summer had a 2- to 5-yr, 30-min return interval.

	 This project is ongoing as funding allows.Aerial photographs will

again be flown for the study area prior to the 2012 monsoon. This

imagery will capture the first full year of erosion with snapshots in

the August/September timeframe when the resource photos were

flown. DEMs have not yet been developed but coarse-scale mapping

of the deposits has occurred. Although there are several limitations

to thisstudy, including detailed hillslope and channel measurements,

results from this study will help provide a broad understanding of

post-fire erosion in the Chiricahua Mountains. Data from other studies (Gibson and others this volume) will be combined with the aerial

imagery data to provide a clearer picture of sediment sources. In the

long-term, these data will help inform resource decisions regarding

post-fire land management.

Figure 3—Ward Canyon in the headwaters of the West Turkey Creek drainage. The top photo is a NAIP 2010 image 

showing the post-Rattlesnake burned area and erosion. After the 1994 fire, post-fire runoff carved a ~10-m deep 

gully. By 2010 the gully banks had laid back and the gully was partially filled. Bottom photo was taken during 

late summer as part of the 2011 USFS resource photo acquisition program. This photo shows post-Horseshoe 2 

Fire erosion from mid August storms with new rills (yellow arrow) and gullies (orange arrow), and re-incision and 

headward migration of previously existing gullies (white arrows).
USDA Forest Service Proceedings RMRS-P-67. 2013 361

Post-Wildfire Erosion in the Chiricahua Mountains Youberg, Neary, Koestner, and Koestner

Acknowledgments

	 We thank the Arizona Geological Survey, the Rocky Mountain

Research Station, the Coronado National Forest, especially Jennifer

Ruyle, Bob Lefevre, and Salek Shafiqullah, and the Southwestern

RegionalGIS/PhotogrammetryProgram,particularlyCandaceBogart

and Bart Mathews, for supporting this project.

References

Carroll, M. 2010. Movement of channel-borne sediments in the 2010 Schultz

Fire burn area. A Thesis Submitted in Partial Fulfillment Of the RequirementsfortheDegreeofMasterofScience InEngineering,NorthernArizona

University, May 2011. 60 p.

DeBano, L.F.; Neary, D.G.; Ffolliott, P.F. 1998. Fire’s Effects on Ecosystems.

John Wiley & Sons, New York. 333 p.

Jenkins, S.E., Sieg, C.H., Anderson, D.E., Kaufman, D.S., and Pearthree,

P.A., 2011 Late Holocene geomorphic record of fire in ponderosa pine

and mixed-conifer forests, Kendrick Mountain, northern Arizona, USA:

International Journal of Wildland Fire. 20(1): 125-141.

Koestner, K.A., Koestner, P.E., Neary, D.G. (in press). Estimating post-fire

peak flows following the 2010 Schultz Fire, Coconino National Forest.

Proceedings of the Arizona-Nevada Academy of Sciences 2011 Annual

Meeting. 3 p.

Lefevre, R.E., and Neary, D.G.. 1999. Rucker Lake: A history of recent conditions affecting a southeastern Arizona watershed. Hydrology and Water

Resources in Arizona and the Southwest. 29: 23-30.

Moody, J.A., and Martin, D.A., 2009. Synthesis of sediment yields after

wildland fire in different rainfall regimes in the western United States:

International Journal of Wildland Fire. 18:96-115.

MTBS. 2012. 1994 Rattlesnake Fire, Arizona, Monitoring Trends in Burn

Severity Data Access Center(http://www.mtbs.gov/dataaccess.html).

Neary, D.G.;Ryan, K.C.; DeBano, L.F., eds. 2005 (Revised 2008). Fire effects

on soil and water. Gen. Tech. Rep. RMRS-GTR-42, Volume 4. Fort Collins, CO: USDAForest Service, Rocky Mountain Research Station. 250 p.

Robichaud, P.R.; Beyers,J.L.; Neary, D.G. 2000. Evaluating the effectiveness

of post-fire rehabilitation treatments.Gen.Tech.Rep.RMRS-GTR-63.Fort

Collins,CO:USDAForestService,RockyMountainResearchStation.85p.

Swetnam,T.W. 2005. Fire historiesfrom pine-dominant forest in the Madrean

Archipelago, Proceedings RMRS-P-36. Fort Collins, CO: USDA Forest

Service, Rocky Mountain Research Station: 35-43.

U.S. Forest Service. 2011. Burned Area Report, Horseshoe 2 Burned Area

Emergency Recovery Assessment, Coronado National Forest, Douglas

Ranger District: 28.

Youberg, A. 2011. Horseshoe 2 Fire, Burned Area Emergency Response

(BAER) Assessment, Geologic Hazard Assessement: Coronado National

Forest, Douglas Ranger District: 15.

Youberg, A., Koestner, K.A., Schiefer, E., and Neary, D.G. 2011. Hazard

analysisfor post-fire debris-flowpotentialinArizona.In: 2011FallMeeting

of the AGU, San Francisco, CA, Abstract EP31B-0821.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
362 USDA Forest Service Proceedings RMRS-P-67. 2013

Youberg, Neary, Koestner, and Koestner Post-Wildfire Erosion in the Chiricahua Mountains
USDA Forest Service Proceedings RMRS-P-67. 2013 363

Tools for Management
364 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 365

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Mapping and Assessing the Environmental Impacts of 

Border Tactical Infrastructure in the Sky Island Region

Caroline Patrick-Birdwell

School of Geography and Development, University of Arizona, Tucson, Arizona

Sergio Avila-Villegas, Jenny Neeley, and Louise Misztal

Sky Island Alliance, Tucson, Arizona

Abstract—In this project we mapped the different types of border barriers, identified impacts of border 

infrastructure on public and private lands and conducted spatial analyses within the approximately 200 

miles of international border in the Sky Island region. The Sky Island region, bisected by the U.S.-Mexico 

border, is critically important for its biodiversity and continental connectivity. Only on the Arizona side is the 

border lined with protected areas such as San Bernardino and Buenos Aires National Wildlife Refuges, San 

Pedro Riparian National Conservation Area, Coronado National Memorial, and several Wilderness Areas. In 

recent years the construction of border barriers, walls and other tactical infrastructure built to deter human 

and vehicle traffic across the border has had substantial impacts on ecological processes and created an 

impermeable boundary with effects that reach far beyond the footprint of the border wall. Due to the Department of Homeland Security’s waiver of all legal requirements along significant portions of the U.S.-Mexico 

border, construction of border infrastructure proceeded without the requisite environmental impact studies 

or input from public land managers or private landowners. With the use of Geographic Information Systems 

we analyzed and graphically compiled results from over-flights along the border generating photographic 

data of the current state and effects of infrastructure, field visits to obtain complementary information and 

ground-truth aerial photographs, and observations of the direct effects of barriers to wildlife movements and 

the flow of ecological and hydrologic processes.

	 From a cartographic perspective, the United States-Mexico border

is often depicted as a fairly straight line, drawn to differentiate the

countries to the north and south. In ecological terms, the line is far

from simple. Hundreds of miles of walls, fences, and other barriers

havebeeninstalledthat are impactingwildlifemovement,watersheds,

and ecosystem health and stability. In response to these construction

projects,SkyIslandAlliance tookthe leadindevelopingageodatabase

and map of the border in the Sky Island region as the first steps to

understand the effects of infrastructure installation and to make this

information available to the public.

	 OnOctober 26, 2006,PresidentGeorgeW.Bush signed into lawthe

Secure Fence Act. The stated goals for this act included construction

of hundreds of miles of additionalfencing along our Southern border;

authorization of additional vehicle barriers, checkpoints, and lighting to help prevent people from entering the United States illegally;

and authorization for the Department of Homeland Security (DHS)

to increase the use of advanced technology like cameras, satellites,

and unmanned aerial vehicles to reinforce our infrastructure at the

border. Since the passage of theAct, the installation and maintenance

of border tactical infrastructure (TI) has become a central component

of DHS border activities (Whitehouse 2006).

	 In orderto accomplishTIinstallation,theDepartment ofHomeland

Security (DHS) relied on provisions from earlier legislation known

as the Real ID act of 2005, which gave DHS authority to waive all

legal requirements for building border infrastructure, and proceeded

with TI construction without having to go through an environmental

impact analysis required by NEPA, the National Environmental

PolicyAct (Ganster 2007). While this has allowed for expedient wall

construction along the US/Mexico border, it has created considerable concern among the public and in conservation communities

over the short and long-term impacts on ecosystems, endangered

and threatened species, wildlife corridors, and watersheds. As figure

1 highlights, approximately 200 miles of TI bisects the unique and

fragile Sky Island bioregion. Sky Island Alliance (located in Tucson,

Arizona) has been engaged in conservation advocacy of the border

region since 1991. In 2010, Sky Island Alliance (SIA) photographed

the extent of existing TI in the Sky Island region, and determined

that a map detailing the types of barriers and other infrastructure

was needed to understand real and potential threats to the area, as

well asfor conservation advocacy purposes. This project represents a

culmination of those efforts. The map produced from this project will

be published in print and online, and will be used as a tool for future

conservation activities and advocacy.Itis also thefirst comprehensive

TI infrastructure map of the region available to the public.
366 USDA Forest Service Proceedings RMRS-P-67. 2013

Patrick-Birdwell and others Mapping and Assessing the Environmental Impacts of Border Tactical Infrastructure . . 

The Sky Islands: An Epicenter of 

Biodiversity

	 The Sky Island region of southeastern Arizona and southwestern

New Mexico, and adjacent states in northwest Mexico (Sonora and

Chihuahua), is characterized by forested mountains surrounded by

intervening desert and grassland. This region hosts a high diversity

of plants, animals and habitats. The region sits at the confluence of

four major ecosystems: the Colorado Plateau, the Sierra Madre Occidental, and theSonoran andChihuahuan deserts.Because the region

encompasses North America’s two largest deserts and is a bridge

between its two major mountain ranges, it is critically important to

continental connectivity. Madrean pine-oak woodland, an important

characteristic of Sky Islands, was designated a biological hotspot by

Conservation International in 2005. The high number of federally

protected areas along the international border—including national

wildlife refuges, wilderness areas, national monuments, riparian

national conservation areas, and national parks in Mexico and the

United States—testify to the region’s unique richness (SIA 2011).

Project Background

	 Creating a border infrastructure map per the needs of SIArequired

a multi-faceted process and an understanding of environmental and

ecological factors of the region. In addition, we researched the political history of the area and how policy was developed to create the

infrastructure, andtheusageoflandforwall construction,bridges, and

easements. Because there was not a unifying source for these varied

data needs, information was amassed from many sources, including

printed publications, government websites, non-profit organizations

includingSIA,theNorthern JaguarProject andDefenders ofWildlife,

border fly-over missions, and field collections. SIA began collecting

data and monitoring the region in 2008, after the passage of the Real

ID Act and the Secure Fence Act and the ensuing construction of the

TI. Since then, it has used staff and volunteers to document wildlife

corridors and monitor erosion and habitat degradation in the region

as a result of construction and increased human activity.

	 In 2010, SIA conducted two photographic fly-over missions to

capture the extentofthe tacticalinfrastructure intheSkyIslandregion.

The flights documented infrastructure from Sasabe, Arizona, to the

easternedgeofNewMexico’s “bootheel.”The aircraft, equippedwith

Figure 1—Project study area.
USDA Forest Service Proceedings RMRS-P-67. 2013 367

Mapping and Assessing the Environmental Impacts of Border Tactical Infrastructure . . Radke

two photographers and a note-taker/navigator flew the extent of the

region (approximately 200 miles) from west to east, on the south side

ofthe border.Themajority ofthe imageswere taken fromthe perspective of looking north, with images also captured from the southern

side of the plane to assist in geo-referencing the photos based on the

area’s topography. As figure 2 shows, the photos from these flyovers

provided an essential visual record of the extent of the TI. The note

taker recorded on a topographic map the changesin the infrastructure

that could be detected from the air. Although this process produced

some excellent documentation of the border infrastructure, it took

considerable time to geo-reference the photosto actual coordinates on

the ground. This was accomplished by comparing the images to the

topographic maps used during the flights and verifying the locations

with Google Earth.

	 After compiling a profile of the TI through the aerial images, we

realized that ground-truthing data collection would be needed to

better understand the types of infrastructure used in areas that were

either not clearly captured in imagery and/or in important conservation areas. In July and August of 2011 we visited several significant

areas along the border, including the Buenos Aires National Wildlife

Refuge (BANWR), the San Pedro Riparian National Conservation

Area (SPRNCA), and the San Bernardino National Wildlife Refuge.

As an example of the importance of these areas ecologically, the San

Pedro is one of the last free flowing rivers in the desert southwest. It

is also a global hotspot for bird and mammal diversity and is certainly

one of the most significant wildlife corridors connecting the United

States and Mexican nations. It is also important to note that the wall

construction in the SPRNCA boundaries represents the first time

such activities had occurred on public land that had been federally

designated as a protected area (Segee and Cordova 2009).

Infrastructure Classifications

	 In the Sky Islands region, most of the existing TI falls into two

main categories: pedestrian fencing and vehicle barriers. There are

sub-categories with style/material differences within the two main

categories.Table 1 outlinesthe different types ofTI documented over

the course ofthis project.Based on our observations, the construction

goals for using different styles depended on topographical features

of the area. For example, vehicle barriers were employed in certain

riparian areas and otherlocationsthat the terrain presented challenges

to constructing pedestrian barriers.

Cartography Goals

	 Once we had assembled our data, we began the task of creating

a map that depicted the area and its relationship to the constructed

tactical infrastructure. The goal for this map wasto provide viewers a

way of seeing this area as an ecological region that transcends political borders, and that the border infrastructure construction bisectsthe

region and itsimportant ecological processes.To accomplish this goal

we did our best to downplay the state and national borders, focusing

instead on the tactical infrastructure as the dividing line (fig. 3). We

also made sure to include important wildlife preserves in Sonora and

Chihuahua. Far too frequently, maps of the U.S./Mexico border only

provide information on the northern side of the border. We did our

best to rectify that approach with the information we had available.

Tactical Infrastructure Footprint

	 Once the data was compiled and assembled into a comprehensive

map, it became possible to establish some baseline footprint data that

can be used for conservation advocacy.

	 The infrastructure information compiled forthismap yielded some

interesting footprint statistics. Table 2 provides statistics on barrier

miles in protected areas. In looking at these figures it is important to

consider the following facts: (1)The mileage listed is an approximate

basedonGIStools,aerialphotography, andgroundtruthingefforts;and

(2) construction is continuing on the border, and this data represents

a snapshot of the situation as of July, 2011. In addition to the linear

miles of federal land that the tactical infrastructure now occupies,

hundreds, if not thousands, of acres of land has also been allocated

and used by Custom and Border Protection (CBP) for infrastructure

purposes under the auspices of the “Roosevelt Reservation.” This

little-known piece of legislation allows for a “public highway” to

run along the U.S./Mexico border. CBP has used this to build access

roads and other TI, and in several locations, have extended the easement well beyond the 60 feet to 120 feet or more. Environmental

stewardship plans produced by CBP estimate a loss of 552 acres

in vegetation communities from the barrier construction activities

(Homeland Security 2008). Conservative estimates from this project

place that figure at more than 900 acres in lost habitat, with over 500

acres on federal land.

	 Basedoninfrastructuremeasurements,thedata collectedduringthis

project highlight the impact of the TI in federal areas. It is important

to remember that this infrastructure, with an average construction

cost of $4.1 million per mile, was placed on U.S. federal land, in

Figure 2—Aerial view south to north of the San Pedro River area bisected by the international border (non-contiguous photos). On the Arizona 

side, the San Pedro is a riparian National Conservation Area. In addition to 6-ft high walls, a wide border road and numerous access roads 

fragment the land (left); Normandy-style vehicle barriers cross the San Pedro River bed (center); washes and arroyos are paved to protect the 

road, but water courses are blocked by pedestrian fences (right) (photos courtesy of Sky Island Alliance, Paul Condon, and LightHawk Inc.).
368 USDA Forest Service Proceedings RMRS-P-67. 2013

Patrick-Birdwell and others Mapping and Assessing the Environmental Impacts of Border Tactical Infrastructure . . 

Table 1—Barrier classifications and characteristics (photos by Sergio Avila-Villegas and Caroline Patrick-Birdwell, 

2011).

Barrier photo Barrier type Approximate 

height Materials Wildlife friendly Water friendly

Pedestrian/ 15 to 18 feet Steel, recycled 

helicopter 

landing mats

No No

Pedestrian/bollard 15 to 18 feet grout-filled steel No No

Pedestrian/bollard 15 to 18 feet grout-filled steel No No

Pedestrian/Waterdrainage Gate15-18 feet grout-filled steel 

posts with gate 

housing

No Variable; 

gates are only 

water friendly 

when lifted in 

advance of a 

storm event

Pedestrian/mesh 15-18 feet Steel with steel/

wire mesh

No No

Vehicle/Normandy 4-6 feet Non-reflective 

steel

Variable Variable

Vehicle/Post and 

Rail

4-5 feet Non-reflective 

steel

Variable Variable

highly senstive ecological hot spots, without benefit of environmental analysis or environmental impact statements. The next phase for

this project is to analyze the data more thoroughly and then use it to

advocate for enlightened border security strategies that consider the

environmental implications of those efforts.

	 Given the size and significance of the border section of the Sky

Island bioregion, this project represents a starting point for further

research. With the infrastructure in place in 62% of the region’s international border, biologists are already seeing signs ofinterruptions

in wildlife corridors as well as erosion and other damage to fragile

riparian ecosystems. The map and data produced for this project are

important first steps for understanding the threats of the border wall

on the environmental health of the region.
USDA Forest Service Proceedings RMRS-P-67. 2013 369

Mapping and Assessing the Environmental Impacts of Border Tactical Infrastructure . . Radke

Figure 3—Infrastructure map (Cartography by Caroline Patrick-Birdwell and Louise Misztal, 2012).

Table 2—Tactical infrastructure mileage/area approximations.

Vehicle Pedestrian Total border Percent covered Total barrier

barrier barrier length w/barrier (study area) Study area

Designated wilderness 4.18 0

Wildlife refuges 3.06 5.44 8.5 100

Conservation areas 1.43 0.3 1.75 100

U.S. Forest Service 17.5 5.65 73 32

National Park Service 0.25 1.8 3.5 58

Other (public/private) 32.96 57.91

Total (all land types) 55.2 71.1 62 126.3 203

Acres used in 60 ft. barrier corridors

Federal land 516

Total 918

CBP predicted vegetation loss 552

References

Ganster, Paul. 2007. Environmental Protection and U.S.-Mexican Border

Security: The Border Fence Issue in Context. In: Ana Córdova and Carlos

de la Parra. A barrier to our shared environment: the border fence between

the United States and Mexico. Tlalpan: Secretariat of Environment and

Natural Resurces: 31-42.

Romero, Fernando. 2008. Hyper border: The contemporary U.S.-Mexico

border and its future. New York: Princeton Architectural Press.

Segee,Brian, andAnaCórdova.2009.ConservationimplicationsofrecentU.S.

border security legislation. In: Laura Lopez-Hoffman, Emily McGovern,

Robert Varady, and Karl Flessa. Conservation of shared environments:

Learning from the United States and Mexico. Tucson: University of Arizona Press: 241-255.

SkyIslandAlliance.[n.d.]skyislandalliance.org(accessedNovember1,2011).

U.S. Department of Homeland Security. 2008.Environmentalstewardship pal

for construction, operation, and maintenance of vehicle fence and related

tactical infrastructure. U.S. Border Patrol, Tucson Sector, Arizona.

White House Archives. [n.d.] whitehouse.archives.gov/news/releases/2006/10/20061026-1.html (accessed October 15, 2011).

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
370 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Using Remote Sensing to Monitor Post-fire Watershed 

Recovery as a Tool for Management 

Jess Clark

USDA Forest Service, Remote Sensing Applications Center, Salt Lake City, Utah

Marc Stamer

USDA Forest Service, San Bernardino National Forest, Fawnskin, California

Kevin Cooper

USDA Forest Service, Los Padres National Forest, Santa Maria, California

Carolyn Napper

USDA Forest Service, Shasta-Trinity National Forest, Mt. Shasta, California

Terri Hogue and Alicia Kinoshita

University of California Los Angeles, Los Angeles, California

Abstract—Post-fire watershed recovery is influenced by numerous variables but one of the most important 

factors is the rate of re-establishment of vegetative cover. Burned Area Emergency Response (BAER) teams, 

along with other agencies (Natural Resource Conservation Service, state, counties, cities, etc.), prescribe 

temporary post-fire mitigation treatments based on expected post-fire responses of watersheds to fire-caused 

damage and based on threats to life, property, and resources associated with watershed damage. The objective of this project was to develop tools to more accurately assess the rate of vegetation regeneration after 

wildfire that will help managers decide if there is a continued need for mitigation measures. We develop a 

decision support tool to aid land managers and emergency response personnel in their evaluation of continued risks posed by recovering watersheds.

Introduction 

	 In the last decade wildfires have increased in both size and severity

(Westerling and others 2006; Snider and others 2007; Westerling and

Bryant 2008). The effects of wildfires potentially impact property

within and adjacent to burn areas several years post-fire via their

adverse effects on watersheds (Jung and others 2009; Kinoshita and

Hogue 2011).As a result, land managers are constantly seeking ways

to assess post-fire watershed responses and their potential impacts

(e.g., flooding, erosion, sedimentation) to federal and non-federal

lands. After a significant wildfire, federal land managers restore and

rehabilitate burn areas in three ways: suppression repair, emergency

response (within 1-3 years after fire), and long-term rehabilitation

(3-10 years after fire). Treatments implemented as part of Burned

Area Emergency Response (BAER) are designed to reduce post-fire

watershed impactsto life, property, and natural and culturalresources.

These treatments, however, are viewed as temporary and are monitored, maintained or even retreated for up to three years after fire

containment (U.S. Department of Agriculture Forest Service 2012).

	 The type and duration of treatments in the aftermath of a fire is

directly related to the degree of watershed impairment. A primary

rehabilitation objective is restoration of vegetation cover to stabilize

slopes and erodible soils, and reduce overland flow. Hillslope erosion

is inversely related to plant cover and is considered minimal when

plant coveris ≥60%(Noble 1965;Orr 1970).For example,treatments

suchasstrawmulchingaremost effective atreducingrilldevelopment

and sediment transport when cover is >60% (Robichaud and others

2010). Cover is especially critical in the first year after the fire when

the risk of erosion is highest (DeBano and others 1998).

	 Remote sensing and geospatial technologies are frequently used by

land managers to guide resource management decisions. Moderate

resolution satellite imagery, most notably Landsat, has proven to be a

valuable informationsource formappingfire severityandits effectson

vegetation (Clark and Bobbe 2006) and monitoring post-fire vegetation recovery (Diaz-Delgado and others 1998; Clark and Kuyumjian

2006;Wittenberg and others 2007).Threewell-knownLandsat image

derivatives used in fire effects mapping are the normalized difference

vegetation index (NDVI) (Tucker 1979),

NDVI = (B4 − B3) / (B4 + B3),

the enhanced vegetation index (EVI) (Huete and others 2002),

EVI = 2.5 * ((B4 – B3) / (B4 + 6 * B3 – 7.5 * B1 + 1)),

andthenormalizedburnratio(NBR)(LòpezGarciaandCaselles1991),

NBR = (B4 – B7) / (B4 + B7).
USDA Forest Service Proceedings RMRS-P-67. 2013 371

Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management Clark and others

where B is the top of atmosphere (TOA) reflectance of the specified

Landsatband.Vegetationindices are commonremote sensingproducts

that are used to highlight a particular vegetation property.

	 While both NDVI and EVI measure the photosynthetic activity

and chlorophyll content in plants, the EVI was developed to correct

for soil background signals as well as atmospheric influences (Huete

and others 2002). The NBR is influenced by chlorophyll activity,

but it is influenced more by moisture content of vegetation and soils

(presence or absence of dry, bare soil and healthy vegetation) (Key

and Benson 2006).

	 BAER teams, and those charged with long-term monitoring of

watershed recovery, utilize these vegetation indices to ensure that

stabilization plans remain on schedule. Since severely burned watersheds often require > 3 years(limit of BAER program’sstewardship)

to return to pre-fire conditions, managers need a method to monitor

their condition (U.S. General Accounting Office 2003). Currently,

however, managers lack a cost-effective way to evaluate watershed

recovery that could help decide when to end temporary protective

treatments. Thus, the objective of this project wasto use remote sensing to monitor post-fire vegetation recovery and to develop a decision

support tool that assists managers in their assessment of risks posed

by the watershed to resource values.

Methods

	 The studyarea includedsixfiresthatburnedbetween2003and2010

and included a variety of elevations, soil burn severities, and stages

of recovery (table 1, fig. 1). The six fires burned in predominately

chaparral and mixed conifer cover types.

	 Data onpercentvegetative coverwere collectedatseverallocations

withintheburnareas.Weutilizedpole-mastphotography,thatis,downlooking camera attached to the top of a telescopingmonopod (fig. 2)to

measure ground cover from heights ranging from 25-30 feet (Gilbert

and others 2009; Smith and others 2000; Vanha-Majamaa and others

2000). Plots were chosen for ease of access, internal homogeneity,

and soil burn severity. We took between four and ten photos per plot

depending on the size ofthe homogeneous patch. Photos were spaced

approximately 15meters apart and interpretedwith custom-builttools

within Esri’s ArcGIS ArcMap© using a random dot grid sampling of

600 points per plot. Plant cover was termed either present or absent

at each point and cover was based only on living plants (both healthy

and dormant/senesced vegetation). Plots were given a single percent

cover value using the count of covered points.

	 A remote sensing data analysis was conducted to relate the photointerpreted field plots to the maximum greenness observed from

satellite imagery during the growing season. To accomplish this, we

compiled all available cloud-free Landsat imagery acquired for each

growing season after the fire. For each Landsat image, we created an

NDVI, EVI, and NBR vegetation index layer. The vegetation index

data for each growing season were further analyzed on a per-pixel

basis to derive maximum observed vegetation index value during

the growing season (Sousa and others 2003). The annual maximum

vegetation index value results capture the spatial and temporal distribution of annuals and other vegetation that green up in the spring

and those whose peak green-up is later in the season.

	 Each plot intersected between 2 and 5 Landsat pixels depending on

the plotsize and orientation. The mean value of the annual maximum

vegetation index pixelsintersecting the plots were calculated for each

year. We then performed regression analysis between ground cover

derived from pole-mast photography and the mean vegetation index

value computed from the three vegetation indices for each growing

season.

Table 1—We analyzed vegetation recovery on 6 burned areas and applied our models to fires in the Madrean Archipelago.

 Soil burn severity Project 

Fire name Year burned Location Acres Elevation range percentagesa applicationb

Old 2003 San Bernardino, CA 91,281 2,000-7,600’ 24, 20, 46, 10 Sample

American River Complex 2008 Foresthill, CA 20,541 2,500-6,700’ 27, 35, 26, 12 Sample

La Brea 2009 Santa Maria, CA 89,489 1,100-5,000’ 13, 24, 53, 10 Sample

Station 2009 La Cańada, CA 160,577 2,000-6,500’ 12, 16, 62, 10 Sample

Bull 2010 Kernville, CA 16,442 2,700-7,400’ 13, 37, 49, 1 Sample

Canyon 2010 Lake Isabella, CA 9,860 2,000-6,000’ 18, 35, 43, 4 Sample

Monument 2011 Sierra Vista, AZ 32,837 4,400-9,500’ 9, 42, 39, 10 Modeled

Horseshoe2 2011 Portal, AZ 222,694 4,500-9,800’ 18, 42, 36, 4 Modeled a

 Percentage of area classified as unburned, low, moderate, and high as estimated from the Burned Area Reflectance Classification (BARC).

b

 Field data was gathered on “sample” fires; models were applied to create predicted cover maps on the “modeled” fires.

Figure 1—The six fires sampled (burn date) included Old (2003), American River Complex (2008), La Brea (2009), Station (2009), Bull (2010), 

and Canyon (2010).
372 USDA Forest Service Proceedings RMRS-P-67. 2013

Clark and others Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management 

Results

	 Regression models produced an acceptable fit for NDVI (N = 53,

R2 = 0.65, p = 0.038) and EVI (N = 53, R2 = 0.63, p = 0.019) but not

for NBR (N = 53, R2 = 0.17, p = 0.004). Fitting a simple linear model

resultedina “reasonablygood” relationshipbetweenground-observed

cover and the NDVI index.

Percent cover = 221.18*MaxNDVI–26.273

The relationship between ground cover and EVI was bestrepresented

by a polynomial model (fig. 3).

Percent cover = -455.22*MaxEVI2 + 519.99*MaxEVI – 47.508

Discussion

	 Despite the utility of NBR for burn severity mapping (Chen and

others 2011), it did not correlate well with field-measured ground

cover (fig. 3). The NBR is best suited for densely forested areas

and, in general, does not perform as well in sparsely vegetated areas

(Miller and Thode 2007). Our data show considerable confusion of

NBR values in the 0-30% observed cover range (fig. 3). This confusion is probably due to the influence of Landsat band 7 in the NBR

algorithm since similar results were not found in the NDVI or EVI

correlations, neither of which use Landsat band 7. Conversely, the

NDVI and EVI had significantly better correlationsthan NBR. These

results have operational significance in providing user flexibility to

apply multiple available remote sensing assets for post-fire recovery

monitoring. Specifically, a limited number of satellite sensors collect data in the 2.1 μm band which is necessary for generating the

NBR. However, several moderate resolution sensors collect data in

the visible/near infrared which isrequired for generating EVI/NDVI.

	 Initial results indicated correlations between EVI and NDVI with

plant cover on the six different fires sampled. Nevertheless, we had

to confine our sampling to chaparral and mixed conifer forests in

California because of project timelines and budget constraints. There

is an obvious need to continue testing in other vegetation types.

Furthermore, we sampled each fire only once which created a single

snapshot in time of the vegetation. To some extent we addressed this

problem by leveraging annual time series satellite imagery and field

photo interpretation that inventoried all living vegetation material

(green and brown).

Additionally, the procedure and models developed through this

initial effort can be enhanced by conducting field observations at

permanent plots on a regular interval in the years following a fire.

This supports multi-temporal assessments of post-fire vegetation

conditions at intervals defined by land managers and facilitate the

ability to assess and quantify rates of vegetation cover change. To this

end, permanent plots have been established on the Bull and Canyon

Fires (fig. 1), the two fires we sampled in this project, for long-term

monitoring. As we obtain additional samples, we will improve the

modeling to better correspond to observed ground cover.

	 There was one impediment to using pole-mast photography—interference by the overstory tree canopy. We found this technique did

not work well in plots with a living overstory canopy (higher than

30 feet) that had little or no live understory. This situation resulted in

high vegetation index values because the satellite sees the top of the

living canopy but low cover values because the photo was captured

beneath the canopy.Thisis not important when the overstory consists

of burned snags because the understory ground cover is viewable by

both the satellite and photography.

Figure 2—Field sampling included an innovative use of pole-mast photography. We took multiple photos in 

a homogeneous plot; percent ground cover was interpreted using a random dot grid overlayed on photos in 

Esri’s ArcGIS ArcMap.
USDA Forest Service Proceedings RMRS-P-67. 2013 373

Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management Clark and others

Figure 3—Both EVI and NDVI were well correlated with photo-interpreted ground cover although NBR 

performed poorly.
374 USDA Forest Service Proceedings RMRS-P-67. 2013

Clark and others Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management 

Figure 4—The Monument Fire burned near the town of Sierra Vista, 

Arizona, in June 2011. This map shows the predicted ground cover in 

three cover classes: 0-30% (red), 30-60% (yellow), and 60-100% (green).

Application

	 To test our model, we applied it to the Monument and Horseshoe

2 Fires that burned in the Madrean Archipelago of Southern Arizona

during 2011.We compiled maximum NDVI/EVI composite layersto

create a continuous raster layer where every output pixel represents

a predicted ground cover value. Broad classes of vegetation cover

(e.g., 0-30%, 30-60%, 60-100%) were applied to the continuous data

for easier interpretation (fig. 4).An initial validation of the classification for the Monument Fire was encouraging and it appeared to be a

potentially useful layer for predicting ground cover in the semi-arid

Southwest.Acquisition of field observation data for model validation

and further assessment of this methodology is planned for other fires

in the region. Also, class thresholds can be adjusted by managers

based on their resource needs and to identify high risk areas.

Decision Support Tool

	 Results from the described methodology can be integrated into

a larger decision support tool. The decision support process (fig. 5)

is a new tool developed by combining the Fuels Treatment Planning Decision Support Process (2009, Fire Science Digest, JFSP, Issue

7), Forest Service Manual 2520 (2523.1), Calculated Risk: a Tool

for Improving Design Decisions by Larry Schmidt (October, 1998

STREAM NOTES), and Assessing Post-Fire Values-At-Risk with a

New Calculation Tool. This tool is intended to be used by managers

to utilize satellite imagery (vegetation indices), to follow watershed

recovery and evaluate potential impacts to values at risk identified

during the BAER assessment.

	 The process described above utilizes composite satellite imagery

to generate percent cover from NDVI values. By factoring in percent

soil cover, soil depth, and type of vegetation cover, land managers

Figure 5—Flow chart for the decision 

support tool.
USDA Forest Service Proceedings RMRS-P-67. 2013 375

Using Remote Sensing to Monitor Post-fire Watershed Recovery as a Tool for Management Clark and others

may identify potential risks at the watershed scale using departures

from pre-fire conditions. Managers know that ≥60% cover typically reduces the potential for rill development and hillslope erosion

(Noble 1965; Orr 1970; Robichaud and others 2010). Therefore,

if the watershed has recovered to ≥60% cover, the associated risk

of erosion in those areas drops to low or moderate, determinations

which could trigger the removal of temporary protective treatments.

If, however, the analysis shows < 60% cover then removal of treatments may increase the risk to high or very high. Determinations of

high or very high might prompt management agencies to re-initiate

a collaborative process with stake holders identified in the BAER

process which might include the National Weather Service (NWS),

government-based Offices of Emergency Services (OES), Natural

Resources Conservation Service (NRCS), U.S. Geological Survey

(USGS), local flood control districts, and private landowners.

Conclusions and Recommendatons

	 Ourmodel appearsto predictthe post-fire recovery of ground cover

well. We believe the cover class maps can be used in concert with

our developed decision support tool, on-the-ground observations,

and good communication between cooperating agencies to help land

managersmakebetterinformeddecisionsregardingexistingprotective

treatments and burned watersheds upslope. The cover class maps and

decision support tool proposed in this project represent a step toward

a more efficient monitoring approach as well as provide a standard

and repeatable protocol for managers throughout the nation. Finally,

we plan to strengthen the model by applying it to other fires to test

its robustness.

References

Chen, X., Vogelmann, J., Rollins, M., [and others]. 2011. Detecting post-fire

burn severity and vegetation recovery using multitemporal remote sensing

spectralindices andfield-collectedcompositeburnindexdata inaponderosa

pine forest. International Journal of Remote Sensing 32(23): 7905-7927.

Clark,J., Bobbe, T. 2006. Using remote sensing to map and monitor fire damage in forest ecosystems. In: M. Wulder, S. Franklin (Eds.). Understanding

forest disturbance and spatial pattern:Remote sensing and GIS approaches.

(pp. 113-131). Boca Raton, FL: Taylor and Francis.

Clark,J., Kuyumjian, G. 2006. Landscape-scale postfire vegetative condition

monitoring using multi-temporal Landsat imagery on the Cerro Grande

Fire. In: Proceedings of the eleventh biennial forest service remote sensing applications conference. (CD-ROM) USDA Forest Service RSAC:

Salt Lake City, UT.

DeBano, L., Neary, D., Ffolliott, P. 1998. Fire’s effects on ecosystems. New

York, NY: John Wiley and Sons Inc. 333pp.

Diaz-Delgado, R., Salvador, R., Pons, X. 1998. Monitoring of plant community regeneration after fire by remote sensing. In: L. Traboud (Ed.),

Fire management and landscape ecology (pp. 315 - 324). Fairfield, WA:

International Association of Wildland Fire.

Gilbert, J., Butt, K. 2009. Evaluation of digital photography as a tool for

field monitoring in potentially inhospitable environments. Mires and

Peat. 5(5): 1-6.

Huete, A., Didan, K., Miura, T., Rodriguez, E., Gao, X., Ferreira, L., 2002.

Overview of the radiometric and biophysical performance of the MODIS

vegetation indices. Remote Sensing of Environment. 83: 195-213.

Jung, H., Hogue, T., Rademacher, L., Meixner, T. 2009. Impact of wildfire on

sourcewater contributionsinDevilCreek,CA: evidence fromend-member

mixing analysis. Hydrologic Processes. 23(2): 183-200.

Key, C., Benson, N. 2006. Landscape assessment (LA)sampling and analysis

methods. In FIREMON: Fire Effects Monitoring and Inventory System,

D.C. Lutes, R.E. Keane, J.F. Caratti, [and others], (Eds.) (pp. 1–55). Gen.

Tech.Rep.RMRS-GTR-164-CD.Ogden,UT:USDAForestService,Rocky

Mountain Research Station.

Kinoshita, A., Hogue, T. 2011: Spatial and temporal controls on post-fire hydrologic recovery in Southern California watersheds. Catena. 87: 240-252.

Lòpez Garcia, M., Caselles, V. 1991. Mapping burns and natural reforestation

using Thematic Mapper data. Geocarta International. 1: 31-37.

Miller, J., Thode, A. 2007. Quantifying burn severity in a heterogeneous

landscape with a relative version of the delta Normalized Burn Ratio

(dNBR). Remote Sensing of Environment. 109: 66–80.

Noble,E. 1965. Sedimentreduction throughwatershed rehabilitation.In: Proceedings of the federal inter-agency sedimentation conference 1093. Misc.

Publ. 970.(pp. 114-123).Washington,DC:U.S.Department ofAgriculture.

Orr. H. 1970. Runoff and erosion control by seeded and native vegetation on

a forest burn: Black Hills, South Dakota. Res. Pap. RM-60. Fort Collins,

CO:U.S.DepartmentofAgricultureForestService,RockyMountainForest

and Range Experiment Station.

Robichaud, P., Ashmun, L., Sims, B. 2010. Post-fire treatment effectiveness

for hillslope stabilization. Gen. Tech. Rep. RMRS-GTR-240. Fort Collins,

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station. 62p.

Smith, S., Garrett, P., Leeds, J., McCormick, P. 2000. Evaluation of digital

photography for estimating live and dead aboveground biomass in monospecific macrophyte stands. Aquatic Botany. 67: 69-77.

Snider, G., Daugherty, P.,Wood, D.The irrationality of continued fire suppression: An avoided cost analysis of fire hazard reduction treatments versus

no treatment. Journal of Forestry 2007. 104: 431-437.

Sousa,A., Pereira,J., Silva,J. 2003. Evaluating the performance of multitemporal image compositing algorithmsfor burned area analysis. International

Journal of Remote Sensing. 24(6): 1219-1236.

Tucker, C., 1979. Red and photographic infrared linear combinations for

monitoring vegetation. Remote Sensing of the Environment. 8: 127-150.

U.S.DepartmentofAgricultureForestService.2012.Emergencystabilizationburned area emergency response. Forest Service Manual Interim Directive

2520-2012-1. Washington, DC: U.S. Department of Agriculture, Forest

Service.

U.S.GeneralAccountingOffice.2003.Betterinformation neededoneffectivenessof emergencystabilizationandrehabilitationtreatments.GAO-03-430.

April 2003. 63p.

Vanha-Majamaa, I., Salemaa, M., Tuominen, S., Mikkola, K. 2000. Digitized

photographs in vegetation analysis – a comparison of cover estimates. Applied Vegetation Science. 3: 89-94.

Westerling,A., Hidalgo, H.,Cyan D., Swetnam,T. 2006.Warming and earlier

spring increaseWestern U.S. wildfire activity. Science 2006. 313: 940-943.

Westerling,A., Bryant B. Climate change and wildfire in California. Climate

Change 2008: 87 (Suppl 1):S231-S249. DOI 10.1007/s10584-007-9363-z.

Wittenberg, L., Malkinson, D., Beeri, O., Halutzy,A., Tesler, N. 2007. Spatial

and temporal patterns of vegetation recovery following sequences of forest

fires in a Mediterranean landscape, Mt. Carmel Israel. Catena 71:76-83.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
376 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Gila River Basin Native Fishes Conservation Program

Doug Duncan

U.S. Fish and Wildlife Service, Tucson, Arizona

Robert W. Clarkson

Bureau of Reclamation, Glendale, Arizona

Abstract—The Gila River Basin Native Fishes Conservation Program was established to conserve native fishes 

and manage against nonnative fishes in response to several Endangered Species Act biological opinions between 

the Bureau of Reclamation and the U.S. Fish and Wildlife Service on Central Arizona Project (CAP) water 

transfers to the Gila River basin. Populations of some Gila River native fish species are rare in the wild and 

appear on the verge of extirpation. The CAP Program provides monies to undertake and support conservation 

actions for five priority fishes and other native fishes in the Gila River basin by implementing recovery plans. 

The Program will last 30 years, and provide more than $16M. In addition, the Program provides monies to 

control and eradicate nonnative fishes and other non-indigenous aquatic organisms. Thus the Program is 

directed toward actions against nonnative aquatic biota where it interferes with recovery of native forms. 

Fund transfers from Reclamation to the Service began in 1997; about $6.8M has been allocated so far and 

Reclamation will transfer about $9.2M the next 17 years. One half of the funding is identified for native 

fish recovery actions, and one half for non-native aquatic biota control actions. Multiple conservation and 

recovery projects benefitting native fish in the Gila basin have been funded so far. In addition, barriers and 

management against nonnative species have been funded and completed.

Introduction

	 The Gila River Basin Native Fishes Conservation Program (GRBNFCP)is an outgrowth ofreasonable and prudent alternatives(RPAs)

developed by the U.S. Fish and Wildlife Service (Service) and U.S.

Bureau of Reclamation (Reclamation) to prevent jeopardy to the

continued existence offederally listed fishesresulting from operation

of the Central Arizona Project (CAP). The RPAs were developed as

part of a biological opinion (USFWS 1994) under the Endangered

Species Act (ESA) section 7 consultation process (16 U.S.C. 1531

et seq.). The CAP is a Bureau of Reclamation constructed aqueduct

that delivers water from the Colorado River to the Gila River basin

in central and southern Arizona (fig. 1). Nonnative fishes and other

aquatic organisms found in the Colorado River and elsewhere in

the system can be transported via the CAP and released (actively

or passively) into the Gila basin where they can negatively impact

native fishes. The presence of nonnative fishes is now considered

the primary obstacle to conservation and recovery of natives in the

region through a myriad of effects including predation, competition,

hybridization, and pathogen and disease transmission (Clarkson

and others 2005; Hubbs 1955; Miller 1961; Minckley 1973, 1985;

Minckley and Deacon 1991; Minckley and Marsh 2009; Moyle and

others 1986; Williams and Sada 1985).

	 It was determined infeasible to prevent nonnatives from entering

and escaping the CAP,so the RPAs were crafted to limit and mitigate

their upstream spread into streams occupied by listed fishes. They

calledfor constructionoffish barrierstopreventupstreammovements

of nonnatives, monitoring to detect the presence and distribution of

nonnatives, and funding to implement recovery plans of native fishes

and control nonnatives. An RPA was also developed to educate the

public about the values of native fishes and the problems that nonnatives create for them. The GRBNFCP is funded by Reclamation

and is directed by the Service and Reclamation, in cooperation with

the Arizona (AZGFD) and New Mexico Game and Fish (NMDGF)

departments.

	 Here we discuss the two funding RPAs: recovery of natives (RPA

3) and management against nonnatives (RPA 4). The Program mission is to undertake and support conservation actions (recovery and

protection) for federal and state listed or candidate fish species native

to the Gila River basin by implementing recovery plans for those

fishes. The two native trouts are specifically excluded, as they are

the focus of other recovery programs. A strategic plan (USFWS and

others 2007)identifiesthe long-term vision forthe Program as well as

broad goals and actions that are expected to be accomplished during

the current 5-year period (http://www.usbr.gov/lc/phoenix/biology/

azfish/pdf/5yearstraplan2008final.pdf).

	 There are five priority-listed native threatened or endangered fish

speciesin the GRBNFCP: Gila topminnow Poeciliopsis occidentalis,

spikedace Meda fulgida, loach minnow Tiaroga cobitis, Gila chub

Gila intermedia, and razorback sucker Xyrauchen texanus. Other

listed or unlisted native fish in the Gila River basin are also eligible

for conservation actions. In addition, a one-time transfer of $100,000

from Reclamation to the Service was made to fund conservation

actions for the recovery of the threatened Chiricahua leopard frog

Lithobates chiricahuensis.
USDA Forest Service Proceedings RMRS-P-67. 2013 377

Gila River Basin Native Fishes Conservation Program Duncan and Clarkson

Figure 1—Map of the Gila River basin.

	 The principal goals of the GRBNFCP are to (1) enhance conservation status of federally listed and candidate fish species in the Gila

River basin;(2) alleviate and diminish threatsfrom nonnative aquatic

species to native fishes; and (3) remove nonnative fishes that might

enter the Gila River basin via the Central Arizona Project or associated

pathways(USFWS and others 2007).While the focus ofthis Program

is recovery of federally listed species, it is recognized that long-term

viability of protected species is accomplished only in the context of

conservation of intact native fish assemblages and their associated

environments.

	 Funding to conserve Gila River basin native warm-water fishes is

limited.MoniesfromtheProgramareprioritizedsuchthatmeaningful,

achievable, and lasting conservation activities benefit native fishes

according to recovery plan goals and other guidance documents(e.g.,
378 USDA Forest Service Proceedings RMRS-P-67. 2013

Duncan and Clarkson Gila River Basin Native Fishes Conservation Program

conservation agreements, habitat conservation plans, State Wildlife

ActionPlans,integratedwatershedmanagementplans,forestmanagementplans,BLMhabitatmanagementplans).Highestpriorityprojects

for the Program are those that are necessary to (1) prevent extinction

and stabilize populations in the wild, and (2) replicate rare populations in the wild. Actions needed to prevent extinction and stabilize

populations in the wild include (1) construct fish passage barriers to

protect existing populations, (2) control nonnative aquatic species

above barriers, (3) establish new and maintain existing populations,

and (4) implement other actions to remove immediate threats and

thereby help prevent extinction. Actions needed to replicate rare

populations in the wild include (1) safeguard streams for replication

of rare populations; (2) where necessary, construct fish passage barriers and renovate streams;(3) undertake captive production, including

development of propagation techniques; and (4) implement other

actions to insure that rare populations are replicated and protected.

	 Additional priority is given to projects that (1) benefit the five

priority species identified in the 1994 and 2008 biological opinions;

(2) benefit multiple species, including all native fishes of the Gila

River basin; (3) provide immediate on-the-ground benefit; or (4) address other activities pertaining to research or management that aid

in conserving native fish populations and habitat.

Process

	 Reclamation transferred $500,000 to the Service during the first 9

years of the Program, beginning in 1997. One half of that amount was

for native fish recovery actions (RPA3), and one half for nonnative

aquatic biota control actions(RPA4). The original biological opinion

called for 25 years of transfers (USFWS 1994). Another biological

opinion (USFWS 2008) contained conservation measuresto fund the

Program an additional 5 years, with annual transfers increasing to

$550,000. Thus, a total of $16,050,000 over 30 years will be available under the GRBNFCP for the recovery of native Gila River basin

fishes, and management against nonnative aquatic species.

	 The strategic plan (USFWS and others 2007) lists the process the

GRBNFCP follows. The Policy Committee (Service, Reclamation,

AZGFD, NMDGF) provides guidance to the Technical Committee,

approves or rejectsrecommended projects, and deals with policy and

political issuesthat may arise during implementation of the Program.

Because theGRBNFCPis a federallyfundedprogramthatimplements

a regulatory document, the Service and Reclamation make the final

decisions on implementation if consensus cannot be reached with the

State partners.

	 A Technical Committee comprised of one biologist from each of

the above four agencies oversees project solicitation, evaluation,

and recommendations for implementation. Ex-officio members are

representatives from the U.S. Bureau of Land Management and the

U.S. Forest Service. Ideas for original projects (called tasks) are

based on recovery plans and are generated through discussions with

ad hoc groups of biologists, agency, academic, and non-governmental

organizations, private fish biologists working in the Gila River basin,

and other entities as appropriate.

Once annual projects are selected, Reclamation transfers funds to 

the Service for disbursement to implementing entities. Some funds are

retained by Reclamation for tasks they are better suited to conduct or

contract(e.g.,fish barrier designs), and theService also has completed

a fewtasks.Tasksthat are recoveryorresearch projectsrequirewritten

reports; completed reports can be found on Reclamation’s web page:

http://www.usbr.gov/lc/phoenix/biology/azfish/projlisting.html .

Overview

	 About $6.8M has been allocated through fiscal year 2011; half to

each fund. The Service has disbursed or spent $2.6M for RPA 3 and

$2.2 million for RPA4, and Reclamation has disbursed or spent $1M

for RPA 3 and $1.1M for RPA 4. A total of $9.5M should be made

available over the next 17 years.

	 EarlyintheProgram’simplementation,individualtaskswerefunded

separately, and were often less than $25,000. The state game and fish

departments that implemented most of those tasks had a difficult

time maintaining dedicated staff with such piecemeal budgets, and

the contracting process to execute so many tasks was burdensome.

Over the past 5 years, tasks performed by the two state wildlife agencies are lump-sum funded under multi-year umbrella agreements

that facilitate retention of dedicated and qualified staffs and reduces

contracting overhead costs. This change has also reduced the amount

of unobligated funding and the lag time to disburse funds.

Arizona comprises about 93% of the 212,000 km2 Gila River basin 

drainage area, with the remaining area in New Mexico and a small 

fraction in Sonora, Mexico. However, the upper Gila River in New 

Mexico remains an important, interconnected stronghold for species 

such as loach minnow and spikedace, and approximately 23% of 

funding has been directed to the state of New Mexico (table 1). 

When funding is broken down by subbasin (table 2), the upper Gila 

has received 22% of the funding, as has the Verde River subbasin. The 

Verde has received similar funding due to the considerable conservation efforts at Fossil Creek, which so far have included renovating 

the stream and removing nonnative fishes, and stocking native fishes. 

Almost a third of the nonnative funds have been spent on tasks in the 

Verde subbasin. The San Pedro and middle Gila subbasins have also 

been well funded when compared to other subbasins, due to projects 

at the Muleshoe Cooperative Management Area and Bonita Creek, 

respectively.

Expenditures directed toward the five priority species have been 

generous for all but razorback sucker (table 3). Conservation opportunities for this species in the Gila River basin are limited, as 

the most appropriate physical habitat has intractable nonnative fish 

issues (Hyatt 2004; Marsh and Brooks 1989; Schooley and Marsh 

2007). There have been many tasks that benefited spikedace and loach 

minnow because their conservation status is of the greatest concern 

among the five priority species. These two species were recently 

uplisted from threatened to endangered status (USFWS 2012). Also, 

a key goal of three of our large restoration projects at Fossil Creek, 

Bonita Creek, and the Muleshoe Cooperative Management Area has 

been the reestablishment of those two fish.

Table 3 also shows that unlisted species received copious funding. 

This is largely because of the several projects that also support populations of candidate-for-listing roundtail chub Gila robusta (USFWS 

2009) and headwater chub Gila nigra (USFWS 2006). Both species

are second priority species under the GRBNFCP strategic plan. The

Fossil Creek project alone involves the two chubs, and four other

unlisted species. In assigning fundsto each speciesin a multi-species

project, we estimated how much each species benefitted from the

project. Because the two chubs were already present in Fossil Creek,

and have done well post-renovation, much of the project costs were

assigned to them.

Table 4 breaks down expenditures by project categories and shows 

that recovery funds (RPA 3) were spent on basic recovery actions: 

repatriation or reestablishment (36%), creation and management of 

refuges (13%), and surveys (11%). We also funded research (23%) 

on the genetics of the roundtail chub complex (Gila spp.) and desert 

pupfish Cyprinodon macularius, and on propagation of chubs, loach 
USDA Forest Service Proceedings RMRS-P-67. 2013 379

Gila River Basin Native Fishes Conservation Program Duncan and Clarkson

Table 1—GRBNFCP expenditures (000s) through 2011, by fund, 

by state.

Fund Arizona New Mexico Unkown/NA Total

Recovery $2,668 $916 $28 $3,613

Nonnative $2,482 $626 $115 $3,223

Total $5,150 $1,543 $143 $6,837

Table 2—GRBNFCP expenditures (000s) through 2011, by fund, by sub-basin.

Fund Upper Santa San Middle Lower Unknown/

Gila Cruz Verde Salt Pedro Gila Gila Unclassified Total

Recovery $912 $200 $487 $639 $481 $330 $25 $591 $3,665

Nonnative $623 $195 $1,026 $229 $448 $467 $28 $203 $3,219

Total $1,535 $395 $1,513 $868 $929 $797 $53 $794 $6,884

Table 3—GRBNFCP expenditures (000s) through 2011, by fund, by species.

Gila Razorback Gila Loach Other

Fund chub sucker topminnow Spikedace minnow listed Unlisted Total

Recovery $439 $70 $448 $910 $614 $241 $992 $3,714

Nonnative $305 $161 $406 $739 $737 $145 $693 $3,186

Total $744 $231 $854 $1,649 $1,351 $386 $1,685 $6,900

Table 4—GRBNFCP expenditures (000s) through 2011, by fund, by project type.

Fund Research Repatriation Refuges Surveys Data Misc Renovation Book Barriers Program Total

Recovery $874 $1,342 $488 $425 $212 $308 $74 - - - $3,723

Nonnative $311 - $55 $276 $20 $106 $1,782 $45 $601 $68 $3,264

Total $1,185 $1,342 $543 $701 $232 $414 $1,856 $45 $601 $68 $6,987

minnow, and spikedace. About 10 percent of the nonnatives fund 

(RPA 4) went to research, largely focused on new and innovative 

control mechanisms of nonnative species. Our investment in research 

hopefully will facilitate species management and implementation of 

recovery actions by this and other programs. Almost three quarters 

of the nonnative management fund went to stream renovations and 

fish barrier projects.

Recovery of Natives (RPA 3)

Tasks under the recovery of natives fund are intended to lead to 

conservation and recovery of Gila River basin native fishes, mainly 

through on-the-ground projects. One of the tasks that underpins the 

entire recovery aspect of the GRBNFCP is development of a hatchery 

to (1) house wild stocks of imperiled populations as insurance against 

extirpation in the wild; and (2) propagate rare populations to assist 

with replicating them into new, protected streams. We developed and 

expanded the existing Bubbling Ponds Native Fish Research Facility in central Arizona to suit the unique needs of the stream-adapted 

native fishes of the Gila basin. We first developed methods to culture 

GRBNFCP-emphasis species that had not previously been propagated 

in a hatchery situation. Next, we acquired samples from most of the 

most–threatened populations as insurance against loss in the wild, and 

began propagating and replicating them into protected wild streams. 

To date, replications of loach minnow, spikedace, Gila topminnow, 

and desert pupfish have been made to seven streams in the basin, and 

many more are planned.

Some of the major support projects for these recovery actions 

funded under RPA 3 included development of a larval fish key, 

characterization of the genetics of the roundtail chub complex (Gila 

robusta, G. intermedia, and G. nigra) and desert pupfish, determination of propagation techniques for loach minnow, spikedace, and the 

three Gila basin chubs, and development of refuge populations for 

Gila chub, topminnow, and pupfish.

Management Against Nonatives (RPA4)

Tasks under the management against nonnatives fund are intended 

to ameliorate threats nonnative aquatic species are known to have on 

Gila River basin native fishes. Foremost among these was support 

for construction of six fish exclusion barriers to protect existing or 

replicated populations of many Program-priority species, evaluation 

of potential to develop new piscicide formulations, and purchases and 
380 USDA Forest Service Proceedings RMRS-P-67. 2013

Duncan and Clarkson Gila River Basin Native Fishes Conservation Program

applications of existing piscicides and detoxification chemicals for 

restoration projects.

The Future

Because more than $9M will be available for native fish conservation in the Gila River basin in Arizona and New Mexico under the 

GRBNFCP, we expect to continue strong recovery programs and threat 

mitigation for these fishes. Additional species listed under the ESA

couldbe coveredunderthe existingprogram,ormayrequire additional

measures if those species’ conservation and recovery needs are not

met by the existing GRBNFCP. We will continue placing barriers to

protect native fish and their habitats, augmenting and reestablishing

native species, and removing problematic nonnative aquatic species

by the most efficacious means necessary.

	 As a result of ESA section 7 consultation on AZGFD’s federally

funded fish stocking program, a significant conservation program is

being developed (USFWS 2011). The so-called CAMP (Conservation and Mitigation Program) hassimilar goals asthe GRBNFCP, but

covers the entire State ofArizona. Over a 10-year period CAMP is to

conserve and recover its priority species and mitigate threatsto those

species. It is expected CAMP will be funded $500,000 annually for

10 years from federal Sport Fish Restoration Funds. Because only

loach minnow is a priority species in both programs, CAMP actions

will cover a broader range of native aquatic species and a larger area.

These complementary programs should prove greatly beneficial for

native aquatic species in Arizona and parts of New Mexico.

	 As demonstrated above, multiple conservation and recovery projects benefitting native fish in the Gila basin have been completed.

The GRBNFCP has been the most significant funding source for

warm-water native fish recovery actions in the basin for more than a

decade. The Program will continue to be a significant contributor to

this important endeavor.

Acknowledgments

	 We want to extend special thanks to our peer reviewers Sally

Stefferud and Paul Marsh. We also wish to acknowledge current and

former GRBNFCP personnel and Policy and Technical Committee

members; Reclamation: Bruce Ellis, Henry Messing; Service: Paul

Barrett, Tom Gatz, Steve Spangle, Sally Stefferud; NMDGF: Eliza

Gilbert, Chuck Hayes, Andrew Monie, David Propst; AZGFD: Rob

Bettaso,BobBroscheid,MikeChilds,Terry Johnson,TonyRobinson,

Mike Senn, Jeff Sorenson, Bruce Taubert, David Ward, Kirk Young;

BLM: Tim Frey, Tim Hughes; Forest Service: Amy Unthank.

References

Clarkson,RobertW.; Marsh, PaulC.; Stefferud, Sally E.; Stefferud,JeromeA.

2005. Conflicts between native fish and nonnative sport fish management

in the southwestern United States. Fisheries. 30(9): 20-27.

Hubbs,Carl L. 1955. Hybridization between fish speciesin nature. Systematic

Zoology. 4:1-20.

Hyatt, MatthewW. 2004.Assessment ofColorado pikeminnow and razorback

sucker reintroduction programs in the Gila River basin. Phoenix, AZ: Arizona Game and Fish Department. Final Report to U.S. Fish and Wildlife

Service, Cooperative Agreement No. 1448-20181-02-J849.

Marsh, Paul C.; Brooks,J. E. 1989. Predation by ictalurid catfishes as a deterrenttore-establishmentofhatchery-rearedrazorbacksuckers.Southwestern

Naturalist. 34(2): 188-195.

Miller, Robert Rush. 1961. Man and the changing fish fauna of the American

Southwest. Papers MichiganAcademy Science,Arts, Letters. 46:365-404.

Minckley, W. L. 1973. Fishes of Arizona. Phoenix, AZ: Arizona Game and

Fish Department. 293 p.

Minckley, W. L. 1985. Native fishes and natural aquatic habitats in U.S. Fish

andWildlife Region II west of the Continental Divide. Tempe,AZ: Department of Zoology, Arizona State University; Albuquerque, New Mexico:

Report to U.S. Fish and Wildlife Service. 158 p.

Minckley, W. L.; Deacon, James. E., eds. 1991. Battle against extinction:

Native fish management in the American west. Tucson, AZ: University of

Arizona Press. 517 p.

Minckley, W. L.; Marsh, Paul. C. 2009. Inland fishes of the greater southwest: Chronicle of a vanishing biota. Tucson, AZ: University of Arizona

Press. 426 p.

Moyle, P. B.; Li, H. W.; and Barton; B. A. 1986. The Frankenstein effect:

impact of introduced fishes on native fishes in North America. In: Stroud,

R. H., ed. Fish culture in fisheries management. Bethesda, MD: American

Fisheries Society: 415-426.

Schooley, Jason. D.; Marsh, Paul C. 2007. Stocking of endangered razorback

suckers in the lower Colorado River basin over three decades: 1974–2004.

North American Journal of Fisheries Management. 27: 43-51.

Title 16 U.S.C. Conservation: Chapter 35—Endangered Species.

U.S. Fish and Wildlife Service. 1994. Final biological opinion on the transportation and delivery of Central Arizona Project water to the Gila River

Basin (Hassayampa, Agua Fria, Salt, Verde, San Pedro, middle and upper

Gila Rivers, and associated tributaries) in Arizona and New Mexico. Albuquerque, NM: 2-21-90-F-119. 41 p.

U.S. Fish and Wildlife Service. 2006. Endangered and threatened wildlife and 

plants; 12-month finding on a petition to list a distinct population segment

of the roundtail chub in the lower Colorado River basin; and to list the

headwater chub as endangered or threatened with critical habitat. Federal 

Register. 71(85): 26007-26017.

U.S.FishandWildlifeService.2008.Biologicalopinion:reinitiatedbiological

opinion on transportation and delivery of Central Arizona Project water

to the Gila River basin in Arizona and New Mexico and its potential to

introduce and spread nonindigenous aquatic species. Phoenix,AZ:Arizona

EcologicalServicesOffice,02-21-90-F-119,02-21-91-F-406,22410-2007-

F-0081. 162 p.

U.S. Fish and Wildlife Service. 2009. Endangered and threatened wildlife and 

plants; 12-month finding on a petition to list a distinct population segment

of the roundtail chub (Gila robusta) in the lower Colorado River basin;

proposed rule. Federal Register. 74(128): 32352- 32387.

U.S. Fish and Wildlife Service. 2011. Biological and conference opinion

for Wildlife and Sport Fish Restoration Funding of Arizona Game and

Fish Department’s Statewide and urban fisheries stocking program for

2011-2021. Phoenix, AZ: Arizona Ecological Services Office, 22410-

2008-F-0486. 781 p.

U.S. Fish andWildlife Service. 2012. Endangered and threatened wildlife and

plants; endangered status and designations of critical habitat for spikedace

and loach minnow. Federal Register. 77(36): 10810-10932.

U.S. Fish and Wildlife Service, U.S. Bureau of Reclamation, New Mexico

Game andFishDepartment, andArizonaGame andFishDepartment. 2007.

Gila River Basin Native Fishes Conservation Program: Revised Strategic

Plan 2008-2012. Phoenix, AZ. 16 p.

Williams, Jack E.; Sada, Donald W. 1985. Status of two endangered fishes,

Cyprinodon nevadensis mionectes and Rhinichthys osculus nevadensis,

from two springs in Ash Meadows, Nevada (U.S.A.). Southwestern Naturalist. 30: 475–484.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 A-1

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Mapping Ecological Systems in Southeastern Arizona

Jim Malusa and Donald Falk

School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona

Larry Laing

Environmental Consultant, Gold Canyon, Arizona

Brooke Gebow

The Nature Conservancy, Southeastern Arizona Preserves, Hereford, Arizona

Abstract—Beginning in 2007 in and around the Huachuca Mountains, the Coronado National Forest 

and other partners have been mapping ecosystems at multiple scales. The approach has focused on 

identifying land type associations (LTA), which represent the sum of bedrock and superficial geology, 

topography, elevation, potential and existing vegetation, soil properties, and local climatic variables. 

This mapping effort has been extended into the FireScape program, in which multiple partners utilize 

ecological land type mapping as a framework for fire planning across the Sky Island bioregion. Land 

type association maps for the Catalina-Rincon mountains (available at www.azfirescape.org) are 

used for managing ecological units (e.g., mixed conifer on granitic soils) typically no smaller than a 

thousand acres, and often much larger. Land type associations compliment raster-based sources of 

information such as LANDFIRE. Not surprisingly, the success of the project depends on an accurate 

depiction of vegetative and physical setting reality, not just interpretations of remote imagery. LTAs 

provide an intuitive and informative method of characterizing complex landscapes for planning and 

ecosystem management.

Introduction

The notion of a landscape as a piece of land with a certain character has persisted for well over a thousand years (Calder 1981). That 

character might be of a lush river valley, a stony ridge, or a desert 

plain. These are natural units, with characteristic landforms supporting 

equally characteristic biota. For instance, in southeastern Arizona you 

might find creosotebush, little-leaf sumac, and viscid acacia dominating the alluvial fans at the toe of a limestone mountain. Landform, 

soils, geology, climate, hydrologic regime, and biota make what is 

recognized as an ecological system (Tansley 1935).

Mapping of ecological systems, or ecosystems, began with Humbolt 

and Bonbland’s 1807 “Essay on the Geography of Plants,” and has 

proceeded in earnest ever since. There remains much to do, in part 

because ecosystems exist at myriad scales, from course systems covering millions of square miles (e.g., humid temperate) to fine (e.g., 

mixed conifer on Mount Lemmon metasediments). To advance a single 

standard for ecosystem mapping at multiple scales, the United States 

Forest Service has adopted the National Hierarchical Framework of 

Ecological Units (table 1; Cleland and others 1997). At course scales, 

the hierachy adopts Bailey’s 1995 classification. In this paper we focus 

on the finer scales, particularly the landscape scale and its associated 

ecological unit, the land type association.

Land type associations are typically no smaller than 1000 acres (404 

ha), and are often much larger. They integrate landform and ecology 

into cohesive units that are intuitively apparent. For example, the 

typical structure of a rolling oak woodland on an alluvial fan—grass 

on the south aspects and trees on the north—is captured as a repeating 

pattern within a single land type association. However, when viewed 

with remotely sensed vegetation pixels, the pattern is fragmented by 

the assignment of at least eight different vegetation types (fig. 1). 

Mapping land type associations is standard practice in much of 

the United States. In the northern Great Lake states they are used for 

landscape–scale assessments and planning, targeting opportunities for 

critical wildlife habitat restoration, and for on-the-ground silvicultural management (Almendinger and others 2000). The entire State 

of Missouri has been mapped with land type associations (Nigh and 

Schroeder 2002). In Arizona, however, landscape mapping is a recent 

development. Spurred by large wildfires in 2002-2003 that burned 

across jurisdictional bounds, Arizona land managers recognized the 

need for multi-agency landscape-level treatments to reduce fuel loads 

and reintroduce fire as a natural process in those ecosystems where it 

played a historic role. These efforts included landscape level mapping 

that began with the Huachuca Mountains and environs (Laing and 

others 2005), and continues today with the multi-agency FireScape 

project encompassing millions of acres of southeastern Arizona (www.

azfirescape.org).
A-2 USDA Forest Service Proceedings RMRS-P-67. 2013

Malusa, Falk, Laing, and Gebow Mapping Ecological Systems in Southeastern Arizona

Table 1—National hierarchy of ecological units (Cleland and others 1997).

Ecological unit Planning and 

analysis scale Principal map unit design criteria Purpose, objectives, and general use

Domain

Global

Continental

Regional

Broad climatic zones or groups (e.g., dry, 

humid, tropical)

Broad applicability for modeling and 

sampling.

Strategic planning and assessment. 

International planning.

Division

Regional climatic types (Koppen 1931, 

Trewatha 1968) Vegetational affinities (e.g., 

prairie or forest) 

Province Patterns in dominant potential natural 

vegetation (PNV), macroclimate, geology

Section

Subregion

Geomorphic province, geologic age, 

stratigraphy, lithology, regional climatic 

data, phases of soil orders, suborders, or 

great groups, PNV Strategic, multiforest, statewide, and 

multiagency analysis and assessment.

Subsection

Geomorphic process, surficial geology, 

lithology, phases of soil orders, suborders, 

or great groups, subregional climatic data, 

PNV—formation or series

Land type association Landscape

Geomorphic process, geologic formation, 

surficial geology, elevation, phases of soil 

subgroups, families, or series, local climate, 

PNV

Forest or area wide planning, and 

watershed analysis.

Land type

Land Unit

Landform and topography (elevation, aspect, 

slope gradient, and position), phases of soil 

subgroups, families, or series, rock type, 

geomorphic process, PNV Project and management area 

planning and analysis.

Land type phase Phases of soil subfamilies or series, landform 

and slope position, PNV

Figure 1—Oak woodland near Sonoita, Arizona, at about 5,000 feet elevation. This landscape is mapped as a single land type association, the Oak Tree Canyon dissected fans. In contrast, the same area is 8 different vegetation types represented as 30 meter square 

pixels in the lower image (raster data from LANDFIRE).
USDA Forest Service Proceedings RMRS-P-67. 2013 A-3

Mapping Ecological Systems in Southeastern Arizona Malusa, Falk, Laing, and Gebow

Methods

The National Hierarchical Framework of Ecological Units (Cleland 

and others 1997) assigns the relative importance of mapping criteria 

at different spatial scales (table 1). Note that the mapping criteria for 

the smallest units have little in common with the largest unit. For 

example, although local landform is important in recognizing land 

type associations, it plays little or no role in distinguishing larger 

ecosystems, where regional climate is paramount (fig. 2). In general, 

climate, as modified by topography, is the dominant criterion at upper levels, while geomorphic process, soils, and potential natural 

communities are more important at lower levels. The comprehensive 

“Terrestrial Ecological Unit Inventory Technical Guide” is an in-depth 

compendium of mapping techniques (Winthers and others 2005). 

We mapped at three levels in southeastern Arizona: land type associations, land types, and ecological units. Land type associations 

are based primarily on bedrock and superficial geology, topography, 

elevation, potential and existing vegetation, soil properties, and local 

climatic variables (fig. 3). Land types are subdivisions of land type 

associations. Land types are based on more localized or finer scale 

topographic, geologic, soil, and plant species association properties 

than in land type associations (fig. 4). These units have land surface 

variations and other physical and biological properties that influence 

hydrologic function, moisture distribution and retention, which affect finer scale plant community composition and distribution. Land 

types range from having no or very few weakly developed drainage 

features, to being densely dissected by deeply incised washes and 

streams. Drainage characteristics (rocky, silty or sandy; confined or 

not; etc.) are strongly dictated by the land types within which they 

originate or through which they flow. 

Finally, for management and analysis purposes, we aggregated 

similar land type associations into ecological landscape units, or 

simply ecological units. As used in this project, ecological units are 

groupings of land type associations with broadly defined similarities 

in vegetation, topography, geologic and soil features. They provide a 

“big picture” view of the ecosystems. Ecological units are intended 

to provide a framework for the assessment of existing conditions and 

departures, desired conditions, and environmental implications.

Figure 2—An example of the relative importance of mapping unit criteria 

at different scales. For instance, at the level of Section, which typically 

encompasses thousands of square miles, regional climate is important, 

but landform is not. The importance of criteria are reversed at the level 

of land type association.

Figure 3—Three land type associations along Ash Creek, near the Rincon 

Mountains. The foreground is a broad floodplain with deep soils supporting 

mesquite, velvet ash, sycamore, and emory oak. The ridge on the right is 

largely composed of schist, and holds primarily ocotillo, mesquite, and 

beargrass. The ridge on the left is composed of granite and holds blue 

oak, hopbush, and manzanita.

Figure 4—Two land types from the same land type association: (a) the 

dominant vegetation is acacia and prickly pear; (b) palo verde and cholla.

a

b
A-4 USDA Forest Service Proceedings RMRS-P-67. 2013

Malusa, Falk, Laing, and Gebow Mapping Ecological Systems in Southeastern Arizona

Case Studies

Greater Huachuca Area

In southeastern Arizona, landscape scale mapping began with the 

Greater Huachuca Fire Management Group (the Coronado National 

Forest, the National Park Service, and Fort Huachuca (U.S. Army)) 

(Gebow and Lambert 2005). The group aimed to develop a management plan that would allow cross-juristicional treatments to return the 

region to something closer to its historical fire regime. The resulting 

map covered a half-million acres (Laing and others 2005).

The first step was to collect the relevant layers of data: soils, geology, topography, elevation, climate, existing vegetation, and remote 

sensing imagery. The analysis of the data began with determining 

which landforms are distinctive at the landscape level. For instance, 

assorted mountain landforms with bedrock at or near the surface are 

distinguished from the deep valley fill deposits. Alluvial fans and terraces adjacent to the mountains were separated from the valley floor. 

Initial mapping of landforms was aided by imagery from Landsat 

Thematic Mapper (TM) and USDA National Agriculture Imagery 

Program (NAIP) (imagery available at http://aria.arizona.edu/). We 

also used a shaded relief map that was digitally manipulated to the 

angle and position of the sun.

The landform patterns detected in the initial step usually coincide 

strongly with geologic materials and soil characteristics provided 

by National Resource Conservation Service (NRCS) soil surveys 

(available at http://soils.usda.gov/). Various biotic communities were 

determined through the use of the imagery, topography, existing 

vegetation maps (e.g., Southwest ReGAP, available at http://earth.gis.

usu.edu/swgap/) and local experts. Associated fire regimes and condition classes were assessed by available literature (e.g., Kaib 1998). 

Precipitation and temperature data were compiled from the USFS 

General Ecosystem Survey. The last step was to validate the units 

on the ground and to make appropriate corrections. The final units, 

developed with feedback from the fire-planning group, represented 

15 land areas that were relatively consistent in terms of geologic 

material, landform, and vegetation patterns. 

Land Type Associations of Southeastern 

Arizona

The Huachuca Area map prompted the National Park Service, the 

U.S. Forest Service and U.S. Geological Survey to conduct additional 

mapping of 109 land type associations over nearly two million acres 

in southeastern Arizona (Cleland and others 2008). The study area, 

which included the Greater Huachua Area, is bounded to the south 

by the international border, to the west by the Santa Cruz River, to 

the east by the San Pedro River, and by Redington Pass to the north, 

between the Catalina and Rincon Mountains (fig. 5). The methodology was similar to that used to map the Greater Huachuca area, using 

GIS layers pertaining to geology, soils, elevation, slope, and existing 

vegetation. However, because of increased variation in local climate 

over the larger study area, we calculated potential evapotranspiration 

(PET) using the Thornthwaite (1948) equation (White and Host 2002). 

Using PRISM data (Daly and Taylor 2000) we also calculated mean 

monthly temperatures for the time period of 1961 through 1990. These 

data were included in the description of each land type association, 

along with physiography, soils, geology, vegetation, and fire history 

over the last 35 years, during which 1,307 fires burned a total of 

243,039 acres. Land types were also deliniated and briefly described 

within the context of their respective land type associations.

Catalina–Rincon FireScape

The Coronado National Forest, University of Arizona, and other 

partners sponsor the ongoing FireScape project (www.azfirescape.

org). FireScape works to advance the science of fire management 

from a landscape and ecological perspective in southeastern Arizona. 

The most comprehensively mapped lands to date are 700,000 acres 

of the Catalina and Rincon Mountains outside Tucson, a study area 

that included the Little Rincons and the east slope down to the San 

Pedro River. Methods were similar to prior phases (above), with two 

exceptions: the addition of draft geology maps provided by the Arizona 

Geological Survey (Reynolds, 2007), and significantly more time in 

the field. We photographed all but one land type association, and took 

notes on the associated vegetation. The data on these 300 photo points 

and 200 species were uploaded onto the website, where they appear 

on dynamic maps of the mapping products (www.azfirescape.org).

We mapped 120 land type associations in the Catalina and Rincon 

Mountains. We also subdivided land type associations into 327 land 

types, as well as aggregating land type associations into ecological 

units for planning and public education (fig. 6). At lower elevations 

(i.e., pediments, fans and valleys), the land types were based primarily 

on soil surveys and drainage features; at higher elevations (usually 

roadless and steep and lacking detailed soil surveys), the land types 

are recognized by surficial and bedrock geology, vegetation, and gross 

physiographic differences. For example, the east face of the Rincons 

between 4000 and 6000 ft (1524 to 1851 m) is home to Madrean oaks 

and pinyon pine, two species of similar size but different response 

to fires (oaks resprout, pines don’t). Hiking the mountain revealed 

a clear distinction in substrate preference between the two species, 

with pinyon dominating habitat with exposed bedrock, and the oaks 

preferring colluvium (landslides). The distinction was readily apparent 

to the eye, but less so from the current generation of remotely sensed 

pixel vegetation maps, such as LANDFIRE and ReGap landcover 

(cited above), which rendered the landscape as 30 meter pixels.

Conclusions 

Have the ecosystems maps been useful? Yes. The map of the 

Greater Huachuca area, for instance, provided the 15 mapping units 

used as “project areas” in a successful three-agency, whole-landscape 

Environmental Assessment (EA) for fire treatments (EA available 

at http://www.azfirescape.org/content/huachuca-firescape-projectdocuments/). The Huachuca units similarly served as the basis for 

a Biological Opinion that addressed effects of fire treatments on 10 

Federally listed species. Likewise, the ecological units map of the 

Catalina and Rincons is being used to guide the identification of 

treatments, and the analysis of the effects of those treatments to meet 

enviromental compliance requirements. For example, the LANDFIRE 

fuels layer can be combined with the ecological units layer to examine 

fire behavior at the landscape scale.

However, comparing LANDFIRE pixels (raster data) and ecological unit polygons (vector data) can be a problem; the mapping units 

often don’t match up. The ecological unit called “desert-oak transition on granitics,” for example, has no straighforward equivilant in 

the LANDFIRE vegetation lexicon. Consequently, as the FireScape 

project has moved on to the Chirichaus, Dragoon, and Dos Cabezas 

mountains in the southeastern Arizona, we have adopted names 

already in use by LANDFIRE (FireScape maps available at http://

www.azfirescape.org/chiricahua-dragoons-dos-cabezas/).

With this adjustment, ecological system maps and remotely sensed 

pixels can not only be used in tandem, but also to check on their 
USDA Forest Service Proceedings RMRS-P-67. 2013 A-5

Mapping Ecological Systems in Southeastern Arizona Malusa, Falk, Laing, and Gebow

Figure 5—Land type associations of southeastern Arizona, with digital elevation model. The international border is the southern boundary (from 

Cleland and others 2008).
A-6 USDA Forest Service Proceedings RMRS-P-67. 2013

Malusa, Falk, Laing, and Gebow Mapping Ecological Systems in Southeastern Arizona

Figure 6—Ecological units of approximately 700,000 acres of the Catalina and Rincon Mountains, Arizona. Polygon boundaries within ecounits 

show the smaller land types and land type associations.
USDA Forest Service Proceedings RMRS-P-67. 2013 A-7

Mapping Ecological Systems in Southeastern Arizona Malusa, Falk, Laing, and Gebow

accuracy. When two maps disagree, it is time to get into the field and 

take a closer look.

Acknowledgments 

Thanks to Dave Cleland for his brains and charm; to Jim Jordan for 

his kindness; to Rick Watson, George Robertson, and Dave Diamond 

for humor and vision in the field; and to reviewers George Ferguson 

and Shirley Papuga for making this better than it was. 

References

Almendinger, J.C.; Hanson, D.S.; Jordan, J.K. 2000. Land type associations 

of the Lake States. St. Paul, MN: State of Minnesota, Department of 

Natural Resources.

Bailey, R.G. 1995. Description of the ecoregions of the United States. 2d ed., 

rev.. Misc. Publ. No. 1391 (rev.). Washington, DC: U.S. Department of 

Agriculture, Forest Service. 108 p.

Brown. D. E., Lowe, C.H., and Pase, C.P. 1979. A digitized classification system 

for the biotic communities of North America, with community (series) and 

association examples for the Southwest. Journal of the Arizona-Nevada 

Academy of Science. 14 (supplement 1): 1-16.

Calder, W. 1981. Beyond the view: Our changing landscapes. Melbourne: 

Inkata Press.

Cleland, D.T.; Avers, P.E.; McNab, W.H.; Jensen, M.E.; Bailey, R.G., King, T.; 

Russell, W.E. 1997. National hierarchical framework of ecological units. 

In: Boyce, M. S.; Haney, A., ed. 1997. Ecosystem management applications for sustainable forest and wildlife resources. New Haven, CT: Yale 

University Press: 181-200.

Cleland, D.T.; L. Laing,; Watson, R.L.; Malusa, J.; Jordan, J.K.; Diamond, 

D.D,; and G. Robertson. 2008. Land type associations of southeast Arizona. 

361 p. Online: http://www.azfirescape.org/home/related_activities/.

Daly, C. and Taylor, G.. 2000. United States average monthly or annual mean 

temperature, 1961-90. Oregon State University, Spatial Climate Analysis 

Service (SCAS/OSU); Corvallis, OR: Climate Source, Inc.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein. NOTE: This paper was 

originally part of this session and belatedly moved to this location, so page numbers are not sequential within these proceedings.

Kaib, J. 1998. Fire history in riparian canyon pine-oak forests and the intervening desert grasslands of the Southwest borderlands: A dendroecological, 

historical, and cultural inquiry. Tucson, AZ: The University of Arizona. 

234 p. Thesis.

LANDFIRE 1.1.0 Existing vegetation type layer. U.S. Department of the 

Interior, Geological Survey. Online: http://landfire.cr.usgs.gov/viewer/ 

[Accessed May 11, 2012].

McNab, W.H.; Cleland, D.T.; Freeouf, J.A.; Keys, J.E., Jr.; Nowacki, G.J.; 

Carpenter, C.A., comps. 2005. Description of ecological subregions: sections of the conterminous United States [CD-ROM]. Washington, DC: U.S. 

Department of Agriculture, Forest Service. 80 p.

Nigh, T.A.; Schroeder, W.A. 2002. Atlas of Missouri ecoregions. Jefferson 

City, MO: Missouri Department of Conservation. 212 p.

Reynolds, S. 2007. Digital geologic map of Arizona. Phoenix, AZ: Arizona Geologic Survey. Online: http://www.azgs.az.gov/. [Accessed May 11, 2012].

Tansley, A. G. 1935. The use and abuse of vegetational concepts and terms. 

Ecology. 16(3): 284-307.

Thornthwaite, C. W. 1948. An approach toward a rational classification of 

climate. Geographic Review. 38:55-94.

USGS National Gap Analysis Program. 2004. Provisional digital land cover 

map for the southwestern United States. Version 1.0. Logan, UT: Utah State 

University, College of Natural Resources, RS/GIS Laboratory. Online: 

http://earth.gis.usu.edu/swgap/. [Accessed May 11, 2012].

White, M. A. and G. E. Host. 2002. An assessment of potential evapotranspiration for the northern Lake States. Online: http://www.nrri.umn.edu/gla/

pet.htm. [Accessed on Sep. 21, 2006].

Winthers, E.; Fallon, D.; Haglund, J.; DeMeo, T.; Nowacki, G.; Tart, D.; 

Ferwerda, M.; Robertson, G.; Gallegos, A.; Rorick, A.1; Cleland, D. T.; 

Robbie, W. 2005. Terrestrial ecological unit inventory technical guide. 

Washington, DC: U.S. Department of Agriculture, Forest Service, Ecosystem 

Management Coordination Staff. 245 p.
USDA Forest Service Proceedings RMRS-P-67. 2013 381

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Potential for Extending Major Land Resource Areas into 

Northern Mexico

Roy S. Mann

USDA-NRCS Retired, Range Management

Philip Heilman and Jeffry Stone

USDA-ARS, Southwest Watershed Research Center

Abstract—There is a significant history of cooperative efforts between Mexico and the United States on 

natural resource management issues. Mexico and the United States have jointly conducted research and 

developed range management technologies. Bringing these technologies together and improving technical 

communications are an ongoing process. This paper discusses a potential tool that can provide a common 

denominator for both countries to more easily frame, describe, and share data relative to rangeland resources. 

The objective is to present possibilities for utilizing current data and provide a vehicle that can facilitate 

technical communications. Existing maps including climate and elevation were used to define probable areas 

of Chihuahua and Sonora that would be similar enough to Major Land Resource Areas (MLRAs) of Arizona 

to consider them an extension of those MLRAs. Reconnaissance surveys were made to compare soils and 

vegetation to those described in Arizona. Comparisons were also made between Ecological Sites (ESDs) used 

in the United States with those developed in Mexico by COTECOCA. A preliminary map was developed that 

represents probable boundaries of MLRA 41 if extended from the U.S. border into the states of Chihuahua 

and Sonora. Some sites were mapped to test application of U.S. ecological site descriptions with on-ground 

conditions in Chihuahua. The potential for extending MLRA boundaries from the United States into Mexico 

are feasible and realistic. This would facilitate the direct use of Ecological Site Descriptions across borders 

and improve exchange of rangeland data between the two countries.

Background and Objectives

	 Our presentation today is not the first discussion of this subject.

There have been two previous accounts of the effort to explore the

possibilities of extendingMajorLandResourceArea (MLRA)boundaries from the southern United States border into Mexico. These are

Rebecca MacEwen and others paper (2005) Defining Boundaries 

Across Borders: A Case Study Extending a Major Land Resource Area 

Into Mexico, and Philip Heilman and others(2000) A Framework for 

Cooperation Across the U.S./Mexico Border.

	 Theprojectdemonstratesthe cooperative natureofseveral agencies

and individuals.All activities were under the leadership of the Southwest Watershed Research Center of the USDA-ARS. The principal

partner in Mexico was the Instituto Nacional de Investigaciones de

Forestales,Agricolas, y Pecuarias(INIFAP).Alicia Melgoza played a

criticalrole in assistingwith scheduling activitiesinMexico, advising

on conditions and characteristics of vegetation, and helpingwith plant

identification. Antonio Chavez, former Director of Campo Experimental La Campana, made the facilities of La Campana available to

serve as a headquarters site for those involved in the project. Rafael

Fierros of Comisión Tecnico Consultiva para la Determinación Regional del losCoeficientes deAgostadero (COTECOCA),Chihuahua

devoted his time and assistance to the effort, regarding COTECOCA

site identification and plant identification.

	 The USDA Natural Resources Conservation Service (NRCS)

provided excellent technical expertise with the participation of Dan

Robinett, Range Conservationist, Tucson, Steve Barker, State Range

ConservationistPhoenix, andDonBreckenfeld,SoilScientist,Tucson.

In addition, the producers of Ejido Nuevo Delicias, Chihuahua,

cooperated by making their lands available for site correlation work.

This paper will present an overview of the activities and findings in

an attempt to identify limits of an MLRA boundary when extended

from the U.S. border in Arizona to the states of Chihuahua and Sonora. Emphasis will be on an appreciation and understanding of the

materials and data bases available that indicate the compatibility of

the concept with other data bases and agencies rather than just the

effort made to test extending MLRAs into Mexico.

Classification System

	 MLRAs are part of a hierarchical classification system that is

important in classifying soils and ecological sites. The system was

developedbytheSoilConservationService,nowNRCS,andpublished

as USDAHandbook 296 in 1965.The handbook wasrevised in 1978,

and published in 1981. This version was again revised and updated

and published in 2006. The classification system has important and
382 USDA Forest Service Proceedings RMRS-P-67. 2013

Mann and others Potential for Extending Major Land Resource Areas into Northern Mexico

valuable application, particularly related to rangelands in terms of

ecologicalsitework, andsoil/site correlation.The systemcomponents

are shown below:

NRCS Hierarchical Classification System:

	 LAND RESOURCE REGIONS

	 	 MAJOR LAND RESOURCE AREAS

	 	 	 LAND RESOURCE UNITS (or Common ResourceAreas)

	 	 	 	 ECOLOGICAL SITES

	 	 	 	 	 SOIL SERIES

Since the 1960’s, NRCS has been using the soil and ecological

site databases as the basic units of a hierarchical natural resource

classification system called Major Land Resource Areas (MLRA’s). 

This system provides a basis for making decisions about national and

regional agricultural concerns, helps identify needs for research and

resource inventories, provides a broad base for extrapolating the results

of research within national boundaries, and serves as a framework for

organizing and operating resource conservation programs”(Fox and

others 1999).

Land Resource Regions

	 The broadest category in the hierarchy isLand Resource Regions.

The region of most interest for this paper is Region D,Western Range

and Irrigated Region shown in figure 1. It makes up 549,725 square

miles(1,424,480 square kilometers). Thisisthe largest of all the land

resource regions in land area. It is a semi-desert or desert region of

plateaus, plains, basins, and many isolated mountain ranges.

Major Land Resource Areas (MLRAs)

	 Regions are further subdivided into Major Land Resource Areas

(fig. 2). This level of classification is the subject of this document.

	 MLRA lines end at the international border. The characteristics

that the lines represent do not end at the border, but extend to some

unknown extent into Mexico. This recognition was the fundamental

rationale for exploringwhere the boundariesmightliewithinMexico.

Some important aspects of the MLRA system are:

•	 It is nationwide in scope.

•	 It is updated and refined periodically. Some states have refinements that reflect county level determinations.

•	 It is not an isolated concept useful only to NRCS. MLRAs have

been correlated with other agency classification systems.

	 In addition to use by the Bureau of Land Management in its use

of ecological sites, it has also been correlated with other agency

classification systems. The information shown below illustrates the

correlation that has been done to date. USDA Handbook 296 (2006)

cross-references MLRAs with Environmental Protection Agency’s

(EPA) Level III Ecoregions, and with United States Forest Service

(USFS) ecological units for the conterminous United States. A few

crossreferences are shown in table 1.The fact that MLRAs have been

Figure 1—Region D: Western Range and 

Irrigated Region (source: USDA NRCS 

Agriculture Handbook 296).
USDA Forest Service Proceedings RMRS-P-67. 2013 383

Potential for Extending Major Land Resource Areas into Northern Mexico Mann and others

correlated to other agency classification systems broadens their utility and expands their range of application. Similar applications and

interest are shown in The Nature Conservancy’s work in the Apache

Highlands Ecoregion, Gori and Enquist (January 2003).

	 In our primary area of interest, the following figure illustrates the

MLRAs asthey occurinArizona. One can see that in the Southeastern

corner of Arizona lays MLRA-41, Southeastern Arizona Basin and

Range, which is the primary focus of this paper. MLRA 41 is further

subdivided into three Common Resource Areas (Climatic Zones), or

Land Resource Units (fig. 3):	

•	 Common Resource Area 41-1 Mexican Oak-Pine Woodland

and Oak Savannah

•	 Common Resource Area 41-2 Chihuahuan-Sonoran Desert

Shrub Mix

•	 Common Resource Area 41-3 Southern Arizona Semidesert

Grassland

The extension of MLRAs into Mexico would allow the direct use of

Ecological Site Descriptions (ESDs) for “on-the-ground” ecological

interpretations. In addition to being a valuable grazing management

tool, ecological site descriptions provide a documented biodiversity

benchmark in the list of plant species that are known to occupy a

particular ecologicalsite.In additionto providinga list of documented

plants, the ESD also indicatesthe potential primary production range

of each species, or groups of species. This provides a new level of

information subject to interpretation for biodiversity studies.

Acitivies and Methods

	 Initial work consisted of reconnaissance surveys, focusing on observations of vegetation and soil characteristics as

compared with that of ecological sites described in Arizona.

A primary tool used in the evaluation of MLRA boundaries was the

application of Geographical Information System (GIS) technology.

Virtually all the activities of the project were correlated with and

incorporated into a GIS database that proved invaluable in evaluating

and displaying findings.

Table 1-Cross reference of MLRAs, USFS, and EPA.

MLRA USFS EPA

39 Arizona and New Mexico Mountains M313A White Mountains- 23 Arizona/New Mexico Peaks- 

 San Francisco Mogollon Rim

40 Sonoran Basin and Range 322B Sonoran Desert 81 Sonoran Basin and Range

41 Southeastern Arizona Basin and Range 321A Basin and Range 79 Madrean Archipelago

41 Southeastern Arizona Basin and Range 321A Basin and Range 24 Chihuahuan Deserts

42 Southern Desertic Basins, Plains, 

 and Mountains 321A Basin and Range 24 Chihuahuan Deserts

Figure 2—Major Land Resource Areas of the United States (source: U.S. Department of Agriculture Handbook 296), .
384 USDA Forest Service Proceedings RMRS-P-67. 2013

Mann and others Potential for Extending Major Land Resource Areas into Northern Mexico

Figure 3—MLRAs and CRAs of Arizona.
USDA Forest Service Proceedings RMRS-P-67. 2013 385

Potential for Extending Major Land Resource Areas into Northern Mexico Mann and others

	 Based on information developed from the initial reconnaissance

trips and othersources, a map was developed by Heilman (2000) and

MacEwen(2005)showinga suggestedextensionofMLRAboundaries

(fig. 4). This preliminary map was largely developed by digitizing

vegetation maps of Sonora and Chihuahua from Brown and Lowe’s

(1994) Biotic Communities of the Southwest. Ultimately, the Brown

and Lowe based map turned out to be an excellent first approximation

that provided valuable guidance for field determinations.

	 Further field work included investigations by Don Breckenfeld,

NRCS Soil Scientist. Soil profiles were described and compared to

known soil series in Arizona. All of the soil profiles described were

determined to be very similarto soilseries associated with MLRA-41

(fig. 5).

	 Existing mapsrelative to climate and elevation were used to define

probable areas ofChihuahua andSonora thatwould be similar enough

to Major Land Resource Areas (MLRAs) of Arizona to consider

them an extension of those MLRAs. Comparisons were also made

between Ecological Site Descriptions used in the United States with

site descriptions developed in Mexico by COTECOCA.

	 Hydrologic studies were also included in the field activities. A

rainfall simulator was used to evaluate hydrologic conditions as

related to ecological conditions and soils on several sites within the

projected MLRA area. The technology and procedures are described

by Stone and Paige (2003) (fig. 6).

	 As another evaluation of site similarities, a small ranch just south

of La Campana was mapped using NRCS ESDs. Most of the sites

were very similar to ecological sites in Arizona, and should be considered to be adequate for management interpretations (fig. 7). The

Arizona site descriptions used in developing the above map, and a

corresponding translation name used are as indicated in the table 2:

Comments Regarding Range Sites, Forage 

Production Sites, and Ecological Sites

	 Both the United States and Mexico have invested major resources

in theirrespective effortsto define and classify rangeland unitsforthe

purposes of conducting inventories, analysis of rangeland resources,

and asrange management tools.The mostsignificant units developed

Figure 4—Possible extent of MLRA 41 boundaries (from Heilman and 

others 2000).

Figure 5—Describing soil, Chihuahua.

Figure 6—Rainfall simulator studies, Chihuahua.

forthese purposes have been theEcological Site concept in theUnited

States, and the Sitios de Productividad Forrajera (Forage Production

Sites) in Mexico. The Range Site (now Ecological Site) concept

was adopted by the USDA Soil Conservation Service (now Natural

ResourcesConservation Service [NRCS])in 1949, based on the work

of E.J. Dyksterhuis (1949).

	 The principal application of Range Sites was in conducting inventories andrangelandanalysesduringtheprocessofrangemanagement

planning with ranchers on privately owned lands. The purpose was

to help individual ranchers manage rangeland resources to improve

rangeland conditions and increase economic returns for the rancher.

	 The primary purpose of the COTECOCA sites (fig. 8), at the time

they were developed and mapped, wasto establish grazing capacities

for the different major plant community types for the entire nation.

Heilman and others (2000) made several important observations

regarding the resolution differences between ecological sites and

forage production sites:
386 USDA Forest Service Proceedings RMRS-P-67. 2013

Mann and others Potential for Extending Major Land Resource Areas into Northern Mexico

Figure 7—Ecological sites mapped in Chihuahua.

Table 2-Site names and map symbols

Arizona Site Name Translation Map Symbol

Shallow Hills 41-1 Lomas – Suelos Poco Profundo LPP

Loamy Hills 41-1 Lomas Francosa LF

Loamy Upland 41-3 Llano Francoso LlF

Granitic Hills 41-3 Lomas Graníticas LG

Sandy Loam Upland 41-3 Franco Arenosa FA

Shallow Upland 41-3 Llano – Suelos Delgados LlD
USDA Forest Service Proceedings RMRS-P-67. 2013 387

Potential for Extending Major Land Resource Areas into Northern Mexico Mann and others

Defining sites on an ecological, rather than forage, basis leads to 

a finer resolution of sites in the U.S. In Arizona, 503ecological sites 

are defined over a total area of 29m hectares. In Chihuahua, 64 forage 

production sites are defined for 24m hectares. Per unit area then, there 

are almost 7 times as many ecological sites defined in Arizona as there 

are forage production sites defined in Chihuahua. Both approaches 

distinguish areas with the potential for homogeneous stands, such as 

sacaton bottoms, as separate sites. However, the Natural Resources 

Conservation Service defines more sites in areas with heterogeneous 

plant communities, even though those sites could all have the same 

forage production capability.

Results

	 A refined map of probable MLRA boundaries was developed that

represents estimated boundaries ofMLRA—41ifextendedsouthfrom

the U.S. borderinto the states ofChihuahua and Sonora. Figure 9 shows

the adjustments made to the original map that reflect, hopefully, a

more accurate definition of MLRA boundary lines.

Conclusions

	 In keeping with the subject of this conference “Biodiversity and

Management,” we hope that the materialsthat have been presented in

this paper will provide some insight in recognizing the opportunities

that exist for further, and broader in scope, collaboration between the

United States and Mexico.

	 Although the effort described in this paper demonstrates the potential for extending MLRAsinto Mexico itshould not be considered

definitive.Boundarylinesthatweredevelopedneedfurtherrefinement

and additional field evaluations. However the lines can be considered

to be a reasonably accurate first approximation, and can be utilized

with a fair degree of confidence. The potential for extending MLRA

boundariesfromtheUnitedStatesintoMexicoisfeasible andrealistic.

This would facilitate the direct use of ESDs across borders and improve exchange ofrangeland data between the two countries. Mexico

would have a direct benefit from the use of MLRAs as a valuable

and useable management tool through the direct use of ecological

site descriptions in range management planning. COTECOCA site

descriptionsshould not be overlooked.They represent descriptions of

all grazing sites within the country, and have been mapped on a state

basis. It is probable that Brown and Lowe maps can be effectively

applied in future efforts to extend MLRA boundaries in other parts

of Mexico.

	 This is an opportune time to work with Mexico in the refinement

of site descriptions. The NRCS is in a continuing process of revising

EcologicalSiteDescriptions andcomparisons couldbemadebetween

COTECOCA descriptions resulting in correlating the two systems.

Thiswould be valuable in strengthening the ecologicalinterpretations

of COTECOCA sites, and would allow for Mexican researchers,

rangeland managers, planners, and others to tap into an existing, rich

Figure 8—A portion of Chihuahua showing COTECOCA site mapping
388 USDA Forest Service Proceedings RMRS-P-67. 2013

Mann and others Potential for Extending Major Land Resource Areas into Northern Mexico

Figure 9—Adjustments made in projected MLRA boundaries in Sonora and Chihuahua.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

database that will increase the value and utility of a neglected, but

valuable resource. Utilizing MLRAs in Mexico would be a valuable

addition to international communications, research, and exchange of

data. MLRAs can serve as a common denominator for classifying

and framing technical data that currently is not being utilized. The

wheel does not have to be reinvented.

References

Brown, D.E., and C.H. Lowe. 1994. Biotic communities of the Southwest.

University of Utah Press, Salt Lake City, UT.

Comisión Tecnico Consultiva para la Determinación Regional del los Coeficientes de Agostadero. 1978. Chihuahua, Secretaria de Agricultura y

Recursos Hidraulicos.

Dyksterhuis, E.J. 1949. Condition and management of range land based on

quantitative ecology. Journal of Range Management. 2(3): 104-115. 

Fox, H. Dale, Mary R. Kidwell, Leonard J. Lane, and Mark A. Weltz. 1999.

MODSS’99 Conference; Brisbane, Australia 1-6 August 1999.

Gori, David F. and CarolynA. F. Enquist.January 2003.An assessment of the

spatial extent and condition of grasslands in central and southern Arizona,

Southwestern New Mexico and Northern Mexico. Prepared by The Nature

Conservancy, Arizona Chapter.

Heilman,Philip,JeffryStone,AliciaMelgozaCastillo,[andothers].May2000.

Bi-national cooperation for water management research on rangelands in

Chihuahua. (Unpublished).

Heilman, Philip, Alicia Melgoza Castillo, Roy S. Mann, [and others]. 2000.

A framework for cooperation across the U.S./Mexico border. Rangelands.

22(3).

MacEwen,Rebecca,RoyS.Mann,PhilipHeilman,[andothers].2005.Defining

boundaries across borders: A case study extending a major land resource

area into Mexico. In: Gottfried, Gerald J.; Gebow, Brooke S.; Eskew, Lane

G.; andEdminster,CarletonB., compilers.Connectingmountainislands and

desert seas: biodiversity and management of the Madrean Archipelago II.

ProceedingsRMRS-P-36.FortCollins,CO:U.S.DepartmentofAgriculture,

Forest Service, Rocky Mountain Research Station: 525-528.

Stone, J.J. and G.B. Paige. 2003. Variable rainfall intensity rainfall simulator

experiments on semi-arid rangelands. In: Proceedings; First Interagency

Conference on Research in the Watersheds; Oct. 27-30, 2003; Benson,

AZ: 83-88.

USDA Natural Resources Conservation Service. 1997. (Rev.1 2003) National range and pasture handbook. Washington, DC: U.S. Department

of Agriculture, Natural Resources Conservation Service, Grazing Lands

Technology Institute.

USDASoilConservationService. 1981.Land resource regions andmajorland

resource areas of the United States. Agric. Handb. 296. Washington, DC:

U.S. Department of Agriculture, Soil Conservation Service.

USDASoilConservationService. 2006.Land resource regions andmajorland

resource areas of the United States, the Caribbean, and the Pacific Basin.

Agric. Handb. 296. Washington, DC: U.S. Department ofAgriculture, Soil

Conservation Service.
USDA Forest Service Proceedings RMRS-P-67. 2013 389

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Into the Third Dimension: Benefits of Incorporating LiDAR 

Data in Wildlife Habitat Models

Melissa J. Merrick and John L. Koprowski

Wildlife and Fisheries Science, School of Natural Resources and the Environment, University of Arizona, Tucson

Craig Wilcox

USDA Forest Service, Coronado National Forest, Safford, Arizona

Abstract—LiDAR (Light detection and ranging) is a tool with potential for characterizing wildlife habitat 

by providing detailed, three-dimensional landscape information not available from other remote sensing 

applications. The ability to accurately map structural components such as canopy height, canopy cover, 

woody debris, tree density, and ground surface has potential to improve wildlife habitat models because 

animals interact and respond to three-dimensional habitat features. Prior to LiDAR, accurate measurements 

of structural features were difficult to obtain over large areas as other remote sensing data are based on twodimensional spectral responses. The Southwest harbors a large diversity of unique vegetation communities, 

each with an associated wildlife assemblage with various management needs. Managers can use LiDAR to 

accurately characterize vegetation and landscape structural characteristics for entire districts or management 

units. Data surfaces derived from the LiDAR point cloud can be readily incorporated into species-specific or 

multispecies habitat models. Although LiDAR has received much attention in characterizing forest structure, 

fewer studies (n = 29) have suggested or incorporated this technology to improve wildlife habitat models 

specifically. Herein we provide a review of current LiDAR applications in wildlife habitat models, provide 

future directions, and detail how LiDAR can increase our ability to represent the world that animals experience.

Introduction 

	 Managing landscapesfor diversity and persistence ofwildlife is dependentonourunderstandingofvegetative,structural,physiographic,

and bioclimatic needs of species and how these factors contribute to

vital parameterssuch as growth,survival, and coexistence (Morrison

and others 2006). Increasingly, managers are faced with the need to

monitor, conserve, and predict a diversity of plant and animalspecies

andtheirspecifichabitatrequirements overbroadspatialscales;scales

at which ground-based sampling is cost prohibitive, or infeasible due

to remoteness. The vegetative communities and associated faunas in

the Madrean Archipelago are extremely diverse, and harbor many

endemic and endangered species(Lasky and others 2011; Poulos and

others 2007),requiring substantial monitoring efforts overlarge areal

extents. To conserve and manage the unique biotic assemblages on

state and federallandswithin theMadreanArchipelago,improvement

ininventoryofspeciesdiversity,identificationofhabitatrequirements

for a variety oftaxa—fromarthropodsto largemammals—identification of areas of particular conservation importance, and continued

monitoring of these areas to ensure species viability and probability

of persistence would be beneficial. 

	 An important management tool is the ability to correlate the presenceof a specieswithparticularphysiographic andvegetative features

thought to be important components of the species’ habitat. Many

of these species-specific physiographic and vegetative features can

be characterized with remote sensing data and, therefore, mapped

over large areas to produce a habitat model (Lurz and others 2008).

Habitatvariablesinsuchmodels are typicallyrepresentedwithremote

sensing data obtained from passive sensorsintercepting wavelengths

of reflected light or heat from the earth’ssurface. These remote sensing products are only capable of representing the complexity of the

landscape in two dimensions and do not directly measure structure

(Vierling and others 2008; Lefsky and others 2002), which limits our

ability to accurately describe and predict wildlife-habitat relationships. Active sensortechnologies,such aslight detection and ranging

(LiDAR), can improve wildlife habitat models by more accurately

describing fine-scale, spatially explicit, three-dimensional structural

features related to animal use, occurrence, and reproductive success

(Vierling and others 2008) and measure these features across entire

management units, mountain ranges, or districts (Reutebuch and

others 2005). 

	 LiDARdataisgenerallycollectedfromalaser-emitterscannerlinked

to an accurate global positioning system and inertial measurement

unit (Reutebuch and others 2005); the resolution and quality of the

data depends on both the scanner and the pulse density (Evans and

others 2009;Reutebuch and others 2005). LiDARdata can be broadly

categorized into two classes depending on the type of sensor: largefootprintwave-formdata inwhich the pulse-return intensity overtime

is digitized, and small-footprint discrete return data (fig. 1) in which

the spatial coordinates at which each laser pulse intersects an object

are recorded (see Anderson and others 2005; Evans and others 2009;
390 USDA Forest Service Proceedings RMRS-P-67. 2013

Merrick, Koprowski, and Wilcox Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models

Lefsky and others 2002;Reutebuch and others 2005; andVierling and

others 2008 for excellentreviews). There are positives and negatives

to each method (Evans and others 2009) and both provide unique

structural information not available from spectral or thermal sensor

technology and can greatly improve our ability to classify landscape

features and describe how animals relate to them. Herein we provide

a review of current LiDAR applications in wildlife habitat models. 

Methods

Literature Review and Synthesis

	 We conducted a search of published literature referenced in the ScienceCitation Index viaThompsonReutersWeb of Science, accessing

all years through 30 March 2012. We used five search routines: (1)

wildlife habitat AND LiDAR in “Title,” (2) habitat AND LiDAR in

“Title,” (3) wildlife habitat in “Topic,” LiDAR in “Title,” (4) habitat

in “Topic,”LiDARin “Title,” and (5) habitatANDLiDARin “Topic.”

We tabulated the number of references returned in each search iteration. We selected references relevant to studies of wildlife-habitat

relationships for further review. For each selected reference, we

summarized main objectives, methodology, main conclusions, and

management implications eitherstated or implied. We also examined

whetherthe studywastaxa specific,whetherLiDAR-derivedvariables

were correlated with landscape features ofinterest and whether or not

incorporation of LiDAR-derived variables enhanced or significantly

improved predictions and classification. We compiled these data to

identify current uses of LiDAR in habitat models, commonly utilized

LiDAR-derived variables, and new ideas or emerging techniques. 

Results

	 Many studies refer to LiDAR as a tool potentially applicable to

wildlife habitat assessment, monitoring, classification, and prediction, as indicated by the substantial number of references returned

via our most general search criteria (fig. 2; habitat AND LiDAR in

“Topic,” n = 154), indicating the growing interest in and awareness

of LiDAR technology. Relatively few references explicitly mention

use of LiDAR in wildlife habitat applications (fig. 2; n = 18). However, a considerable subset of total referencesspecifically addressthe

applicability of LiDAR data to predict species presence, diversity,

reproductive performance, or for creating variables that represent

important structural features useful in wildlife habitat models. We

reviewed 59 studies relating the use of LiDAR data to wildlife

habitat characteristics and summarized some common research aims

(table 1). The stated aims included assessment of LiDAR’s ability to

identify or quantify organisms, predict species presence, diversity,

or performance, and predict physical and vegetative characteristics

associated with species presence.

	 Of the 59 publications that we reviewed, the majority (45 or 76%)

assessedthepredictive capabilityofLiDAR-derivedvariablesrelative

to field-based measurements; ofthese, all but two reported significant

relationships betweenLiDAR-derived andfield-basedmeasurements

(table 1). Just over half (31 or 53%) of the publications focused on a

particular species or group of organisms (e.g. breeding forest birds,

tree species, forest spiders, fish communities). Of these, all but two

publications focused on using LiDAR to describe important habitat

characteristics, animal relationships to structural features, and predicting occurrence based on these structural affinities. Avian habitat

relationships were the most common area of taxon-specific focus

(table 1; 13 or 42%). We detected no use of the term “LiDAR” within

publications from wildlife-specific journals such as The Journal of 

Wildlife Management or Wildlife Research.

	 Many LiDAR-derived variables are strongly correlated with standard, field-based habitat measures, allowing for accurate predictions

at multiple spatial scales (table 2; Hyde and others 2005; Lefsky

Figure 1—A visualization of discrete, multiple return LiDAR data: 

(A) LiDAR point cloud draped with georeferenced aerial imagery, 

(B) Height of LiDAR pulse returns above the ground showing all 

vegetation structure and density, (C) Height of LiDAR pulse returns 

≤2 meters above the ground showing shrub/sapling vegetation 

structure and density, (D) raster layer created from mean canopy 

height surface, 3 m pixel resolution.
USDA Forest Service Proceedings RMRS-P-67. 2013 391

Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models Merrick, Koprowski, and Wilcox

Figure 2—Number of published journal articles referencing LiDAR, habitat, and wildlife habitat 

returned from Web of Science search criteria including all years through 30 March 2012. 

Table 1—Summary of common research aims from 59 studies relating the use of LiDAR data to wildlife habiat characteristics

Stated aims # results % of stated aims Additional characteristics # studies

– Assess potential to identify or quantify organisms Predictive? 45

 based on structural association 3 3.3

– Assess potential to predict species performance 3 3.3 Significant relationship between LiDAR

 based on structural associations data and variable of interest? 43

– Assess potential to predict physical characteristics 8 8.7 Incorporate spectral data? 12

– Review or comment 9 9.8 Improvement over spectral data alone? 9

– Discuss applications 11 12.0 Multiscale? 10

– Assess potential to predict vegetation 

 characteristic 17 18.5 Taxa specific? 31

– Classification and mapping 17 18.5 Taxa considered:

– Assess potential to predict species presence or 

 diversity based on structural associations 24 26.1 amphibians 1

 reptiles 1

Total aims 92 mammals 2

 arthropod 4

fish 4

plants 6

 birds 13

and others 2002; Vierling and others 2008). Some variables, such as

canopy cover, tree or vegetation height, and canopy volume within

various height categories, were obtained directly fromthe point cloud

or waveform intensity, whereas other LiDAR-derived variables are

secondarily estimated based on the relationship between primary

LiDAR metrics and plot-based measurements such as basal area,

biomass, and leaf area index (table 2; Dubayah and others 2000;

Lefsky and others 2002; Ruetebuch and others 2005; Vierling and

others 2008). Still other LiDAR-derived variables provide novel

information that is difficult to obtain from field-based measurements

suchashighlyaccurate terrainmodels andsecondarilyderivedmetrics

ofslope, aspect, and rugosity (ameasure ofsurface height variability).

Information unique to LiDAR data includes measures of pulse-return

intensity, which can be used to distinguish among live and dead trees

(Bater and others 2009; Kim and others 2009) and identify ephemeral

wetlands beneath forest canopy cover (Julian and others 2009), as

well as vertical structural complexity and volume. Vertical structural

complexity and volume can then be used to model fine-scale differences in canopy use within bird communities (Clawges and others

2008), or specific nesting requirements of a single species (Goetz

and others 2010). Additionally, new variables may be developed for

describing vegetative composition, complexity, and physiographicvegetative associations based on ratios, linear relationships, or other

novel combinations, which may further refine our ability to describe
392 USDA Forest Service Proceedings RMRS-P-67. 2013

Merrick, Koprowski, and Wilcox Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models

Table 2—Examples of LiDAR-derived metrics and applications for incorporating LiDAR variables in wildlife habitat models. Primary 

LiDAR metrics can be obtained directly from the LiDAR point cloud. Secondary LiDAR derivatives must be modeled based on the 

relationship between primary metrics and ground-based training data. Referen ces include studies incorporating metrics and/or 

applications.

LiDAR-derived metric

Primary or 

secondary? Applications LiDAR method References

Canopy surface model Primary Height of trees or other features without 

terrain subtracted. Depict elevation of 

canopy surface above sea level. 

Discrete return, 

waveform

Andersen and others 

2005; Graf and others 

2009

Canopy cover/closure Primary Produced from canopy surface model, 

important in habitat models for many 

bird and forest obligate species, but also 

those that prefer open areas

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002; Hyde and others 

2005; Martinuzi and 

others 2009

Canopy/vegetation 

height model

Primary Height of trees or other features with 

terrain subtracted. Identify forest seral 

stages, habitat preferences for species 

with varying affinities for canopy, portions 

of the canopy, or stand age

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002; Hyde and others 

2005; Graf and others 

2009

Canopy/vegetation 

profiles

Primary “ Waveform Dubayah and Drake 

2000; Lefsky and others 

2002; Goetz and others 

2010

Canopy base height Primary Identify inhabitable canopy area when 

subtracted from canopy height

Discrete return, 

waveform

Andersen and others 

2005

Canopy volume Primary Identify differences in canopy structure 

among forest age classes, distinguish 

among canopy structural affinities of 

various species

Waveform Lefsky and others 2002

Coefficient of variation 

vegetation height

Primary Identify forest seral stages and be used 

with field data to predict occurrence of 

snags and woody debris

Discrete return Bater and others 2009

Digital terrain model 

(DTM)

Primary Delineate streambeds, create 

very accurate DEM, slope, aspect, 

ruggedness layers

Discrete return, 

waveform

Reutebuch and others 

2003; Graf and others 

2009

Digital Elevation Model Primary Elevation and other interpolated surfaces: 

slope, aspect, rugosity commonly used in 

habitat models

Discrete return, 

waveform

Goetz and others 2010; 

Martinuzi and others 

2009

Foliage height diversity Primary Parse LiDAR vegetation returns within 

height intervals to estimate shrub or 

foliage density, important for ground 

nesting birds or other species that 

respond to foliage density at varying 

heights

Discrete return, 

waveform

Clawges and others 

2008; Martinuzi and 

others 2009

Return intensity Primary Distinguish between live and dead 

biomass as dead trees,vegetation, and 

water have low return intensity compared 

to live trees and vegetation

Discrete return Kim and others 2010

Standard deviation 

of vegetation height/ 

mean absolute deviation 

height

Primary Index of stand structural complexity, age, 

seral stage, used to identify snags

Discrete return, 

waveform

Graf and others 2009; 

Martinuzi and others 

2009

Vertical distribution of 

canopy structure

Primary Estimate above-ground biomass, identify 

forest seral stage

Waveform Dubayah and Drake 

2000; Goetz and others 

2010
USDA Forest Service Proceedings RMRS-P-67. 2013 393

Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models Merrick, Koprowski, and Wilcox

Above-ground biomass Secondary Provides an estimate of forage availability 

and, when combined with multispectral 

band information, forage quality

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002; Hyde and others 

2005

Basal area Secondary Provides information on forest structure 

and density, an important variable in 

habitat models of forest-dwelling species

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002

Canopy complexity/

diversity

Secondary Describes the vertical complexity of 

vegetation structure, important for 

arboreal species

Waveform Lefsky and others 2002; 

Goetz and others 2010

Diameter Breast Height 

(DBH)

Secondary Provides information on stand age, tree 

maturity could serve as a proxy for food 

or nest site availability

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002

Leaf area index (LAI) Secondary Provides an estimate of forage availability 

and, when combined with multispectral 

band information, forage quality

Discrete return, 

waveform

Dubayah and Drake 

2000; Lefsky and others 

2002

Timber/vegetation 

volume

Secondary Identify habitable areas within tree or 

shrub cover

Discrete return, 

waveform

Dubayah and Drake 

2000; Clawges and 

others 2008

Vertical distribution ratio Secondary Differenctiate among areas with high 

and low understory canopy structure, 

important for species sensitive to 

understory vegetative cover

Waveform Goetz and others 2010

Table 2—Continued

important habitat features and predict presence of organisms on the

landscape (Vierling and others 2008).Below, we briefly examine four

application domains common to wildlife studies that have benefitted

from incorporation of LiDAR-derived variables; we also provide a

summaryofLiDAR-derivedvariables applicable tostudiesofwildlifehabitat relationships (table 2). 

Habitat Mapping

	 LiDAR’s ability to accurately characterize physical terrain and

vegetative structure can contribute to landscape classification efforts

(Lefsky and others 2002). Classification techniques often include

classification and regression trees, machine learning algorithmssuch

as Random Forest (Martinuzzi and others 2009), and image segmentation via object based image analysis routines (Arroyo and others

2010). Adding three-dimensional structure increases a classification

routine’sabilitytoresolveamongsimilarvegetativetypesandfine-scale

physiographic features. When structural information such as canopy

height, return intensity, and rugosity are combined with multispectral

data from satelliteslike Landsat ETM, QuickBird, IKONOS (Arroyo

andothers2010;Bradburyandothers2005;Clawges andothers2008)

or hyperspectral data, classification accuracy is improved. With sufficient training data, LiDAR can be used to accurately classify, map,

and model important structural habitat variables over broad spatial

scales (Bradbury and others 2005; Lefsky and others 2002). Resulting classifications or maps can then be incorporated into predictive

models of species presence or use of an area (Graf and others 2009;

Martinuzzi and others 2009). 

Predicting Pre sence or Use

	 The ability to predict an organism’s presence and monitor for its

continued persistence is a central goal among land managers and

wildlife biologists alike (Hyde and others 2005). Because LiDAR

data directly measuresstructural featuresthat animalsrespond to (i.e.

selectfor) athierarchicalspatialscales,LiDAR-basedparametershave

potential to improve the predictive capability of species distribution

and probability of use models (Bradbury and others 2005; Seavy and

others 2009). LiDAR-derived variables can be used to correlate finescale structural associations measured in the field and map these over

large spatial extents, indicating areas where a species or community

is likely to be found. For example, many species require dead trees

(snags)for nesting (bluebirds,woodpeckers, andmany tree squirrels),

and may only use an area if such features are available. LiDAR can

be useful in identifying or predicting the presence of snags, either

based on the tendency for olderforeststructuralstagesto harbor more

woody debris (Bater and others 2009; Martinuzzi and others 2009),

or by the change in the intensity of LiDAR pulse returns based on the

amount of dead wood in a plot (Kim and others 2009). Once maps of

snag-class presence or absence were created for a single mountain,

Martinuzzi and others (2009) used this information to predict the

mountain-wide probability of use for several bird species based on

their affinities for snags of particular sizes.

	 Availability of preferred microclimates can allow species to persist in otherwise extreme environments (Suggitt and others 2011).

LiDAR’s ability to accurately map fine-scale topographic features

has potential for creating variables based on species’ microclimate

needs, and thermally important areassuch assuitable sitesfor rearing

neonates and likely locationsfor ectothermburrows could bemapped

at broad scales.
394 USDA Forest Service Proceedings RMRS-P-67. 2013

Merrick, Koprowski, and Wilcox Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models

Correlates of Habitat Quality

	 Defining and improving habitat quality are topics of considerable

interest in the wildlife literature and major goals for efforts such as

land management, conservation, restoration, and endangered species

recovery. In reality, however, habitat quality is difficult to assess, especially with metrics meaningful to the organism of interest (Hinsely

and others 2006). Because of the difficulty in measuring structurally

complex landscapes, habitat quality isofteninferred frommeasures of

reproductive success within a defined area. If reproductive successis

stronglycorrelatedwithspecificphysiographicorvegetationstructural

features, then LiDAR can be used to define areas of predicted high

reproductive success and habitat quality. LiDAR-derived metrics of

canopy height were strongly correlated withmean chick bodymassin

Parus major over a 7-yearstudy, and the direction of thisrelationship

changed depending on springtime temperatures (Hinsley and others

2006). With this complicated interaction between climate, position in

canopy, and nestling success established, it is conceivable that mean

nestlingmass and inferred habitat quality could be predicted annually

over an entire forest based on LiDAR data and climate predictions.

Correlates of Biodiversity

	 Withthe threatof climate change andassociateddisturbance events,

there is an increased need forrapid biological assessments ofstate and

federallandstobothinventoryandmonitorspeciesdiversityandfocus

conservationeffortsonareas associatedwithhighbiodiversity(Bergen

and others 2009; Lesak and others 2011). Structural complexity and

heterogeneityinbothvegetationandphysiographyare associatedwith

higher levels of plant and animal diversity relative to less complex

sites (Bergen and others 2009). Therefore, LiDAR-derived variables

should aid in identifying areas of high biodiversity and in predicting

biodiversity of certain species assemblages based on their structural

affinities(Bergen and others 2009). Forest dwelling beetle diversity

and richness were correlated with LiDAR-derived measurements

of elevation and tree height (Müller and Brandl 2009). Songbird

species richness in deciduous forests was related to LiDAR-derived

measures of forest structure such as canopy and midstory height and

midstory density, with intra-guild species diversity (e.g. ground vs.

aerial foragers, edge vs. interior specialists) described by different

forest structural metrics (Lesak and others 2011).

Discussion

Proven Tool

	 LiDARtechnology has become increasingly accessible and affordable over the last 10-15 years and continues to improve, providing

valuable datasets that can be used by resource managers toward

a variety of applications, from forestry and watershed science, to

wildlife conservation and management over large spatial extents

(Lefsky and others 2002; Ruetebuch and others 2005; Vierling and

others 2008). While not all LiDAR data is equal (Evans and others

2009), it is evident that LiDAR technology and its various applications are no longer just theoretical. LiDAR technology has proven a

useful tool when applied to many wildlife-habitat studies, and will

continue to be refined. Every article we reviewed touted LiDAR’s

ability to quantify (often accurately) landscape and aquatic structural

features that organisms identify as habitat, to map fine scale habitat

associations over large areas, and to improve predictive models of

species presence, use, reproductive success, and overall biodiversity.

Combining LiDAR and multispectral datasets allows for maximal

exploration of species’ biotic and structural associations to more

adequately represent the real world that animalsinhabit and to which

they respond. 

Promising Future

	 In the MadreanArchipelago, LiDAR data collection has begun for

the Coronado National Forest, and LiDAR-derived metrics promise

to improve forest inventory and monitoring efforts, including habitat

assessment and modeling for a variety of wildlife species. The sheer

size and topographic and biotic diversity of national forests like the

Coronado are representative of the challenges this vast and complex

region poses in terms of conservation and management, and also

represent an opportunity to demonstrate the benefits of incorporating

LiDAR data to accurately describe, quantify, monitor, and conserve

its great biological diversity and natural resources.

Acknowledgments

	 We thank Brett Mitchell, Denise Laes, and Steven Dale at the

Remote Sensing Application Center for instruction on using LiDAR

data and facilitating discussions about potential uses ofLiDARon the

Coronado National Forest. Tom Mellin and BrianWakeling provided

helpful comments that significantly improved this manuscript. 

References

Andersen, H.; McGaughey, R.; and Reutebuch, S. 2005. Estimating forest

canopy fuel parameters using LIDAR data. Remote Sensing of Environment 94:441-449.

Arroyo, L. A.; Johansen, K.; Armston, J.; and Phinn, S. 2010. Integration of

LiDARandQuickBirdimageryformappingriparianbiophysicalparameters

and land cover types in Australian tropical savannas. Forest Ecology and

Management 259:598-606.

Bater, C. W.; Coops, N. C.; Gergel, S. E.; Lemay, V.; and Collins, D. 2009.

Estimation of standing dead tree class distributions in northwest coastal

forests using lidar remote sensing. Canadian Journal of Forest Research

39:1080-1091.

Bergen, K. M.; Goetz, S. J.; Dubayah, R. O.; [and others]. 2009. Remote

sensing of vegetation 3-D structure for biodiversity and habitat: Review

and implications for lidar and radar spaceborne missions. Journal of Geophysical Research 114:1-13.

Bradbury, R. B.; Hill, R. A.; Mason, D. C.; [and others]. 2005. Modelling

relationships between birds and vegetation structure using airborneLiDAR

data : A review with case studies from agricultural and woodland environments. Ibis 147:443- 452.

Bässler, Claus; Stadler, J.; Müller, J.; Förster, B.; Göttlein, A.; and Brandl, R.

2010. LiDAR as a rapid tool to predict forest habitat types in Natura 2000

networks. Biodiversity and Conservation 20:465-481.

Clawges, R.; Vierling, K.; Vierling, L.; and Rowell, E. 2008. The use of

airborne lidar to assess avian species diversity, density, and occurrence in

a pine / aspen forest. Remote Sensing of Environment 112:2064 - 2073.

Dubayah, R. O.; and Drake, J. B. 2000. Lidar remote sensing for forestry.

Journal of Forestry 98:44-46.

Evans,J. S.; Hudak,A. T.; Faux, R.; and Smith,A. M. S. 2009. Discrete return

Lidar in natural resources: Recommendations for project planning, data

processing, and deliverables. Remote Sensing 1:776-794

García-Feced, C.; Tempel, D. J.; and Kelly, M. 2011. LiDAR as a tool to

characterize wildlife habitat: California spotted owl nesting habitat as an

example. Journal of Forestry 109:436-443.

Goetz, S. J.; Steinberg, D.; Betts, M. G.; [and others]. 2010. Lidar remote

sensing variables predict breeding habitat of a Neotropical migrant bird.

Ecology 91:1569-76.
USDA Forest Service Proceedings RMRS-P-67. 2013 395

Into the Third Dimension: Benefits of Incorporating LiDAR Data in Wildlife Habitat Models Merrick, Koprowski, and Wilcox

Graf, R.; Mathys, L.; and Bollmann, K. 2009. Habitat assessment for forest

dwelling species using LiDAR remote sensing: Capercaillie in the Alps.

Forest Ecology and Management 257:160-167.

Hinsley, S.A.;Hill,R.A.;Bellamy, P.E.; andBalzter,H. 2006.The application

of Lidar in woodland bird ecology: Climate, canopy structure, and habitat

quality. Photogrammetric Engineering & Remote Sensing 72:1399-1406.

Hyde, P.; Dubayah, R.; Peterson, B.; [and others]. 2005. Mapping forest

structure for wildlife habitat analysis using waveform lidar: Validation of

montane ecosystems. Remote Sensing of Environment 96:427-437

Julian, J. T.; Young, J. A.; Jones, J. W.; [and others]. 2009. The use of local

indicators to improve LiDAR-derived predictions of potential amphibian

breeding ponds. Journal of Geographic Systems 11:89-106.

Kim, Y.; Yang, Z.; Cohen, W. B.; Pflugmacher, D.; [and others]. 2009. Distinguishing between live and dead standing tree biomass on the North Rim of

GrandCanyonNationalPark,USAusing small-footprintlidar data.Remote

Sensing of Environment 113:2499-2510.

Lasky, J. R.; Jetz, W.; and Keitt, T. H. 2011. Conservation biogeography of

the US-Mexico border: a transcontinental risk assessment of barriers to

animal dispersal. Diversity and Distributions 17:673-687.

Lefsky, M. A.; Cohen, W. B.; Parker, G. G.; and Harding, D. J. 2002. Lidar

remote sensing for ecosystem studies. BioScience 52:19-30.

Lesak, A. A.; Radeloff, V. C.; Hawbaker, T. J.; [and others]. 2011. Modeling

forest songbird species richness using LiDAR-derived measures of forest

structure. Remote Sensing of Environment 115:2823-2835.

Lurz, P.W.W.; Koprowski, J.L.; and Wood, D.J.A. 2008. The use of GIS and

modelling approaches in squirrel population management and conservation: a review. Current Science 95:918-922.

Martinuzzi, S.; Vierling, L. A.; Gould, W. A.; [and others]. 2009. Mapping

snags and understory shrubs for a LiDAR-based assessment of wildlife

habitat suitability. Remote Sensing of Environment 113:2533-2546.

Morrison,M.L.;Marcot,B.; andMannan,W. 2006.Wildlife-habitatrelationships:conceptsandapplications. 3rd ed.IslandPress,WashingtonDC.416p.

Müller, J.; and Brandl, R. 2009. Assessing biodiversity by remote sensing

in mountainous terrain: The potential of LiDAR to predict forest beetle

assemblages. Journal of Applied Ecology 46:897-905

Poulos, H. M.; Taylor,A. H.; and Beaty, R. M. 2007. Environmental controls

on dominance and diversity of woody plant species in a Madrean, Sky

Island ecosystem, Arizona, USA. Plant Ecology 193:15-30.

Reutebuch, S. E.;Andersen, H.; and McGaughey,R.J. 2005. Light Detection

and Ranging (LIDAR):An emerging tool for multiple resource inventory.

Journal of Forestry 103:286-292.

Seavy, N. E.; Viers, J. H.; and Wood, J. K. 2009. Riparian bird response to

vegetation structure:Amultiscale analysis usingLiDARofmeasurements

canopy height. Ecological Applications 19:1848-1857.

Suggitt,A.J.; Gillingham, P. K.; Hill,J. K.;[and others]. 2011. Habitatmicroclimatesdrivefine-scalevariationinextreme temperatures.Oikos120:1-8.

Vierling, K. T.; Vierling, L. A.; Gould, W. A.; [and others]. 2008. Lidar:

shedding new light on habitat characterization and modeling. Frontiers

in Ecology and the Environment 6:90-98.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
396 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 397

Grasslands
398 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 399

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Sustaining the Grassland Sea: Regional Perspectives 

on Identifying, Protecting and Restoring the Sky Island 

Region’s Most Intact Grassland Valley Landscapes

Gitanjali S. Bodner and Peter Warren

The Nature Conservancy in Arizona

David Gori, Karla Sartor, and Steven Bassett 

The Nature Conservancy in New Mexico

Abstract—Grasslands of the Sky Islands region once covered over 13 million acres in southeastern Arizona 

and adjacent portions of New Mexico, Sonora, and Chihuahua. Attempts to evaluate current ecological 

conditions suggest that approximately two thirds of these remain as intact or restorable grassland habitat. 

These grasslands provide watershed services such as flood control and aquifer recharge across the region, and 

continue to support dozens of species of concern. Prioritizing conservation interventions for these remaining 

grassland blocks has been challenging. Reliable data on condition and conservation value of grasslands in the 

region have not been systematically summarized. State and national boundaries further complicate efforts 

to identify where the best remaining habitats and populations of grassland obligate species still exist. We 

present results of an effort to merge grassland condition assessments, compile information on target species 

locations, and identify “priority grassland valley landscapes” across the region. We evaluate these priority 

landscapes in terms of the number of target species, critical threats, and enabling conditions for long-term 

conservation success such as activity by local cooperative groups dedicated to sustaining their landscapes. 

Lastly, we discuss the opportunities and challenges of designing and implementing effective restoration 

activities in these large multi-jurisdictional landscapes. 

Introduction

	 The grasslands of central and southern Arizona, southern New

Mexico, and northern Mexico, referred to here as Sky Island grasslands,formthe “grasslandseas” thatsurroundsmallforestedmountain

ranges, transitioning into desertscrublands as elevations drop, or into

foothill woodlands or chaparral as elevations rise. At the continental

scale, Sky Island grasslands form a unique setting where the large

blocks of Great Plains and Chihuahuan Desert grasslands east of

the Rockies and Sierra Madre Mountains spill over a low spot in

the Continental Divide. This ecotone geography and the ecological

gradients associated with “Sky Island mountains and grassland seas”

add tremendous floral and faunal diversity to the region as a whole

(McClaran and Van Devender 1997). Grassland valley landscapes

here are often recognized as distinct and important places by their

respective human communities,who self-organize to varying degrees

in order to benefit from and protect the ecological and economic

values of these places.

	 SkyIslandgrasslandshaveundergonedramaticvegetationchanges

over the last 130 years including encroachment by shrubs and trees,

loss of perennial grass cover and spread of non-native species(Humphrey 1987; Bahre 1991). The causes for these vegetation changes

have been the subject of debate and range from changes in regional

climate tohumanimpactsincludingpoorlymanagedlivestockgrazing

and suppression ofwildfires(Humphrey 1958;Buffington andHerbel

1965; Hastings and Turner 1965; Cable 1967; Wright 1980; Bahre

1985, 1995; Swetnam 1990; Brown and others 1997; McPherson and

Weltzin 2000).

	 Changes in grassland composition and structure have not occurred uniformly across the borderland region, nor are they solely

the consequence of past climatic events or human impacts but are

ongoing today (Archer 1989; Bahre 1991; Brown and others 1997;

Ceballos and others 2010). These changes, combined with increased

fragmentation due to exurban development and agricultural conversion, have caused significant declinesin grassland species, especially

wide-ranging ones.

	 Although the above studies, as well as others, have documented

change to grasslands locally, there have been only a few attempts to

characterize the extent of change or assess current grassland condition of borderland grasslands at broaderscales(Cox and Ruyle 1986;

Enquist and Gori 2008; Yanoff and others 2008).

	 In 2009, The National Fish and Wildlife Foundation (NFWF)

launched its Sky Island Grassland Initiative, a 10-year plan to protect

and restore grasslands and embedded wetland and riparian habitats

in the Sky Island region (fig. 1). The Initiative emphasizes grassland

restoration, protecting threatened land and water, and restoring
400 USDA Forest Service Proceedings RMRS-P-67. 2013

Bodner and others Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . .

populations of target wildlife species(NFWF 2009). The Foundation

anticipates investing millions of dollars to achieve these goals and

expects to leverage additional millions in federal, state and private

funds. The question is: where will this investment yield the greatest

returns in terms of restoring grassland health and recovering wildlife

species across the region?

	 This report attempts to answer this question at a regional scale

by integrating two recent spatial assessments of the historical extent

and current condition of Sky Island grasslands and savannas as well

as compiling information on the occurrence of grassland-dependent

wildlife and species of concern. Using this information and an

expert-based approach, we identify 12 priority landscapes where the

potential for restoring grasslands and recovering target species have

the greatest probability of success. We identify and evaluate these

priority landscapes based on their size, condition, landscape context, number of target species and natural communities, and human

enabling conditions for long-term conservation success. The latter

factor includes the existence of partnerships, such as local cooperative groups dedicated to sustaining their landscape; land protection

efforts; and ongoing ecological restoration and management.

Methods

Mapping Grassland Condition

Two recent assessments of historical grassland extent and current

condition form the basis of this Sky Island GrasslandAssessment and

cover most of the historical and remaining grasslands and savannas

in the Sky Island region: theApache Highland GrasslandAssessment

(AHGA; Gori and Enquist 2003) and the New Mexico Rangeland

Ecological Assessment (REA; Yanoff and others 2008). The AHGA

defined a series of grassland condition classes using expert opinion

Figure 1—Current condition of Grasslands in the Sky Island focal area.
USDA Forest Service Proceedings RMRS-P-67. 2013 401

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . . Bodner and others

and the peer-reviewed literature to define threshold values for shrub

cover. Mapping of these condition classes was done by 24 range

management specialists from federal and state agencies, academic

institutions and non-governmental organizations for the U.S. and a

combination of experts and LANDSAT satellite imagery analysisfor

Mexico.Mappingwas at approximately 1:100,000 to1:250,000 scale.

AHGAcondition classes were defined by percent woody plant cover,

compared to historical condition, and whether the grass species were

native,non-native,uplandorriparian(table1).TheAHGAconsidered

fire to be a major natural disturbance that controlled woody plants

and maintained grass dominance historically. The AHGA included a

comprehensive accuracy assessment using 234 field sampling points

in the U.S. portion of the Apache Highlands ecoregion. The overall

accuracy rate of the final map was greater than 77%.

	 The New Mexico Rangeland Ecological Assessment (REA) was

limited to southern New Mexico and includes the northeastern portion of the Sky Islands project area. Mapping was completed by 70

experts at approximately 1:23,000-1:100,000 scale.Condition classes

in the REA are vegetation states in the state-and-transition models

that accompanyNaturalResourcesConservationService’s ecological

site descriptions (NRCS 2009b). REA condition classes are similar

to those of the AHGA in that most altered classes reflect increases in

woody plant cover over historical condition. However, REA classes

are finer-scale, being associated with ecologicalsites, and stressrelative rather than quantitative woody cover, the distribution of grasses

between or under woody plants, grass composition dominated by

long-lived perennial grasses versussparse ruderal perennial or annual

species, and non-vegetative indicators such as soil erosion (table 1).

The REA did not map non-native grasses, as the spread of Lehmann

and Boer lovegrass into southwest New Mexico was limited before

2007 (Gori and Enquist 2003; Schussman and others 2006).TheREA

did not include an accuracy assessment.

	 For this analysis, the two assessments were combined into one

spatial dataset by grouping the finer-scale REA condition classes up

to AHGA classes (table 1). The combined spatial data set does not

identify grasslandsin Mexico outside oftheApache Highlands ecoregion. To represent these grasslands, we used the Instituto Nacional

de Estadística y Geografía (INEGI) land cover map that contains no

information on the current condition of grasslands or on the spatial

extent of former grasslands. Using the combined data set, we summarized the historical extent and current condition of grasslands and

savannas across the project area (fig. 1).

Mapping Sensitive Species and Natural 

Communities

ToassistintheidentificationandevaluationofSkyIslandgrasslands,

we summarized existing information from a variety ofsources on the

distributionofsensitivegrasslandspecies andriparian-aquatic species

that occur in wetland habitats embedded within grasslands (table 2).

ThesespecieswereidentifiedastargetsinNFWF’sSkyIslandBusiness

Plan (NFWF 2009).In addition, we summarized occurrence information for several natural communities that have declined significantly

(>90% from historical extent) over the last 100+ years due primarily

to human impacts: ciénega wetlands, sacaton riparian grasslands,

and Chihuahuan black grama grasslands(Hendrickson and Minckley

1984; Humphrey 1960; Yanoff and others 2008). Occurrence records

obtainedfromtheNaturalHeritageProgramsinArizona,NewMexico,

and Sonora were screened to identify grassland and riparian-aquatic

species. Natural Heritage data were not available for Chihuahua.

Table 1—Sky Island Grassland Assessment condition classes and the original AHGA and REA classes.

Original assessment class Source Sky Island assessment class

Native grassland and savanna with low woody 

 cover a AHGA No or low woody increase, native 

 grassland and savanna

Native grassland with or without ruderal 

 grasses REA “

Native savanna with or without ruderal grasses REA “

Bottomland sacaton (Sporobolus wrightii, S. airoides) AHGA

 grassland REA “

Woody-invaded native grassland and savanna with AHGA Medium woody increase, native 

 medium woody cover b grassland and savanna

Woody-invaded native grassland with or REA “

 without ruderal grasses

Woody-encroached native savanna with or REA “

 without ruderal grasses

Non-native grassland and savanna with low woody AGHA No or low woody increase, non-native 

 cover, where non-native perennial grasses, grassland and savanna

 Lehmann and Boer lovegrass, are common or dominant a

Non-native grassland and savanna with medium woody cover, AHGA Medium woody increase, non-native grassland

 where non-native perennial grasses, and savanna

 Lehmann and Boer lovegrass, are common or dominant b

Former grassland and savanna with high woody 

 cover c AHGA High woody increase, former grassland

 and savanna

Woody-dominated former grassland REA “

Woody-dominated former savanna REA “

Highly eroded former grassland REA “

Highly eroded former savanna REA “

a

Woody cover < 10%

b

Woody cover 10-35%, with cover of mesquite or juniper, Prosopis and Juniperus, <15% c

Woody cover >15% cover mesquite and juniper combined and/or >35% total woody cover; perennial grass cover always <3%; type conversion 

from grassland or savanna to shrubland.
402 USDA Forest Service Proceedings RMRS-P-67. 2013

Bodner and others Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . .

Tapping Expert Knowledge to Delineate and 

Evaluate Highlight Priority Landscapes

In the process of compiling the information above, we recognized

that these sources did not fully capture the collective knowledge that

we knew to exist for the region. In September 2010, we assembled

several experts to both expand our understanding of grasslands

across the region and to explicitly delineate focal areas. Participants

included: Angel Montoya, Peregrine Fund and Partners for Fish and

Wildlife; Dan Robinett, NRCS retired; Miles Traphagen, Turn of 

The Century Monitoring Inc.; and from The Nature Conservancy:

Gita Bodner, Dave Gori, Peter Warren, and Steven Yanoff, assisted

by Anne Bradley and Lara Miller.

	 Participants were asked to map areas they viewed as particularly

promising for sustaining the region’s grasslands over the long term,

drawing on their own knowledge and a variety of supporting information provided at the workshop. We seeded the discussion with

suggested criteria foridentifying, delimiting, and evaluating “priority

grassland landscapes” and with polygons used in the NFWFBusiness

Plan (NFWF 2009), but participants were free to map and describe

attributes of these delineated areas as they saw fit. Suggested criteria

includedsizeofgrasslandblocks, ecological conditionofthoseblocks,

presence of embedded streams and wetlands and target wildlife species, fragmentation versus connectivity of grassland habitat and of

native vegetation more broadly, presence of intractable threats, and

existence of conservation efforts such as local community partnerships. Spatial data sets provided to participants included information

on vegetation classifications, AHGA and REA grassland condition

assessments, soil classifications, species occurrence locations and

habitat models, land protection status, and results of various groups’

efforts to identify priority conservation areas for other purposes .

Oncepolygonsweredrawnandmetwithgroupagreement,participants

were asked to fill out a matrix of conservation value and feasibility

for each. Criteria evaluated included many of the same ones used

to inform the drawing of the polygons, but this exercise required

participants to rate criteria for each site as low, medium, or high and,

where necessary, explain those ratings. This was done as a group effort, with discussion of rankings along the way. Filling out the matrix

together also spawned discussions about the characteristics of each

grassland landscape, and about additional sources of information

about distributions of target species and communities.

Results and Discussion

	 Hereafter,wewillreferto“grasslands andsavannas” collectively as

“grasslands” to streamline the presentation and discussion of results.

Historic Distribution and Current Condition 

of Sky Island Grasslands

	 ExpertsintheUnitedStates andMexicoidentified13,902,000acres

as current or former grassland, which corresponds to almost half of

the area we analyzed (table 3; fig. 1). We assume that this represents

a conservative figure for the historic distribution and extent of grasslands in this area. Furthermore, grasslands were highly connected

historically, allowing wide-ranging species, like bison, pronghorn,

and grassland birds, as well as species with more limited dispersal

capabilities, like black-tailed prairie dogs, to move freely within and

between these habitats.

	 Only 2.6 million acres or 18.9% of these grasslands are currently

dominated by native grasses and relatively open and shrub free. By

comparison,shrubencroachmenthasoccurredonover8,159,000acres

or 58.7%of current and former grasslands.Approximately 4.9million

acres or 35.1% of current and former grasslands have experienced

more shrub encroachment but still fall into classes where this change

is considered reversible, for example, with prescribed fire or other

brush control methods. However, shrub cover and associated soil

erosion has exceeded a threshold on over 3.2 million acres, producing

a type conversion from grassland to shrubland on almost one quarter

of the historic grasslands and savannas in this region. Such areas are

considered difficultif notimpossible to return to open grassland states

Table 2--Spatial data collected and data sources for analysis of NFWF target and high-priority grassland and riparian-aquatic species 

occurrences in priority grassland landscapes.

Biological feature Data source

Ciénega Ciénega database and spatial layer developed by TNC-AZ 

 based on literature, agency reports, expert knowledge 

 and field reconnaissance 

Sacaton riparian grasslands Mapped by experts, AHGA (Gori & Enquist 2003; 

 Enquist and Gori 2008) and REA (Yanoff and others 2008)

Black grama grasslands Mapped by experts: historical extent and current condition on

 Sandy and Shallow Sandy ecological sites; REA 

 (Yanoff and others 2008)

Black-tailed prairie dog Database and spatial layer developed by TNC-AZ based on

 (Cynomys ludovicianus) colonies literature, agency reports & expert knowledge.

Pronghorn (Antilocapra americana) Spatial layer developed by TNC-AZ & NM based on Brown and 

 Ockenfels (2007), AZGFD (2010) and expert knowledge

Jaguar (Panthera onca) Spatial layer developed by TNC-NM and AZ based on

 agency reports and expert knowledge

Bison (Bison bison) Spatial layer developed by TNC-NM and AZ based on agency

 reports and expert knowledge 

Mexican gartersnake (Thamnophis eques megalops) Location information cited in Rosen and others (2001)

Sensitive grassland and riparian/aquatic species Spatial information on species occurrences from Natural 

 Heritage Programs in AZ and NM and Centro de Datos para 

 la Conservación de Sonora; sensitive grassland and riparian-aquatic 

 species and subspecies have global ranks of G1-G3 and T1-T3
USDA Forest Service Proceedings RMRS-P-67. 2013 403

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . . Bodner and others

on any substantial scale, though small patches might conceivably be

recovered with heavy investment of resources.

	 The spread of non-native perennial grasses within grasslands has

also been substantial(table 3).Boerlovegrass and, to a greater extent,

Lehmann lovegrass are common or dominant on at least 1.5 million

acres such that non-native grasslands with little to moderate woody

increase comprise 11% of this area’s current and former grasslands.

When theAHGAwas completed in 2003, non-native grasslands were

largely restricted to Arizona. However, Lehmann lovegrass appears

to be spreading into southwestern New Mexico asit is now present in

long-termmonitoring plots where it had not previously been recorded

(P. Sundt 2009). Implications of this spread are mixed, with some

wildlife species more impacted than others (Bock and others 1986;

Albrecht and others 2008). Native grasses remain present in many

invaded areas, albeit at lower density. Fire regimes and hydrology

can be affected but are not as radically transformed as in the case

with many plant invasions (Emmerich and Cox 1992).

Identification, Delineation, and Evaluation of 

Priority Landscapes

Participantslargely agreed on locations of “best” remaining grassland landscapes (fig. 2), Polygon boundaries were adjusted based on

additionalknowledgeaboutlocationsofvaluedfeatures(e.g.extending

the Janos polygon north into thePlayasValley to reflectmovements of

a wild bison herd) or ongoing conservation efforts (e.g. expansion of

Muleshoe-Aravaipa polygon to include an area known as the Bonita

grassland where landowners are combating shrub invasion). Experts

suggested addition of polygons (e.g. Buenaventura) to incorporate

areas that met many of the suggested criteria but whose values have

not been widely known among practitionersin thisregion. The group

alsorecommendeddroppingtwopolygons(SantaCruz andSanPedroU.S.) that had been previously delineated but rated low in grasslandspecific values (table 4). Both areas continue to harbor other habitats

of high conservation value, particularly riparian systems, which may

in turn benefit from grassland restoration activities. Nevertheless,

for the purposes of grassland conservation per se, these areas were

seen as being less likely than other landscapes to host large blocks

of grassland habitat into the future. Parts of the initial Santa Cruz

polygon that still have substantial grassland patches were added to

theAltarValley polygon,withwhich expertsidentified relatively high

landscape connectivity (an impression substantiated by themulti-year

movements of a wandering jaguar known as Macho B). In two cases,

participants chose to further refine polygon boundaries by examining

additional information aftertheworkshop.Overlaying draft polygons

on Google Earth proved especially useful for modifying boundaries

to exclude highly modified areas such as center-pivot agriculture in

Chihuahua.

	 The 12final polygonsincorporated almost allthe substantial blocks

of highest-qualitygrassland thatthe conditionassessments had identified within the region (83% of overall acreage in native no-to-low

woody increase classes) as well as substantial acreage of surrounding patches deemed restorable grassland (fig. 2). These polygons

also succeed at encompassing many of the other target biological

features associated with grasslands (fig. 3, table 5). For example, all

but two riparian sacaton grassland patches identified in grassland

assessments were included in final polygons. The two that were not

identified are relatively isolated from other grassland blocks, one being on the southwestern edge of the region where terrain changes to

highly dissected Sonoran scrub, and the other being on the northern

end where terrain rises towards the Mogollon Plateau. The latter is

near the Burro Cienega-Hachita polygon but is separated from it by

a large open pit mine.

	 Delineations of these major grassland landscapes largely conformed to units defined by individual valleys. This evaluation bore

out previous observations that the largest remaining blocks of open

grassland tend to be in mid-elevation valley bottoms(e.g. SanRafael,

Empire-Cienega-Sonoita). Grassy hills contribute substantial blocks

in some areas (e.g. Tumacacori and Sierrita portions of the Altar

polygon). In others (North Peloncillos, Southern Sulphur Springs,

Aravaipa-Muleshoe-Willcox), grassland habitat primarily occurs

on bajada or foothills terrain above lowlands that have undergone

vegetation and/or land use conversions. We applied the valley names

most commonly used by local communitiesto referto these grassland

landscapes. Participants discussed combining nearby polygons but

decided that the smaller units more accurately reflect differences

among nearby polygons in enabling conditions, e.g. in community

culture or development threats among nearby polygons.

	 In filling out the evaluation matrix for these polygons, participants

largely agreed on ratings (table 4). Where there was initial disagreement, further discussion was usually enough to bring the group to

consensus on a final rating. One exception came in trying to rate an

aspect of conservation feasibility thatreflectsthe influence that active

communityandpartnergroupshavebypromotingand/orimplementing

conservation measures in a given landscape. Participants agreed on

the importance of this factor, discussed activities of various partner

Table 3—General statistics for grasslands (U.S. and Mexico) in the Sky Island region.

Grassland type # Patches Total acres % All grasslands

No or low woody increase, native grassland 102 2,144,488 15.4

No or low woody increase, native savanna 48 478,953 3.5

Medium woody increase, native grassland and 

 savanna 138 3,212,191 23.1

No or low woody increase, non-native

 grassland and savanna 11 162,417 1.2

Medium woody increase, non-native grassland 

 and savanna 19 1,342,952 9.7

High woody increase, former grassland and 

 savanna 185 3,275,463 23.6

Mix of low and medium woody increase, native

 grassland and savanna 26 328,574 2.4

Undetermined grassland and savanna 5,764 2,956,650 21.3

Total U.S.-MX Grassland/Savanna 13,901,688
404 USDA Forest Service Proceedings RMRS-P-67. 2013

Bodner and others Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . .

Figure 2—Priority grassland landscapes, showing ecological condition.

groups andmodifiedsomepolygon boundariestoreflect partnerwork,

but ultimately decided that this factor could not be readily distilled

into a comparative rating.In all areas, participantsrecognized that(1)

additionalinformation orincreased conservation efforts could change

these ratings, (2) that all 12 polygons hold substantial promise for

conservation of grasslands and associated species, and (3) that both

opportunities and needs vary among the set.

	 Discussion among participants also drew out observations about

a number of ecological gradients present across the region, added to

information on targetspecies, described both threats and conservation

valuesofparticularlandscapesingreaterdetail, andaddedinformation

about partners and their conservation activities (elaborated in Gori

and others 2012).

Applications for Protection and Restoration

	 In each of these landscapes, successful grassland conservation

will need to include three key components: (1) strong partnerships

within the local community, which in turn support, (2) land protec-
USDA Forest Service Proceedings RMRS-P-67. 2013 405

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . . Bodner and others
406 USDA Forest Service Proceedings RMRS-P-67. 2013

Bodner and others Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . .

Figure 3—Priority grassland landscapes, showing select species and biological features of interest.

tion to prevent further fragmentation of the surviving grasslands, and

(3) active ecological management to restore and maintain grassland

health through time. These are addressed further in Gori and others

(2012), highlighted below to show the broad utility of information

presented here.

	 Overlaying polygons on land management highlights the role of

particular institutions in sustaining grasslands across the region and

within each individual landscape. For instance, these maps suggest

that if the U.S. Forest Service wants to invest in maintaining open

grasslands, this agency might focus on areas within the San Rafael

polygon; if the agency’s goal is to restore recoverable grassland,

working in the Altar-Tumacacori might provide the best opportunities. Of federal agencies, BLM has a particularly large role to play

acrossthe U.S. portion of thisregion. While each of these landscapes

includes a mix of public and private lands, success in landscapes

like the Empire-Cienega-Sonoita depends heavily on management

effects on BLM lands. By contrast, outcomes in landscapes such as

the Animas-San Luis Valley depend almost entirely on the actions of

private landowners.

	 Comparing polygons to maps of land management and protection

status (Gori et al 2012) highlights places where grassland blocks are

vulnerable to fragmentation by human infrastructure and disturbance,

aswell askeyopportunitiestoprotectlongtermconnectivityofnatural

vegetation for the many wildlife species that are affected by more

dramatic landscape changes such as proliferation of roads, houses,

mines, or intensive agriculture. Several jurisdictions, for instance,

combine to provide protection across nearly the entire San Rafael

Valley in the United States; adding easements or other conservation

status to the last remaining large block of unprotected private land

at the top of the valley would secure the ongoing value of these

previousinvestments. Successfully completing long-standing efforts

to bring publicly identified parcels of Arizona State Trust land into
USDA Forest Service Proceedings RMRS-P-67. 2013 407

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . . Bodner and others
408 USDA Forest Service Proceedings RMRS-P-67. 2013

Bodner and others Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . .

conservation status would greatly benefit five of the 12 priority grassland landscapes identified here. Conserving these State Trust lands

would, for example,safeguard the grassland block in the center of the

Empire-Cienega-Sonoita, and secure wildlife connectivity acrossthe

I-10 corridor, Catalina-Rincon Mountains, and the San Pedro River

to the Aravaipa-Muleshoe-Willcox landscape.

	 Overlaying polygons on maps of grassland habitat condition highlights key opportunities to protect or restore connections for species

such as pronghorn whose movements are affected by habitat changes

such asshrub encroachment (Brown and Ockenfels 2007). However,

for planning and implementing restoration or management interventions, more information is needed. Closer examination of Ecological

Site maps (Bestelmeyer and others 2009) can provide many insights

into conservation value and management options within landscapes.

Finer-scale mapping of ecological condition (e.g. by ecologicalstate)

can then informselection of project locations, treatmentmethods, and

monitoring (Tiller and others 2012).

	 Future effortsto gather information from additional experts would

nodoubtimprove the resolution, accuracy, andrichnessofthe regional

picture presented here. We recognize, for example, that challenges

of collecting data across state and international boundaries create inconsistent coverage forsome communities and species, and welcome

refinements. The large proportion of grasslands in Mexico currently

depicted as “unknown condition,” for example, highlights the need

to compile consistent information about these areas. We encourage

readersto treat this assessment, and otherslike it, asliving documents

to be modified and added to as time goes on, as ecological and social

conditions change, asspeciesrangesshift, and aswe learnmore about

all these factors.

	 In reviewing the status and trends of Sky Island grasslands, it is

clear that maintaining them as functioning systems represents one

of the greatest needs, and greatest challenges, of any conservation

issue in the borderlands. These grasslands are an essential link that

sustains the wildlife in the mountains and the perennial waters in the

valley bottoms. Yet successful grassland conservation is complex,

requiring simultaneous progress on several fronts: protecting the

land from fragmentation, managing the land to control shrubs and

maintain grass cover, and supporting community organizations and

other partnerships that get the work done. We hope that this analysis,

combined with efforts to make progress on these fronts and evaluations of what is working best, helps us all rise to this challenge.

Acknowledgments

	 We would like to thank the individualslisted above for expert input

during this analysis; Steven Yanoff for integrating the two grassland

assessment layers; Anne Gondor for help accessing spatial data for

Mexico; and the many, many experts who provided input into the

AHGAandREA. Funding forthis work was provided by the National

FishandWildlifeFoundation,TheNatureConservancy,PimaCounty,

and Freeport McMoran.

References

Albrecht,EricW.;Geiger,E.L.;Litt,A.R.;McPherson,G.R.;Steidl,R.J. 2008.

Fire as a tool to restore biodiversity in ecosystems dominated by invasive

grasses. Unpublished report on file at: U.S. Department of Defense, Legacy

Resource Management Program.

Archer, Steve. 1989. Have southern Texas savannas been converted to woodlands in recent history? American Naturalist. 134: 545-561.

AZGFD(ArizonaGame andFishDepartment).2011.Arizona statewidepronghorn management plan. 101 p. Report available online: http://www.azgfd.

gov/h_f/documents/2011%20Pronghorn%20Management%20Plan%20

-%20December%203%202011.pdf [April 27, 2012]

Bahre, Conrad J. 1985. Wildfire in southeastern Arizona between 1859 and

1890. Desert Plants. 7: 190-194.

Bahre,ConradJ.1991.Alegacyof change:historichumanimpactonvegetation

of the Arizona borderlands. Tucson: University of Arizona Press. 231 p.

Bahre, Conrad J., 1995. Human impacts on the grasslands of southeastern

Arizona. In: Mitchel P. McClaran; Van Devender, T.R., eds. The desert

grassland. Tucson: University of Arizona Press, Tucson, AZ: 230-226.

Bestelmeyer,BrandonT.;Tugel,A.J.;Peacock,G.L.;[andothers].2009.Stateand-Transitionmodelsforheterogeneouslandscapes:Astrategyfordevelopment and application. Rangeland Ecology and Management. 62(1): 1-15.

Bock, Carl E.; Bock, J.H.; Jepson, K.L.; Ortega, J.C. 1986. Ecological effects of planting

African lovegrasses in Arizona. National Geographic Research. 2 (1986):

456-463.

Brown, David E.; Ockenfels, R.A. 2007. Arizona’s pronghorn antelope: a

conservation legacy. Phoenix: Arizona Antelope Foundation. 190 p.

Brown, James H.; Valone, T.J.; Curtin, C.G. 1997. Reorganization of an arid

ecosysteminresponse torecent climate change.ProceedingsoftheNational

Academy Sciences. 94: 9729-9733.

Buffington, Lee C.; Herbel, C.H. 1965. Vegetational changes on a semidesert

grassland Range from 1858 to 1963. Ecological Monographs. 35: 139-164.

Cable, Dwight R. 1967. Fire effects on semidesert grasses and shrubs.Journal

of Range Management. 20: 170-176.

Ceballos, Gerardo; Davidson, A.; List, R.; [and others]. 2010. Rapid decline

of a grassland system and its ecological and conservation implications.

PLoS One 5:1-12.

Cox, Jerry R.; Ruyle, G.B. 1986. Influence of climatic and edaphic factors on

the distribution of Eragrostis lehmanniana Nees inArizona, USA. Journal

Grassland Society of South Africa 3: 25-29.

Emmerich,WilliamE.;Cox,J.R.1992.Hydrologycharacteristicsimmediately

after seasonal burning on introduced and native grasslands. Journal Range

Management. 45.5 (1992): 476-479.

Enquist,CarolynA.F.;Gori,D.F.2008.Applicationofanexpertsystemapproach

for assessing grassland statusin the U.S.-Mexico borderlands: implications

for conservation and management. Natural Areas Journal. 28(4): 414-428.

Gori, David F.; Enquist, C.A.F. 2003. An assessment of the spatial extent and

condition of grasslandsin central and southernArizona, Southwestern New

Mexico and Northern Mexico. The Nature Conservancy of Arizona. 28

p. Report available on line: http://azconservation.org/dl/TNCAZ_Grasslands_Assessment_Report.pdf [April 27, 2012]

Gori, David F.; Bodner, G.; Sartor, K.; Warren, P.; Bassett, S. 2012. Sky

Island grassland assessment: Identifying and evaluating priority grassland

landscapesfor conservation and restoration in the borderlands. 60 p.Report

available online: http://azconservation.org/downloads/category/grassland

[October 1, 2012]

Hastings,JamesR.;Turner,R.M.1965.The changingmile.Tucson:University

of Arizona Press. 317 p.

Hendrickson, Dean A.; Minckley, W.L. 1984. Ciénegas-vanishing climax

communities of the American Southwest. Desert Plants. 6(3): 130-175

Humphrey, Robert R. 1958. The desert grassland: A history of vegetational

change and an analysis of causes. Botanical Review 24: 193-253.

Humphrey, Robert R. 1960. Forage production on Arizona ranges. V. Pima,

Pinal and Santa Cruz Counties. University ofArizonaAgricultural Experiment Station Bulletin. 302: 1-138.

Humphrey,RobertR. 1987. 90 years and 535 miles: Vegetation changes along

the Mexican border.Albuquerque: University of New Mexico Press. 448 p.

McClaran, Mitchel P.; Van Devender, T.R., eds. 1995. The desert grassland.

Tucson: University of Arizona Press. 346 p.

McPherson, Guy R. 1995. The role of fire in desert grasslands. In: Mitchel P.

McClaran;VanDevender,T.R.,eds.Thedesertgrassland.Tucson:University

of Arizona Press: 130-151.

McPherson, Guy R.; Weltzin, J.F. 2000. Disturbance and climate change in

the United States/Mexico borderland plant communities: a state-of-theknowledge review. Gen. Tech. Rep. RMRS-GTR-50. Fort Collins, CO:

U.S. Department ofAgriculture, Forest Service,Rocky MountainResearch

and Experiment Station. 24 p.
USDA Forest Service Proceedings RMRS-P-67. 2013 409

Sustaining the Grassland Sea: Regional Perspectives on Identifying, Protecting . . . Bodner and others

Minckley,ThomasA.;Turner, D.;Weinstein, S.R., in review.The relevance of

wetland conservation in arid regions:Are-examination of vanishing climax

communities in the American Southwest. Journal of Arid Environments.

NFWF (National Fish and Wildlife Foundation). 2009. Sky Island grassland

business plan. 34 p. Report available online: http://www.nfwf.org/AM/

Template.cfm?Section=Wildlife_and_Habitat12andTEMPLATE=/CM/

ContentDisplay.cfmandCONTENTID=14106 [April 27, 2012].

NRCS (U.S. Natural Resources Conservation Service). 2009. Homepage of

the Ecological Site Information System, U.S. Department of Agriculture,

U.S. Natural Resources Conservation Service, [Online]. Available: http://

esis.sc.egov.usda.gov/ [April 27, 2012].

Rosen, Philip C.; Wallace, E. J.; Schwalbe, C. R. 2001. Resurvey of the

Mexican Garter

Snake (Thamnophis eques) in Southeastern Arizona. Unpublished report on

file at: Arizona Game and Fish Department and U.S. Fish and Wildlife

Service, Phoenix, Arizona. 64 p.

Sundt, Peter. 2009. Rangeland monitoring of the DiamondARanch: Western,

Central, and Eastern Divisions, 1993-2008. Unpublished report on file at:

The Nature Conservancy in New Mexico, Santa Fe, NM.

Schussman, Heather; Geiger, E.; Mau-Crimmins, T.; Ward, J. 2006. Spread

and current potential distribution of an alien grass Eragrostis lehmanniana

Nees., in the southwestern USA: comparing historical data and ecological

niche models. Diversity and Distributions. 12: 582-592.

Swetnam, Thomas W., 1990. Fire history and climate in the southwestern

United States. In: Krammes, J.S., tech. coord. Effects of fire management

of southwestern natural resources. Gen. Tech. Rep. RM-191. Fort Collins,

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Forest and Range Experiment Station: 6-16.

Wright, Henry A. 1980. The role and use of fire in the semidesert grass-shrub

type. Gen.Tech.Rep.INT-85. Ogden, UT: U.S. Department ofAgriculture,

Forest Service, Intermountain Forest and Range Experiment Station. 24 p.

Yanoff, Steven; McCarthy, P; Bate, J.; Miller, L.W.; Bradley, A.; Gori, D.

2008. New Mexico rangeland ecological assessment. 73 p. The Nature

Conservancy in New Mexico. Report available online: http://nmconservation.org/projects/rangeland_ecological_assessment/ [April 28, 2012].

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
410 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Sacaton Riparian Grasslands of the Sky Islands: Mapping 

Distribution and Ecological Condition Using State-andTransition Models in Upper Cienega Creek Watershed

Ron Tiller, Melissa Hughes and Gita Bodner

The Nature Conservancy in Arizona

Abstract: Riparian grasslands dominated by Sporobolus wrightii (big sacaton) were once widely distributed 

in the intermountain basins of the Madrean Archipelago. These alluvial grasslands are still recognized as key 

resources for watershed function, livestock, and wildlife. The upper Cienega Creek watershed in SE Arizona is 

thought to harbor some of the region’s most extensive sacaton stands. Documenting their extent and ecological state is important for informing management in this valley and for contributing to a clearer picture of this 

community’s status across the region. Our objectives were to map the distribution of sacaton; qualitatively 

assess stands of sacaton into ecological states; and test mapping and assessment methods for use in other 

valley bottoms in the region. We used a two-step approach: interpretation of aerial photography and soil maps 

followed by field reconnaissance. Field work consisted of qualitative, rapid assessments of ecological state 

using the Natural Resource Conservation Service’s State-and-Transition (S&T) models for the Loamy Bottom 

Ecological Site. The most open and productive state of Sacaton Grassland occupies 54 percent of alluvial 

habitats evaluated, with remaining acreage in various states of degradation, recovery, or transition to Mesquite 

Bosque woodland. Our observations in the Cienega Creek watershed suggest potential modifications to the 

S&T models that may more accurately reflect site potential, likelihood of transitioning to other states, and 

management strategies tailored to maintaining or improving sacaton grassland conditions across the region.

Introduction and Rationale

	 Ripariangrasslandsdominatedbybigsacaton(Sporobolus wrightii)

once formed nearly pure stands of robust perennial grass along the

Southwest’s perennial and intermittent streams (Bahre 1991; Brown

1982; Humphrey 1958). The sacaton riparian habitat is distinct from,

yet shares features with, neighboring riparian habitats and upland

grassland associations. Individual plants and small clumps of this

grass can be found in many settings. In the right soils and alluvial

settings, however, big sacaton, and to a lesser extent alkali sacaton

(S. airoides), creates a unique habitat formation sometimes called

a sacaton “flat” or “bottom” with distinct history and management

needs. Developing maps of sacaton grassland extent and ecological

condition can provide a foundation from which to develop informed

managementobjectives andactionsthatmaintain,restore, andpromote

the resilience of this important community.

Historical Geography

Declining Distributions

	 Sources estimate sacatongrasslandsnowoccupylessthan5percent

of their original distribution across the region (Humphrey 1960).

Commonly cited causes of decline include natural phenomena and

anthropogenic influences from as early as the late 1800s (Cooke and

Reeves 1976; Cox and others 1983; Humphrey 1958, 1960). With

channel entrenchment, reductions in overbank flooding, and the advent ofindustrialized farming, vast expanses of Sporobolus grassland

became ideally suited for agriculture and many thousands of acres

were converted to crops by the mid-1900s.

Current Geography and Condition

	 Despite steep declines,stands of bottomland sacaton grassland are

stillfound scatteredacrosstheirformerrangewiththe larger,more intact

examples most recognizable. Fine stands of big sacaton grasslands can

still be found in upper reaches of gently sloping valleyslike the upper

CienegaCreekbasin,upperBabocomariRiver,upperSanPedroRiver

in Mexico, and upper Santa Cruz River basin near Lochiel. Condition of sacaton flats in the region varies widely from mature stands

surviving on eroded former floodplain terraces or under the canopy of

mesquite woodlands, to exceptionally productive stands with little bare

soil or woody cover between plants that exceed 6 ft. in height.

	 These stands are recognized as key resources for watershed function, livestock, and wildlife and there is a growing interest in their

conservation and management inArizona. In the past 30 years, largescale riparian conservation areas have been establishedwithin several

watersheds that harbor extensive sacaton flats. Some of the finest

remaining examples exist on theLasCienegas NationalConservation

Area (LCNCA)managed bytheBureau ofLandManagement(BLM).

The BLM isresponsible for this and other conservation areas and has

been particularly active in seeking direction regarding the status of

these grasslands and guidance for management.
USDA Forest Service Proceedings RMRS-P-67. 2013 411

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

	 Limited efforts have been made to map the contemporary distribution of sacaton and associated communities in the Sky Island region.

These vary in their spatial extent and the degree to which they treat

sacaton flats as a distinct entity (table 1). We know of no efforts,

however, thatsystematically evaluate ecological condition ofsacaton

grasslands in a focal watershed, let alone the region. Our case study

examined alluvial habitats on the upper Cienega Creek watershed

(fig. 1) to (1) map the distribution of bottomland sacaton stands in

the study area; (2) qualitatively assign sacaton stands into ecological

states to understand dynamics and inform management; and (3) test

methodsformapping and evaluating sacaton communities elsewhere.

Methods

	 This mapping effort followed a framework laid out by the Natural

Resources Conservation Service (NRCS), first mapping distribution

of plant communities associated with particular soil and climate settings (Ecological Sites) (Briske and others 2006) and then mapping

“states” or ecological condition within those (Bestelmeyer and others

2010). To map sacaton bottomlands, we used a two-step approach:

interpretation of aerial photography and field reconnaissance. In

preparation for fieldwork, we digitized provisional polygons based

on aerial photos using three pieces of GIS information inArcMap 10:

(1) BLM’slayer of ecologicalsites mapped by NRCS specifically for

Las Cienegas (BLM 2003); (2) county soil maps and ecological sites

NRCS associated with these soils (soildatamart.nrcs.usda.gov); (3)

and, aerial photogaphs from the USDA’s 2007 National Agricultural

Imagery Program (NAIP; datagateway.nrcs.usda.gov). The first two

layers guided our focus on alluvial soils capable of supporting bottomland sacaton stands (fig. 1). The third layer provided 1-m (3-ft.)

resolution, 4-band digital aerial photographs. With the near infrared

band expressed to provide greater differentiation offeatures, we drew

polygons based on color, texture, and vegetation differences apparent

across the alluvial ecological sites. Some polygons extended beyond

soil map units to capture the full extent of target plant communities.

	 Field verification focused on areas with greatest concentration

of sacaton grasslands (fig. 2). Using handheld GPS units (Trimble

Juno SB) equipped with GIS (ArcPad 8), ground-level information

was recorded at one or more locations within a polygon depending

on the homogeneity or variability apparent between field maps and

on-the-ground observations. Polygons were adjusted as necessary

where sacaton stretched beyond or compressed within the boundaries

of digitized projections. Unique features within a polygon were also

catalogued. These included erosional features and other plant associations (e.g., tobosa or blue grama grasslands, cottonwood-willow

forest, and rabbitbrush (Ericameria scrublands).

	 To evaluate current condition ofsacaton standsin the field, we used

rapid-assessment qualitativedescriptionsofhabitattype,markedGPS

Table 1—Other vegetation mapping efforts with a variety of purposes and spatial extents that included upper Cienega Creek 

watershed.

Agency What and where they surveyed What they found in Cienega watershed

The Nature Conservancy (Gori and 

Enquist 2003)

Regional grassland condition survey 

combining expert knowledge and field 

reconnaissance.

723 acres of sacaton riparian grassland 

without condition evaluations.

Bureau of Land Management and 

NRCS (BLM 2003)

Empire and Empirita allotments and 

Las Cienegas NCA. Includes Cienega 

Creek and tributaties in an ecological 

site map for Las Cienegas.

7,600 acres we could classify as alluvial 

for ecological sites: Loamy Bottom, Sandy 

Bottom Swale, and Loamy Swale.

AZ Game and Fish Dept. (Kubly and 

others 1997)

AZ statewide riparian vegetation along 

the then-known extent of mapped 

perennial streams.

71 acres of sacaton, 384 acres of 

mesquite woodland.

USGS, AZ GAP vegetation mapping 

(Halvorson 2002)

AZ statewide riparian and upland 

vegetation communities; Cienega 

Creek, Empire Gulch, Gardner 

Canyon, and substantial upland 

areas outside of sacaton-dominated 

associations. This survey extended 

beyond strictly riparian stands.

4,680 acres of sacaton-scrub grassland

Pima County (Harris 2000) Riparian communities within Pima 

County; included portions of Cienega 

watershed.

9,290 acres of semi-desert grassland 

(sacaton and upland together BLP series 

143.1; Brown and others 1979), 427 acres 

of mesquite bosque (series 224.52), and 

149 acres of abandoned agricultural fields 

(presumably mixed former sacaton and 

cienega habitats).

Pima County, aerial reconnaissance 

(Fonseca 2000)

Pima County, including upper 

watershed of Cienega Creek beyond 

its borders into Santa Cruz County, and 

sites in Altar Valley, sacaton focus.

732 acres of sacaton grassland 

communities discriminated from upland 

grasslands, 2,909 acres of mesquitesacaton association, and 971 acres 

of cottonwood-willow forest-sacaton 

association.
412 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Figure 1—Areas of prospective sacaton grassland and mesquite bosque habitat in upper Cienega Creek watershed (beige 

lines), digitized from 2007 NAIP aerial imagery. These provisional stands are shown superimposed on alluvial Ecological 

Sites (colored polygons) previously mapped by NRCS and BLM.
USDA Forest Service Proceedings RMRS-P-67. 2013 413

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

Figure 2—Field verification of extent and condition: Lavender polygons represent areas field-verified and classified according to 

ecological condition as of September 2011. Purple dots illustrate point locations where field notes were recorded and voucher 

photographs of features and ecological condition were taken. This field work covered 3,732 acres of alluvial habitats in 560 polygons.
414 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

points, snapped voucher photographs, and assigned state categories

based on visual estimates of the following: sacaton grass cover

(both Sporobolus species were treated as equals), woody plant cover

(mesquite and other species), evidence of erosion, and mechanisms

affecting change (e.g., fire/drought, gully erosion, agricultural conversion) (table 2). Ecological condition categories corresponded to

states initially described in the NRCS Loamy Bottom Rangeland

State-and-Transition (S&T) models(rangeland: R041XC312AZ and

(forestland: F041XC310AZ). However, to more accurately reflect

what we found on the ground, we concluded it would be necessary to

refine this model by splitting certain statesinto phases orsub-statesto

reflect different site potential, likely transitional pathways, and need

for more specific prospective management strategies (fig. 3).

Results

	 Reviewing aerial imagery and soil data yielded 6,563 acres of

alluvial ecological sites with potential sacaton or mesquite bosque

communities. We field verified 3,874 acre of this digitized area (figs.

2 and 4). Of the area found to be sacaton rangeland, 72 percent had

previously been mapped as Loamy Bottom rangeland, 3 percent

Loamy Bottom forestland, 5 percent as Loamy Swale, and 8 percent

as Sandy Wash (R041XC316AZ). The remaining areas consisted of

upland soils (7 percent) and Sandy Bottom (5 percent; cottonwoodwillow forest; F041XC317AZ).

	 The more straightforward areas to delineate were core alluvial

habitatswhere sacatonormesquite communitiestraversedfloodplains

from side to side. The more difficult boundariesto outline were those

at the upper limits of sacaton distribution along the main stem of

Cienega Creek and its tributaries. In these areas, sacaton grassland

generallytransitionedtoBouteloua gracilis(bluegrama)orPleuraphis 

mutica (tobosa grass) grasslands in areas with finer surface soils, or

to xeroriparian habitats in areas with coarser soils (alluvial fans and

washes). At these outer reaches we made the greatest adjustments to

the final boundaries of sacaton and mesquite bosque habitats once

they were field verified.

	 Fieldsurveys enabledustoclassifypointlocations andfullpolygons

into states as shown in figure 5. Sample photographs are included in

figure6.USDA,NRCS=S&Tmodelswerehelpfulfor assigningstates

to rangelands/forestlands, but not all of our observed areasfell within

Table 2—Combined Loamy Bottom Ecological Sites (rangeland and forestland), 12-16 inch precipitation zone, in tabular format for quick field 

reference. Letters represent original models’ states; numbers represent revised model’s sub-state distinctions. An asterisk (*) denotes naming 

convention differs from Mesquite-Sacaton state described in Loamy Bottom rangeland S&T model.
USDA Forest Service Proceedings RMRS-P-67. 2013 415

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others Figure 3—Modified State-and-Transition model for MLRA 41-3 Loamy Bottom Rangeland and Forestland, Cienega, Sandy Wash, and Sandy Bottom Ecological Sites (12-16 inch precipitation zone) based on observations at Las Cienegas National Conservation Area. Our subdivisions of NRCS’s states are based on site capacity (soils and hydrology), divergent ecological dynamics, and on management implications for sacaton rangeland. This model also documents several connections among alluvial ecological sites, with transition pathways that are not well illustrated in the NRCS S&T models for Loamy Bottom. Note, for example, the transitional pathways that can transform a sacaton grassland into a mesquite bosque, both of which are highly prized by land managers and wildlife alike. 
416 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Figure 4—Ecological state map showing condition of sacaton stands (green through pink polygons) , plus extent of related 

riparian communities (orange, blue and red polygons with mesquite bosque, cienega, and cottonwood-willow forest respectively). 

Bright green represents highly productive, open sacaton stands in the condition closest to what is considered the “historic climax 

plant community “ (HCPC).
USDA Forest Service Proceedings RMRS-P-67. 2013 417

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

Figure 5A—Ecological state map, close up. Condition classifications (solid color polygons) for an example area that covers 

272 floodplain acres and shows most of the ecological states described in the NRCS State-and-Transition models for Loamy 

Bottom sites. Inclusions of Cienega and Sandy Bottom (cottonwood-willow forest) Ecological Sites occur here. Small circles are 

observation points from which polygons’ ecological states were assigned and voucher photographs were taken; some identify 

point features, e.g. eroding gullies, that occur within other polygons. 
418 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Figure 5B—Ecological state map using the modified model described in Figure 3. This classification shows finer distinctions in condition than are found among NRCS model states. Some model modifications call out distinctions that 

may be especially useful to managers, e.g. between actively eroding state E1 that may require restoration interventions 

versus stabilized erosion E2 that is healing on its own. Others reflect different dynamics observed in the field, e.g. 

sacaton-mesquite C1 with shrub form mesquite encroachment versus tree form mesquite in C2 on a more obvious 

trajectory towards mesquite bosque. 
USDA Forest Service Proceedings RMRS-P-67. 2013 419

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

Figure 6—Example photographs of ecological conditions of Loamy Bottom ecological sites (rangeland and forestland) 

encountered in the upper Cienega watershed. Text conforms to Ecological Site descriptions; class letter assignments 

are those of authors.

Sacaton Grassland (Historic Climax Plant Community; state A): 25-80% sacaton ground over, 0-20% annuals, <20 feet 

depth to water table. 

Sacaton Grassland (state B): 25-60% sacaton ground cover, 1-15% mesquite, <20 ft depth to water table.
420 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Sacaton-Mesquite (state C): 5-40% sacaton ground cover, 5-20% mesquite, other shrubs and succulents, <20 ft depth to 

water table. This state known as Mesquite-Sacaton in Loamy Bottom S&T model.

Exotics (state D) – Exotic perennial grasses (e.g., Johnsongrass, yellow bluestem, and bermudagrass) with/without mesquite, <20 ft depth to water table. Johnsongrass (Sorghum halepense) dominates a patch of former sacaton grassland in 

the upper third of photo. 
USDA Forest Service Proceedings RMRS-P-67. 2013 421

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

Eroded Sacaton (state E): 20-50% sacaton ground cover, no mesquite, exotic grasses present, gully erosion, >20 ft depth 

to water table. 

Annuals-Sacaton Grassland (state F): Trace sacaton cover, no mesquite, 0-10%other shrubs and succulents, annuals 

present, >20 ft depth to water table.
422 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Mesquite Bosque, native annuals (Historic Climax Plant Community; state H): no sacaton, 25-65% large mesquite, 0-5% 

other shrubs and succulents, 25-50 ft to water table.

existing conditions. Blending and refining models for rangeland and

forestlandsites andaddingourobservationsofsub-states andlinkages

between them produced an alternate model (fig. 3) we believe more

accurately represents what we mapped across Las Cienegas NCA.

We offer management suggestions for Loamy Bottom Rangelands

by state based on the revised S&T model, expertise of the authors,

and supporting literature.

	 Sacaton Grasslands(statesA, B1, B2, and B3) occupy about 2,156

acres here. While distributed along the length of the creek, these

stands rarely occupy the entire length or width of a broad alluvial

setting. Sacaton stands are not as homogeneous as anticipated; a

mosaic of other conditions occurs around states A, B1 and B2, including Sacaton-Mesquite and Mesquite Bosque (see states below).

Cienega and Sandy Bottom Ecological Sites also are found within

and around this mosaic. The S&T model and additional research

resultssuggest that Sacaton Grassland statesA, B1, and-B2 primarily

require low-intensity management designed to maintain high grass

cover and low shrub cover. Fire arguably has the greatest value for

managing big sacaton and alkali sacaton ranges (Britton and Wright

1983; Wright and Bailey 1982) provided that frequency, extent, and

seasonality of fire occurrence are considered for their overall health.

Grazing management may affect how sacaton plantsrespond to other

disturbances, with implications for the health of the grass stand. For

maximum forage production, Cox and others (1989) recommend

grazing big sacaton in the spring, not grazing in dry summers and

dry years, and discontinuing fall grazing. The combination of fire

and grazing produces very different effects compared to that of either

activity alone (Vogl 1974).

	 Sacaton patches with moderate cover and appreciable woody plant

encroachment are evident in three states. Sacaton Grassland (state

B3) occurs with low-medium cover of tall mesquite (>12 ft. tall) and

vigorously growing sacaton with moderate cover. Sacaton-Mesquite

conditions C1 and C2 are also evident (see below for C2). State C1

occurs with low-mediumcover ofshrubbymesquite (<12 ft. tall) with

low-medium cover of less vigoroussacaton. In condition B3,sacaton

appeared to co-exist with mesquite while in condition C1 the grass

appeared to be in decline. Both states would benefit from management aimed at reducing woody cover with minimal soil disturbance.

While mechanical grubbing is not recommended (Robinett personal

communication, 2011), studies show that woody plant control with

herbicides can significantly increase grass and forb production. Targeted applications to individual woody plants are best and might be

useful in integrated brush management programs followed with fire.

	 Sacaton-Mesquite (stateC2;knownasMesquite-Sacaton inLoamy

Bottom S&T model) stands with tree-form mesquite (>12 ft. tall)

appear to be in transition towards Mesquite Bosque (State H). This

transitionalpathwayrepresents animportant contribution ofthisstudy

since it is not articulated in existing Ecological Site Descriptions

(ESD) and their respective modelsfor Loamy Bottoms. Our observations suggest that a site-by-site evaluation would need to be made on

whether to control mesquite in these areas or allow them to transition to

bosque, a desirable and rare plant community in its own right.

	 Eroded Sacaton Grassland is evident in two phases at this site:

activelyeroding(E1) and healing/healed(E2).In thiswatershed,State

E1 appears as patches throughout the alluvial landscape, in areas of

headcuts and gullies or sheet erosion. Management actions might

focus on arresting and/or repairing erosion, for example, treating

headcuts that threaten to dewater high quality sacaton grasslands.

Treatments may include a combination of methods(e.g., Zuni bowls,

one-rock dams, and media lunas; Zeedyk and Clothier 2009). Sheet
USDA Forest Service Proceedings RMRS-P-67. 2013 423

Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . . Tiller and others

erosion areas vary from about one to several acre patches in size

interspersedwithin grasslands(A-B2) and sacaton-mesquite (B3-C2)

in the floodplains of Gardner Canyon and Empire Creek. Installation

of controls to slow overland flow and capture sediment may be recommended management actions. Additional measures may include

reseeding and/or re-vegetation with sacaton, or possibly with other

native grass species. Select areas might be rested from grazing to

enable re-establishment of plant cover on bare soils.

	 Eroded sacaton (state E2) occurs as healed or healing gullies or

sheet erosion.We observed areas offormer cutbankswith peeled back

walls well-vegetated with sacaton or occasional wetland plants. In

other places, we saw areas healing from sheet erosion where all but

the original sparsely vegetated shallow cutbank was visible. Since

state E2 is healing or healed of its own devices, we suggest no management actions.

	 Exotic perennials (state D) occur as small patches within states A

and B.Johnsongrass(Sorghum halepense) isthe most often observed

exotic perennial grass, rarely occupying more than an acre at thissite.

Johnsongrass and bermudagrass (Cynodon dactylon) are most often

associated here with cienega wetlands or along stream channels and

gravel bars ofSandyBottomecologicalsites.At present,these grasses

do not appear to threaten overallsacaton grassland condition since they

appear to be more or less confined to wetland and near stream sites.

	 Annuals (state F) is largely restricted to abandoned agricultural

fields.Thesefields arepartiallyrecolonizedbysacaton,butmore soby

mesquite. Small-scale earth-movingmightserve to promote recovery

of sacaton and expansion of extant cienega. Presently, surface water

from a cienega is precluded from reoccupying the southern portion of

the fields by short levees (3-5 ft.) that could be breached. Monitoring

the rate and direction ofsacaton and woody plant encroachment onto

these fieldsis a viable, low budget option for tracking the recovery of

the fields with or without modifications to the levee or canal system.

	 MesquiteBosque comprisesstate H. Most MesquiteBosque found

along Cienega Creek includes big sacaton plants as an understory

component. Mesquite bosques support several trophic levels and

great varieties of invertebrates, birds, bats and other animals because

of the abundance and nutritional quality of food and high structural

diversity. Considering wildlife values and Resource Management

Plan directives for Las Cienegas NCA to protect and maintain mesquite bosque, the best management strategy may be to leave these

woodlands alone regardless of how they came to be formed. Present

day environmental conditions clearly favor mesquite bosque over

sacaton grassland in some areas above, and most areas below, 4,300

ft. elevation.

Discussion

	 We produced an extensive GIS database for referencing our map

of on-site sacaton grassland conditions and their distribution. Sacaton

grasslandispresentonover2,100acres,supportingthe contentionthat

some ofthefiner examplesin southeastArizona occurin this drainage.

This acreage includes pure sacaton flats with those experiencing the

early-to mid-stages of mesquite encroachment. With little over half

of the verified alluvial areas found to be in the most productive states

(A-B2) of Sacaton Grassland, our observations speak to the timeliness of the BLMs request for information on the baseline condition

of sacaton communities and its desire to guide management.

	 Sacatoncommunities arerestrictedtolowgradient, alluvial environments yet, not entirely confined to what had been mapped as Loamy

Bottomecologicalsite.Weobservedappreciable sacatonstands(states

A-C1) in areas mapped as Loamy Swale and Sandy Bottom Wash

EcologicalSites.Possibilities astowhy thismay be include: precision

of ecological site mapping in the mid-1990s was coarse enough that

some sacaton grassland communities plot in Loamy Swale or Sandy

Wash ecologicalsites; and, broad areas mapped as Loamy Swale and

Sandy Wash are actually transitional between these ecological sites

and Loamy Bottom.

	 Alkali sacaton (S. airoides) commonly occurs in states A-B3 of

upper Cienega Creek watershed; it is even the dominant perennial

grass in some areas. Cox (1984, 1985) reports that S. wrightii occurs

on moderately alkaline and slightly calcareous soils, and is usually

replaced by S. airoidesin areaswith very alkaline orsaline soils.Since

the S&T model for Loamy Bottom specifically addresses S. wrightii 

with S. airoides playing only a supporting role (Robinett 2005a), it is

possible that some of the area previously mapped as Loamy Bottom

is actually Saline Bottom (Womack 2005) or a combination of the

two.Our field verifications did not include digging soil pitsto validate

Ecological Site classification at point locations, but we recognize this

is a valuable effort when feasible.

	 The dynamics of change within sacaton grasslands, and the factors

drivingthesechanges,deserve additionalvalidationandresearch.Field

observations and review of the literature for this project suggested

transitional pathways between sacaton grasslands and other riparian

vegetation communities. These are addressed more fully elsewhere

(Tiller and others 2012). Sacaton Grassland and Mesquite Bosque

states transition between each other along successional trajectories.

Oneormore linkagesmightbemadebetweenrangelandandforestland

ecological sites, allowing them to flow fluidly between one another.

	 The vast majority of mesquite bosque stands of the upper Cienega

Creek watershed include big sacaton plants as an understory component, suggesting that these woodland communities may be a product

of recent environmental change (i.e., past 130 years; Bryan 1928).

Sacaton growing in the understory of these woodlands is most likely

a remnant of a former grassland condition and may persist in part

where water tables remain high (Lacey and others 1975). Mesquite

Bosque with trace or no sacaton understory is observed primarily at

the mouth of upland swalesthat drain to sacaton grassland communities via side drainages. Many of these bosques also may be recent in

occurrence since above 4,300 ft. elevation they are not visible in the

earliest aerial photography captured in the mid-1930s.

	 As the field survey and data refinement phases of this project

matured, modifications to improve efficiency in the future became

apparent. Creating dropdown menusfor field recordings using a suite

of site descriptors would increase the consistency and reduce postprocessing time by integrating field data directly into a GIS database.

Future efforts with limited time for field visits might be best served

by reducing field time spent classifying transitional areas and use

aerial imagery to emphasize work in larger areas of specific interest.

	 Our case study represents a substantial contribution to the field

because it provides(1) amore accuratemap ofwhatsacaton rangeland

occurs and its ecological condition at this site; (2) a more inclusive

modelforunderstandingdynamics andmanagementneedsofsacaton,

and related communities, across the region; and (3) refined methods

for evaluating the distribution and condition of sacaton communities

elsewhere. At present, the development of ESDs and S&T models is

moving into a field validation phase (Dan Robinett, personal communication, 2011b). This case study has the potential to aid in the

refinement of these models for use in other sacaton bottomland sites

across the Sky Island region.

Acknowledgments

	 We thank the following individuals for their help with this paper:

Daniel Morgan (AmeriCorps/Southwest Conservation Corps), Amy
424 USDA Forest Service Proceedings RMRS-P-67. 2013

Tiller and others Sacaton Riparian Grasslands of the Sky Islands: Mapping Distribution and Ecological Condition . . .

Markstein(ChicagoBotanicGarden),DanielDriscoll,ChrisJarchow,

and Samantha Hammer (volunteers) provided field support. BLM

staff Leslie Uhr, Marsha Radke, and Eric Baker provided technical

support including loaning us Juno SB GPS units. TNC staff Karla

Sartor,DanMajka,RobMarshall,DorothyBoone helpedwith project

design, logistics, and interpretation. Special thanks to Dan Robinett

(Robinett Rangeland Resources, LLC) and Julia Fonseca (Pima

County) for technical review of this paper. Funding was provided by

TNC and by grants from the National Fish and Wildlife Foundation,

Pima County, Freeport MacMoRan, and the BLM.

References

Bahre, C.J. 1991.Alegacy of change: Historic human impact on vegetation in

the Arizona borderlands. The University of Arizona Press, Tucson. 231 p.

Bestelmeyer,BrandonT., Mosley, K., Shaver, P.L., Sanchez, H.,Briske, D.D.,

Fernandez-Gimenez, M. 2010. Practical Guidance for Developing Stateand-Transition Models. Rangelands. 32(6):23-30.

Briske, D. D., S. D. Fuhlendorf and F. E. Smeins. 2006. A unified framework

for assessment andapplicationof ecologicalthresholds.RangelandEcology

and Management 59:225-236.

Britton, C.M. and H.A. Wright. 1983. Brush management with fire. Pages

61-68 inMcDaniel, K.C., ed. Proceedings: brushmanagementsymposium;

1983 February 16; Albuquerque, NM. Society for Range Management,

Denver, CO.

Brown,D.E.,C.H.Lowe, andC.P.Pase. 1979.Adigitized classification system

for the Biotic communities of NorthAmerica, with community (series) and

associated examples for the Southwest. Journal of the Arizona-Nevada

Academy of Science 14: 1-16.

Brown, D.E. 1982. Biotic communities of the American Southwest-United

States and Mexico. Desert Plants 4:1-342.

Bryan, K. 1928. Change in plant association by change in ground water level.

Ecology 9(4):474-478.

Cooke, R. U. and R. W. Reeves. 1976. Arroyos and environmental change in

the American southwest. Oxford: Clarendon Press.

Cox, J.R. 1984. Shoot production and biomass transfer of big sacaton (Sporobolus wrightii). Journal of Range Management 37(4):377-380.

Cox,J.R. 1985.Above-ground biomass and nitrogen quantitiesin a big sacaton

(Sporobolus wrightii)grassland.JournalofRangeManagement38:273-276.

Cox,J.R. 1988. Seasonal burning andmowing impacts on Sporobolus wrightii 

grasslands. Journal of Range Management 41(1):12-15.

Cox, J.R., R.L. Gillen, and G.B. Ruyle. 1989. Big sacaton riparian grassland

management: seasonal grazing effects on plant and animal production.

Applied Agricultural Research 4(2):127-134.

Cox, J.R., H.L. Morton, J.T. LaBaume, and K.G. Renard. 1983. Reviving

Arizona’srangelands.Journal of Soil and Water Conservation 38:342-345.

Fonseca, J. 2000. Map Guide: Sacaton Grasslands Special Element. Sonoran

Desert Conservation Plan.

Gori, D.F., and C.A.F. Enquist. 2003.An a ssessment of the spatial extent and

condition of grasslandsin central and southernArizona, Southwestern New

Mexico and Northern Mexico.

Halvorson,W. L., K.Thomas, L. Graham, M R. Kunzmann, P.Bennett,C.Van

Riper,C.Drost. 2002.TheArizonaGapAnalysis Project finalreport.USGS

Sonoran Desert Field Station, University of Arizona. Research Performed

Under: Cooperative Agreement No.14-45-0009- 1580, 14-45-0009-1054,

No.14-45-0009-94-1069, KR9-1165-CIV, andArizona CollectionsAgreement HAB 96-117.

Harris, L.K., J.A. Wennerlund, and R.B. Duncan. 2000. Riparian vegetation

mapping and classification, Sonoran Desert Conservation Plan. Pima

CountyGovernment,Tucson,AZ.Contract #07-30-H-127196-0100. 56 pp.

Humphrey, R. R. 1958. The desert grassland: a history of vegetational change

and an analysis of causes. Botanical Review 24: 193-253.

Humphrey, R. R. 1960. Forage production on Arizona ranges: V. Pima, Pinal

and Santa Cruz Counties. University of Arizona Agricultural Experiment

Station Bulletin 302:1-x.

Kubly, D.M., R.A. Winstead, L.J. Allison, C.R. Wahl, S.R. Boe, and J.A.

Wennerlund (editors). 1997. Statewide riparian inventory and mapping

project (executive summary). Nongame Technical Report 111 & 112.

Arizona Game and Fish Department, Phoenix, AZ.

Lacey, J. R., P. R. Ogden, and K. E. Foster. 1975. Southern Arizona Riparian

Habitat: Spatial Distribution and Analysis. University of Arizona Office

of Arid Lands Bulletin 8:1-148.

Robinett, D. 2005a. Ecological site description for Site ID: R041XA312AZ,

Sporobolus wrightii(big sacaton),LoamyBottom, 12-16 inch precipitation

zone, Major Land Resource Area 041 - Southeastern Arizona Basin and

Range. U.S. Department of Agriculture, Natural Resources Conservation

Service.Online:http://esis.sc.egov.usda.gov/esis_report/fsReport.aspx?id=

R041XC312AZ [Accessed 10-2011].

Robinett, D. 2005b. Ecological site description for Site ID: F041XC310AZ,

Prosopis velutina - Prosopis glandulosa var. torreyana / / Sporobolus 

wrightii (velvet mesquite - western honey mesquite / / big sacaton), Loamy

Bottom 12-16 inch precipitation zone, Major Land Resource Area 041 -

Southeastern Arizona Basin and Range. U.S. Department of Agriculture,

Natural Resources Conservation Service. Online: http://esis.sc.egov.usda.

gov/esis_report/fsReport.aspx?id= R041XC310AZ [Accessed 10-2011].

Robinett, D. 2011a. [Personal communication]. Verbal communication with

Ron Tiller. D. Robinett isretired, USDANational Resources Conservation

Service, Tucson, AZ.

Robinett,D.2011b.[Personal communication].LettertoRonTiller,December

11, 2011. On file at: The Nature Conservancy, Tucson, AZ.

Stromberg, J.C. 1993. Element Stewardship Abstract: Sacaton Grasslands.

Tucson, AZ: The Arizona Chapter of the Nature Conservancy.

Tiller, R.; Hughes, M.; Bodner, G. 2012. Sacaton riparian grasslands: Using

state-and-transition models to map distribution and ecological condition

in the upper Cienega Creek Watershed. Tucson, AZ: The Arizona Chapter

of the Nature Conservancy.

U.S. Department of Agriculture, Natural Resources Conservation Service.

2008. Soil Survey Geographic (SSURGO) database for Santa Cruz and

Parts of Cochise and Pima Counties, Arizona (AZ667). Fort Worth, Texas.

U.S. Department of Agriculture, Natural Resources Conservation Service.

2005. Ecological Site Inventory System (ESIS). Major Land Resource

Area 41-3. Online: http://esis.sc.egov.usda.gov.

U.S. Department of Interior, Bureau of Land Management (BLM). 2003. Approved Las Cienegas resource management plan and record of decision.

Arizona State Office and Tucson Field Office.

U.S. Department of the Interior, Fish and Wildlife Service (USFWS). 2002.

Biological and conference opinion summary, effects of the proposed Las

Cienegas National ConservationArea Resource Management Plan in Pima

and Santa Cruz counties, Arizona; October 4, 2002. Phoenix, AZ: U.S.

Department of the Interior, U.S. Fish and Wildlife Service.

U.S. Geological Survey (USGS) and U.S. National Park Service (NPS). 2000.

Vegetation Mapping Program. Online: http://biology.usgs.gov/npsveg/.

[Accessed Dec. 2007].

Vogl, R.J. 1974. Effects of fire on grassland. In: Kozlowski, T.T.; Ahlgren,

C.E., (eds.). Fire and ecosystems. New York: Academic Press: 139-194.

Womack, D. 2005. Ecological site description for Site ID: F041XC315AZ,

alkali sacaton - inland saltgrass (Sporobolus airoides - Distichlis spicata),

Saline Bottom 12-16 inch precipitation zone, Major Land Resource Area

041-SoutheasternArizonaBasinandRange.U.S.DepartmentofAgriculture,

Natural Resources Conservation Service. Online: http://esis.sc.egov.usda.

gov/esis_report/fsReport.aspx?id= R041XC315AZ [Accessed 10-2011].

Wright, H.A.; Bailey, A.W. 1982. Fire ecology: United States and southern

Canada. New York: John Wiley & Sons. 501 p.

Zeedyk, B.; Clothier, V. 2009. Let the water do the work: Induced meandering, an evolving method for restoring incised channels. Washington, DC:

Island Press. 240 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 425

Human Impacts
426 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 427

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Long Distance Commutes by Lesser Long-Nosed 

Bats (Leptonycteris yerbabuenae) to Visit Residential 

Hummingbird Feeders

Debbie C. Buecher

Buecher Biological Consulting, Tucson, Arizona

Ronnie Sidner

Ecological Consulting, Tucson, Arizona

Abstract—Each spring, thousands of female lesser long-nosed bats (Leptonycteris yerbabuenae) migrate 

from southern Mexico to northern Sonora and southern Arizona to have their young and take advantage of 

seasonably available forage resources, including nectar, pollen, and fruit of columnar cacti. Once the pups 

are volant, the population begins to disperse across the grasslands of southeastern Arizona, foraging on 

flowering paniculate agaves (Agave palmeri) and day-roosting in Madrean Sky Islands during their seasonal 

migration back to southern Mexico. Although the grasslands surrounding Arivaca, Arizona, have very low 

densities of naturally occurring agaves, nectar bats are documented visiting area hummingbird feeders to 

obtain sugar water. To better understand foraging patterns by these bats in an area with few natural food 

plants, we radio-tracked 28 L. yerbabuenae during 2010 and 2011. We captured bats at area hummingbird 

feeders and tracked them to a previously unknown day-roost in the Santa Rita Mountains, approximately 40 

km away. We discovered that bats nightly performed a long-distance commute from the Santa Rita Mountains 

to Arivaca, bypassing hummingbird feeders and naturally occurring agaves closer to their day-roost. It may 

be that the large colony size in the day-roost necessitates long-distance dispersal as a mechanism to reduce 

competition for limited food resources.

Introduction

	 The lesser long-nosed bat (Leptonycteris yerbabuenae) was listed

as endangered in the United States by the U.S. Fish and Wildlife

Service (USFWS)in 1988 (Shull 1988).The northernmost population

of this species is migratory, while a southern population near Jalisco,

Mexico, appearsto be present year-round (Ceballos and others 1997;

Cole and Wilson 2006). Female L. yerbabuenae are pregnant when

they arrive in southernArizona from central Mexico duringApril and

May. These bats are highly gregarious and use a limited number of

roosts with colonies numbering in tens of thousands of individuals.

They give birth (typically inMay) atlargematernity roostsinSonoran

desertscrub habitat (Cockrum and Petryszyn 1991), feeding nightly

on the nectar, pollen, and fruit of columnar cacti, including saguaro

(Carnegia gigantea) and organ pipe (Stenocereus thurberi) (Fleming and others 1993). During July and August, once the young are

volant (able to fly), maternity colonies begin to disperse. Wilkinson

and Fleming (1996) suggested two distinct migratory routes for the

species, one along the west coast of Mexico and the other along the

SierraMadreOccidentalwithlittlegeneticmixingbetweenthegroups.

However, more recent DNAanalysis determined that there is general

genetic mixing acrosssouthernArizona and, for conservation efforts,

the two groupsshould be considered one population (Ramirez 2011).

The bats begin a gradualmigratory path eastward,moving up in elevation and shifting to grassland communities where they feed on nectar

and pollen of flowering paniculate agaves, primarily Palmer agaves

([Family Agavaceae, Agave palmeri]; USFWS 1997). Ober and others (2005) radio tracked L. yerbabuenae in the Huachuca Mountains

of southeastern Arizona and determined that these bats used home

ranges with a greater density of flowering agaves (3.6 flowering

agaves/ha) than the general landscape (1.8 flowering agaves/ha). In

addition, nectar bats have discovered that hummingbird feeders offer energy resources during their migration across southern Arizona.

Some researchers have speculated that nectar bats visit artificial food

resourcesin greater numbers during years ofreduced agaveflowering,

particularly after periods of below average winter rainfall (Krebbs

2007, unpublished paper). Both L. yerbabuenae and Mexican longtongued bats(Choeronycteris mexicana) are documented using hummingbird feeders near Portal, Arizona, (Chiricahua Mountains) and

in the Santa Rita Mountainssince the 1970s(Cockrum and Petryszyn

1991).Recently, urban homeswellwithinTucson,Arizona, city limits

have multiple feeders drained each evening between mid-August and

late September (D. Buecher, personal observation 2011; Lowery and

others 2009).

L. yerbabuenae require caves and mine tunnels with specific temperature and relative humidity for day-roosts, as well as bridges and

various abandoned man-made structures, for night roosts (Cockrum

and Petryszyn 1991).Appropriate day roosts may be a limiting factor
428 USDA Forest Service Proceedings RMRS-P-67. 2013

Buecher and Sidner Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) 

for the distribution of L. yerbabuenae because they are documented

flying 24 to 40 km nightly from a day roost to suitable foraging

habitat (Horner and others 1998; Sahley and others 1993). Ober and

others (2005) found that this species has large home ranges (505 ha),

which allows them to access food resources across vast landscapes.

Disturbance and/or loss of roost sites, combined with a gradual decrease in foraging habitat from increasing land development, are the

most serious issues L. yerbabuenae face in Arizona (USFWS 1997).

Roostsin abandonedmine tunnels, caves, or old abandoned buildings

in remote areas provide refuge for bats unless disturbance occurs.

Increased presence of cross border violators(CBVs) along the border

between the United States and Mexico, who use these same structures

forshelter, contributesto disturbance to colonial bats(USFWS 2008).

L. yerbabuenae are also at risk when they frequent human structures

for roosting purposes or to feed at hummingbird feeders because they

are at risk of predation by domestic cats, being harmed by humans,

or killed or injured through collisions with urban structures.

	 Visual surveys during our study indicated no large fields of agave

plants(≥1.8 flowering agaves/ha) in the vicinity ofArivaca,Arizona,

but L. yerbabuenae had been documented using area hummingbird

feeders in late summer. Therefore, we were interested in (1) which

nectar bat species were using feeders; (2) how they used a landscape

withfewnaturalfoodresources, and(3)where theywereday-roosting.

Because bats are nocturnal it is often difficult to monitor nightly feeding behavior. However, radio telemetry helps elucidate the foraging

behavior of L. yerbabuenae (Ober and Steidl 2004; Ober and others

2005), and allows researchers to track bats across both temporal and

spatial scales and identify flight corridors between day and night

roosts (Amelon and others 2009).

Study Area

	 Thisstudywas conductednearthe townofArivaca,PimaCounty,in

southeasternArizona, USA(31o34’N, 111o19’W), an area surrounded

by the Las Guijas Mountains, Cobre Ridge, Tumacacori Mountains,

andCerroColorado Mountains.Arivaca is approximately 29 kmwest

of the Santa Cruz River, 45 km west of the Santa Rita Mountains, and

about 19 km north of the international border with Mexico.Although

there islittle naturalforaging habitatfor nectarivorous batsin the area,

there are significant stands of flowering agaves during the summer

months in the Santa Rita Mountains to the east. That range has at

least four documented lesser long-nosed bat post-maternity roosts.

Despite a scarcity of natural food plants nearArivaca, local residents

have reported that their hummingbird feeders are often visited by

bats in late summer. Arivaca is a small community (~ 900 residents

according to the 2000 census), located at approximately 1100 m in

elevation with homes concentrated on small lots in the historic town

site and homes east of Arivaca on 0.5 ha lots to larger (≥4 ha) lots.

ThehabitatsurroundingArivaca issemidesertgrassland(Brown1994)

characterized by low rolling hills and ephemeral streams. Arivaca

Creek, which contains dense cottonwood galleries, generally flows

from east to west and is located less than 0.40 km south of Arivaca.

The lowundulatingterrainofthe studyareamade constantmonitoring

of telemetered bats sometimes difficult to achieve so radio-tracking

stations were located on high points whenever possible, with views

overlooking Arivaca and nearby canyons.

Methods

	 We captured up to 15 L. yerbabuenae per year (n 2010 = 13 and n 2012

= 15) during post-maternity migration (five each in August, September, and October). We radio tracked the bats during three sessions,

each lasting 2 weeks per month for a total of 6 weeks per year. Bats

were captured at residential hummingbird feeders both in Arivaca

and approximately 6.4 km east of Arivaca. Protocols outlined by the

American Society of Mammalogists (Sikes and others 2011) were

followed while handling all bats, and appropriate scientific collecting

permits were maintained during this study. Because of the potential

threatofWhiteNoseSyndromebeingspreadbetweenbatsbyhumans,

USFWS decontamination protocols outlined for bat researchers were

followed during the capture and handling process (USFWS 2011).

Radio transmitters (#LB-2N from Holohil Systems Ltd., Ontario)

weighing 0.42g were attached to L. yerbabuenae using colostomy

adhesive. Prior to attachment of a transmitter, it was confirmed that

the mass of the transmitter was less than 5% of the bat’s total body

mass (Aldridge and Brigham 1988).

	 During each session of radio tracking, three to five radio-tracking

stations were maintained to monitor telemetered bats. Each station

had a scanning radio receiver (R1000, Communications Specialists,

Inc., Orange, California) and a three or five element Yagi antenna, a

compass, and a long-range radio tomaintain communication between

stations. Each telemetry station was manned prior to sunset unless

personnel were mobile and actively following a signal. To ensure

that the radio receivers were functioning properly, test beacons were

placed along undisclosed bearings and personnel took sightings on

these prior to the evening’s activities.

	 Each night, personnel constantly monitored all radio frequencies

of bats telemetered during each month. If a signal was detected, the

frequency and general bearing was communicated to allstations, and

these stations attempted to monitor the frequency until the signal was

lost.For eachsignaldetected,the frequency,timeofdetection,bearing,

and strength of the signal were recorded on data sheets. Personnel

were able to continue scanning all frequencies while monitoring a

particular frequency. If a new frequency was detected, the monitor

would obtain a bearing for the newly detected frequency and coordinate with other stations. In order to better define foraging areas,

both stationary radio tracking stations and mobile stations were used

to monitor L. yerbabuenae behaviors and habitat use. The program

LOAS 4.0.3.8 (Ecological Software Solutions, LLC) was used to

determine intersections from telemetry station bearings. Because of

the low rolling terrain of the area, three simultaneous points were not

always achieved on bat detections so ‘Best Biangulation’ was used

to determine bat locations. We conducted statistical analysis on the

radio tracking data using the software package JMP4.0 (SAS Institute

Inc. 1996).

Results

	 During the 2-year study, we tracked 28 L. yerbabuenae, including

17 adult females, 7 subadult females, 1 adult male, and 3 subadult

males. The first session of radio tracking each year was conducted in

mid-August,whennectarbats aredocumentedinsoutheasternArizona

grasslands; the lastsession was conducted in early October (USFWS

2007; Sidner 2010). During each studymonth in 2010 and 2011, three

to five L. yerbabuenae were each affixed with a radio transmitter that

was approximately 3% of their body mass and released. Upon release

of a bat, radio telemetry stations immediately began monitoring its

movements. During 2010-2011, we captured 134 bats of seven different species while mist netting for L. yerbabuenae on the patios of

homes and at two flowering agaves (table 1). All species captured

were expected to occur in semidesert grasslands of southeastern

Arizona (Hoffmeister 1986). The diversity of bats was higher than

expected at hummingbird feeders because two residences used for
USDA Forest Service Proceedings RMRS-P-67. 2013 429

Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) Buecher and Sidner

netting had large porchesthat offered night roosting opportunitiesfor

many bat species. Despite the lack of food plants for nectarivorous

bats within the study area, we captured a surprising number of both

L. yerbabuenae (n = 33) and C. mexicana (n = 86) over the course

of the study. In 2010, despite radio tracking for 14 nights per month,

we were unable to continuously track bats over multiple consecutive

nights. During August, we tracked five L. yerbabuenae for only one

night each; in September we tracked five bats for one to three nights

each; and in October we tracked three batsfor one to two nights each.

Because the 2010 radio tracking had such limited results, no statistical

analyses were conducted. However, signals from telemetered bats

were detected in the vicinity of the Santa Rita Mountains to the east,

so we suspected the bats might be using a day-roost in that mountain

range.

	 The radio tracking sessions in 2011 were more successful and we

had sufficient numbers of bearing intersections for statistical analysis. In August 2011, we captured five L. yerbabuenae at a residence

approximately 6.4 km east of Arivaca. Two of the bats had modest

to abundant plant pollen on their heads and face, indicating that they

were successful infinding pollen-bearing flowers,most likely agaves,

before visiting the hummingbird feeders. We tracked these animals

for 3 to 10 nights (χ = 6, SE = 1.18) before the signals were lost.

August bearings on telemetered bats were used to calculate location

intercepts using LOAS (2011) (fig. 1). One bat disappeared from

the area after one night so the behaviors of the remaining four were

analyzed statistically. These bats behaved as a group and the latitudes

and longitudes of their bearing intersections were similar (ANOVA

latitude F3,79 = 1.38, P = 0.26, ANOVA longitude F3,79 = 0.71, P = 0.55 ).

As a group, these animals concentrated their foraging efforts east of

Arivaca. In August 2011, we were able to track telemetered bats to

an abandoned mine in the Santa Rita Mountians where we found five

of our radio transmitters on the floor (four from 2011 and one from

2010). A subsequent exit count documented approximately 7,000 L. 

yerbabuenae emerging from the mine at sunset. After the bats were

gone, a trip into the site found five additional transmitters (one from

2011 and four from 2010).

	 In September, we captured five L. yerbabuenae at a residence in

Arivaca, where the homes are located closer together. We tracked

these animals for two to nine nights (χ = 4.2, SE = 1.72) until the

signals were lost. September bearings on telemetered bats were used

to calculate location intercepts using LOAS (2011) (fig. 2). The

September bats flew more directly to Arivaca, the area where they

were captured, and foraged around homes closer to town. Statistical

analysis of the five bats shows that they behaved as a distinct group

within latitude (ANOVAlatitude F4,146 = 0.88, P = 0.48), but notstrongly

within longitude (ANOVAlongitude F4,146 = 5.88, P <0.001). Statistical

Table 1—Mist netting results from the 2010-2011 radio telemetry survey 

near Arivaca, Arizona.

Scientific name Common name Number

Caught

 Antrozous pallidusa Pallid bat 1

 Eptesicus fuscusa Big brown bat 4

 Choeronycteris mexicanab Mexican long-tongued bat 86

 Leptonycteris yerbabuenaeb Lesser long-nosed bat 33

 Myotis auriculusa Southwestern myotis 2

 Myotis velifera Cave myotis 7

 Tadarida brasiliensisc Brazilian free-tailed bat 2

a

Vespertilionidae

b

Phyllostomidae

c

Molossidae

Figure 1—Gold circles indicate intersection locations from LOAS for five telemetered bats during August, 2011. These points 

indicate where L. yerbabuenae were flying in relation the capture site east of Arivaca
430 USDA Forest Service Proceedings RMRS-P-67. 2013

Buecher and Sidner Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) 

analysisofbehaviordifferencesbetweenbatstelemeteredinAugust(6

kmeast ofArivaca)to batstelemetered during September(inArivaca)

showed a significant difference between the latitude where these bats

foraged (t228 = -12.49, P < 0.001). Some of the bats captured 6.4 km

east of Arivaca did fly as far as Arivaca; however, they consistently

usedmore areas east oftown than those bats captured inArivaca itself.

Bats originally captured at feeders in Arivaca flew more directly to

that area before settling down to forage most of the night. Differences

in longitude are not as pronounced, but still significant (t 228 = -2.73,

P = 0.01), with bats captured east of Arivaca flying more to the north

than bats that were captured in Arivaca.

In October, we captured eight L. yerbabuenae: five at the same

Arivaca residence as in September and three at the same residence

as in August east of Arivaca. The plan for this last month was to radio track five L. yerbabuenae captured at two different sites. It was

hoped that this could elucidate if bats acted as distinct groups and

used different foraging patterns, depending on their capture site. We

tracked the telemetered bats for four to eight nights (χ = 5.8, SE =

0.63) until signals were lost. October bearings on telemetered bats

were used to calculate location intercepts using LOAS (2011) (fig.

3). The October bats did not forage with a distinct configuration, but

appeared to combine both the behaviors ofAugust bats andSeptember

bats. When these five bats are analyzed statistically they behaved

significantly differently across latitude (ANOVA latitude F4,76 = 0.23, P

< 0.001), distributed from east to west between the two capture sites.

However, longitudinally they behaved as a group (ANOVAlongitude F4,76

= 1.24, P = 0.30), staying within a limited north to south range.

	 Once the day roost was found, we were able to estimate average

flight speed for telemetered bats by using a telemetry station along

I-19 to document exittimesfromthe roost,while one ortwo stationsin

theArivacaValley detected the bats once they crossed theTumacacori

Mountains. Eleven bats were documented traversing this distance for

an estimated average speed of 28.3 kph. Sahley and others (1993)

found similar speeds (27.2 kph) by L. yerbabuenae commuting from

a day-roost in Sonora, Mexico, to forage on flowering columnar cacti.

L. yerbabuenae visiting hummingbird feeders in the Tucson basin

(table 2) travelled to foraging areas a bit faster (32.8 kph) although

these animalswere commutingfrommultipleday-roosts(Lowery and

others 2009). All nectar bat flight speeds are significantly faster than

documented for many regional insectivorous bat species (Hayward

and Davis 1964). Sahley and others (1993) suggested that the large

body size and high wing loading of L. yerbabuenaemay be an adaptation for long-distance commutes to food resources across naturally

heterogeneous landscapes.

Discussion

	 We were unable to consistently track telemetered bats in 2010

because of issues related to losing the radio signals almost immediately after bats were telemetered. Unfortunately, when a transmitter

is not detected once affixed, there are three possible reasons: (1) the

bat continued its migration to another foraging area in southeastern

Arizona; (2) the bat migrated towardsits wintersite in Mexico; or (3)

the transmitter was groomed off at a day- or night-roost. In 2010, we

assumed that these bats were continuing their migration from west

to east across southeastern Arizona to Madrean Sky Islands where

there were good stands of flowering agaves. The fact that nectar bats

were captured in semidesert grassland that lacked stands of agaves

was attributed to their learned knowledge that hummingbird feeders

offered sugar-water to ‘power’ them across inhospitable landscapes

(Cockrum and Petryszyn 1991; Lowery and others 2009). However,

in 2011 we were more successful in continually monitoring telemetered bats and we were able to document bats commuting between

their day-roost in the Santa Rita Mountains to hummingbird feeders

Figure 2—Blue circles indicate intersection locations from LOAS for five telemetered bats during September, 2011. These points 

indicate where L. yerbabuenae were flying in relation to the capture site in Arivaca.
USDA Forest Service Proceedings RMRS-P-67. 2013 431

Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) Buecher and Sidner

Table 2—Flight distances and speeds recorded for lesser long-nosed bats reported by various researchers.

Source Year Location Distance Avg. Speed Food Resources

 Buecher/Sidner 2011 Pima Co., AZ 35.4 km 28.3 kph sugar water

 Horner et al. 1998 Sonora, Mexico 32.5 km 15.5 kmh columnar cacti

 Lowery et al. 2009 Pima Co., AZ 28.8 km 32.8 kph sugar water

 Ober et al. 2004 Cochise Co., AZ 19.2 km unknown paniculate agaves

 Sahley et al. 1998 Sonora, Mexico 27.2 km 27.2 kph columnar cacti

 Sidner 1995 Cochise Co., AZ 27.4 km unknown paniculate agaves

Figure 3—Red circles indicate intersection locations from LOAS for five telemetered bats during October, 2011. These points 

indicate where L. yerbabuenae were flying in relation to the two capture sites, one in Arivaca and one east of Arivaca.

in Arivaca. The discovery of both 2010 and 2011 radio transmitters

groomed off in the roost documents bats from both years using the

mine for day-roosting.All bats captured inArivaca during 2011 were

found day-roosting a majority of the time in the Santa Rita roost, approximately 32-38 km east of the capture sites. What is interesting

about this behavior is that these L. yerbabuenae appear to nightly

bypass naturally occurring agaves in the Santa Rita Mountains and

closer hummingbird feeders along the more urbanized I-19 corridor.

Hummingbird feedersinTubac,GreenValley, and evenPatagonia and

Sonoita were closer to the day-roost than the Arivaca feeders (table

3). This seemingly unusual behavior may be an effort by individuals

in a large colony to partition available food resources. The Santa Rita

Mountains have at least four known lesser long-nosed day-roosts

with an approximate regional population of 16,000 bats. Because the

newlydiscoveredSantaRita roosthas approximately7,000bats,these

animals might be required to disperse across a broad landscape in

order to find sufficient food resources. The telemetered animals were

all captured in or near Arivaca, which may bias the results towards

those individuals that have already chosen this area to feed, whereas

other L. yerbabuenae in the day-roost may forage at closer feeders.

Despite a lack of flowering agaves in the Arivaca area, nectar bats

are actually present in good numbers, presumably because they have

‘learned’to utilize sugar water resources at urban homes to maintain

their energetic needs during migration across southern Arizona.

Overall, these animals were highly loyal to this artificial food source

(sugar water) and some bats returned many nights to the area after

capture.

	 Whenthe telemeteredbats are evaluatedas agroup,thebats captured

6.4 km east ofArivaca behaved as a unit and did notforage differently

when measured spatially. Bats captured in Arivaca also behaved as

a unit and travelled farther towards Arivaca to feed. Howell (1979)

hypothesized that L. yerbabuenae foraged in groups as a benefit for

locating food resources in a heterogeneous landscape. Our results

support this hypothesis by showing that bats captured at specific

sites behaved strongly as a group on subsequent nights. When the 2

months (August and September) are compared there is a significant

difference in the foraging areasthe bats used. However, during October, when the sampling for bats was conducted from two residences

6.4 km apart, the combined group foraged across all latitudes and

longitudes reflected by the August and September bats. Since these
432 USDA Forest Service Proceedings RMRS-P-67. 2013

Buecher and Sidner Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) 

bats behave so cohesively on the landscape, it would be interesting

to understand how they initially segregate themselves into foraging

groups. Three possibilities might be (1) kin groups; (2) members of

summer maternity colony cohorts; or (3) winter colony cohorts.

	 The results of our radio tracking study provide important information on activity patterns and foraging behaviors of L. yerbabuenae

on artificial food resources near Arivaca, Arizona. While this area

was not previously documented for having regular visitation at urban

hummingbird feeders by nectar bats, our high capture ratesshow that

feeders are used by both L. yerbabuenae and C. mexicana. However,

one question, still to be determined, is how much do these animals

benefit physiologically from this highly artificial food? The flowers

of columnar cacti and paniculate agave offer nectar, but also large

amounts of pollen, which have amino acids that the bats convert to

proteins (Howell 1974). Horner and others (1998) estimated that the

energy budget of L. yerbabuenae was ~40 kJ/day and it took 80-100

visits to columnar cacti flowers to obtain this energy. It is currently

unknownhowmany visitswouldbeneeded toartificialfoodresources

to obtain an equal amount of energy. Another concern is how hummingbird feedersimpact the naturalforaging behaviors andmigration

patterns of nectar bats.Withoutthe availability of urban hummingbird

feeders, there is very little reason for nectar bats to be foraging in the

grasslandssurroundingArivaca.There has beenwork to quantify how

many agave plants are needed to maintain L. yerbabuenae populations (Ober and others 2005). Although the grasslands surrounding

Arivaca appear to have few agaves, fortunately there are naturally

occurring stands of agaves in many of the Sky Islands to the east of

Arivaca. This allows nectar bat populations to persist in the region

and offers resources required during energetically costly migration

for L. yerbabuenae migrating to winter roosts in Mexico.

References

Aldridge, H.D.J.N. and Brigham, R.M. 1988. Load carrying and maneuverability in an insectivorous bat: a test of the 5% “rule” of radio-tracking.

Journal of Mammalogy 69:379-382.

Amelon, S.K.; Dalton, D.C.; Millspaugh, J.J.; and Wolf. S.A. 2009. Radiotelemetry. Pp. 57-77 in Ecological and behavioral methods for the study of

bats (T.H.Kunz and S Parsons eds). The Johns Hopkins University Press,

Baltimore, MD.

Buecher, D.C. and Sidner,R. 2011.Telemetry study forlesserlong-nosed bats,

SBI-net TucsonWest Project, Block One, Office of Technology Innovation

and Acquisition, U.S. Customs and Border Protection, U.S. Border Patrol,

Table 3—Distances from the Santa Rita day-roost to area hummingbird 

feeders. Bold italic entries indicate feeders definitely used by the 

telemetered bats during 2010-2011 radio tracking. Note that five 

urban areas where feeders are closer to the day-roost than Arivaca, 

Arizona.

Feeder locations Santa Rita roost

miles (km)

 Tubac 7 (11.2)

 Patagonia 13 (20.8)

 Continental 14 (22.4)

 Sonoita 16 (25.6)

 Green Valley 18 (28.8)

 East Arivaca 20 (32)

 Sahuarita 22 (35.2)

 Arivaca 24 (38.2)

West Arivaca 27 (44.8)

Tucson Sector.Arizona. Unpublished paper on file at U.S. Fish andWildlife

Service, Tucson, AZ. 44 p.

Brown,D.E.1994.Warm-temperategrasslands:Semidesertgrassland.Pp.123-

131 in Biotic communities: Southwestern United States and Northwestern

Mexico (D.E. Brown ed.). University of Utah Press, Salt Lake City, UT.

Ceballos G.; FlemingT.;ChávezC.; and Nassar,J. 1997. Population dynamics

ofLeptonycteris curasoae (Chiroptera:Phyllostomidae)inJalisco,Mexico.

Journal of Mammalogy 78(4):1220-1230.

Cole, F.R and Wilson, D.E. 2006. Leptonycteris yerbabuenae. Mammalian

Species No. 797, The American Society of Mammalogists: 1–7.

Cockrum, E.Land Petryszyn, Y. 1991. The long-nosed bat, Leptonycteris:An

endangered species in the southwest? Occasional Paper Number 142. The

Museum Texas Tech University. 32 p.

Fleming, T.H.; Nunez, R.A.; and Sternberg, L. 1993. Seasonal changes in the

diets of migrant and non-migrant nectarivorous bats as revealed by carbon

stable isotope analysis. Oecologia 94:72-75.

Hayward, B. and Davis, R. 1964. Flight speeds in western bats. Journal of

Mammalogy 45:236-242.

Hoffmeister, D. 1986. Mammals of Arizona. University of Arizona Press.

Tucson, AZ. 602 p.

Horner, M.A.; Fleming, T.H.; and Sahley, C.T. 1998. Foraging behavior and

energetics of a nectar-feeding bat, Leptonycteris curasoae (Chiroptera:

Phyllostomidae). Journal of Zoology 244:575–586.

Howell, D.J. 1974. Bats and pollen: physiological aspects of the syndrome of

chiropterophily. Comparative Biochemistry and Physiology. 48:263-276.

Howell, D.J. 1979. Flock foraging in nectar-feeding bats: advantages to the

bats and to the host plants. American Naturalist 114:23-49.

Krebbs, K. 2007. Lesser long-nosed bat (Leptonycteris cursoae) movements

and foraging in central-eastern Arizona. Unpublished paper on file at:

Arizona Game and Fish Departnemt, Phoenix, AZ. 22 p.

LOAS. 2011. Ecological Software Solutions LLC. Hegymagas, Hungary.

Version 4.0.3.8.

Lowery, S.F.,Blackman, S.T., andAbbate, D. 2009. Urbanmovement patterns

of lesser long-nosed bats (Leptonycteris curasoae): Management implications for the Habitat Conservation Plan within the City of Tucson and the

Town of Marana. Unpublished paper on file at: Arizona Game and Fish

Department, Phoenix, AZ. 22 p.

Ober, H.K. and Steidl, R.J. 2004. Foraging rates of Leptonycteris curasoae 

vary with characteristics of Agave palmeri. The Southwestern Naturalist

49(1):68-74.

Ober, H.K., Steidl, R.J.; and Dalton, V.M. 2005. Resource and spatial-use

patterns of an endangered vertebrate pollinator, the lesser long-nosed bat.

Journal of Wildlife Management 69(4):1615-1622.

Ramirez, J. 2011. Population genetic structure of the lesser long-nosed bat

(Leptonycteris yerbabuenae)inArizona andMexico.Masterthesis.Tucson:

University of Arizona. 90 p.

Sahley, C.T., Horner, M.A., and Fleming, T.H. 1993. Flight speeds and mechanical power outputs of the nectar-feeding bat, Leptonycteris curasoae

(Phyllostomidae: Glossophaginae.Journal of Mammalogy 74(3):594-600.

SAS Institute Inc. 1996. JMP start statistics: A guide to statistics and data

analysis using JMP and JMP IN software. Wadsworth Publishing Company, Belmont, CA.

Sidner, R. 1996. Sixth annual monitoring of potential roostsites of the lesser

long-nosed bat (Leptonycteris curasoae) and other species of bats on the

Fort Huachuca Military Reservation, Cochise County, Arizona, May–December 1995. Unpublished paper on file at: USAG Commander through

Engineering and Environmental Consultants, Inc., 122 p.

Sidner,R.2010.Twentiethannualmonitoringofbats especiallythe endangered

lesser long-nosed bat (Leptonycteris yerbabuenae) on the Fort Huachuca

Military Installation, Cochise County, Arizona, April – December 2009.

Unpublished paper on file at: Commander, U.S. Army Garrison, Fort

Huachuca, Arizona. 90 p.

Sikes, R.S., Gannon, W.L., and Animal Care and Use Committee of ASM.

2011. Guidelines of the American Society of Mammalogists for the use of

wild mammals in research. Journal of Mammalogy 92:235-253.

Shull, A. 1988. Endangered and threatened wildlife and plants: Determination of endangered status for two long-nosed bats. Federal Register

53(190):38456-3860.
USDA Forest Service Proceedings RMRS-P-67. 2013 433

Long Distance Commutes by Lesser Long-Nosed Bats (Leptonycteris yerbabuenae) Buecher and Sidner

U.S. Fish and Wildlife Service. 1997. Lesser long-nosed bat recovery plan.

Unpublished paper on file at: U.S. Fish and Wildlife Service,Albuquerque,

NM. 49 p.

U.S. Fish and Wildlife Service (S. Richardson). 2007. Final 5-year review

summary and evaluation for the lesser long-nosed bat. Unpublished paper

on file at: Arizona Ecological Services Office, Phoenix. 43 p.

U.S. Fish and Wildlife Service 2008. Biological Opinion on Secure Border

Initiative (SBInet) Tucson West Tower Project,Ajo, Tucson, Casa Grande,

Nogales andSonoitaStationsAreaofOperation,U.S. BorderPatrol,Tucson

Sector. #AESO/SE, 22410-2008-F-0373. 153 p.

U.S. Fish and Wildlife Service. 2012. White-Nose Syndrome Website. URL

address: www.fws.gov/whitenosesyndrome/research.html. Last updated:

March, 2012.

Wilkinson, G.S. and Fleming, T.H. 1996. Migration and evolution of lesser

long-nosedbatsLeptonycteris curasoae,inferredfrommitochondrialDNA.

Molecular Ecology 5:329-339.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
434 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Effects of Roads on Wildlife in Arizona: How Far Have We 

Traveled?

Hsiang Ling Chen and John L. Koprowski 

Wildlife Conservation and Management, School of Natural Resources and the Environment, The University of 

Arizona, Tucson, Arizona 

Abstract—Roads are conspicuous and pervasive features of landscapes and represent one of the most significant anthropogenic impacts on natural areas and wildlife. The Madrean Archipelago is defined by natural 

levels of fragmentation due to geography; however, human population growth and transportation needs 

threaten to exacerbate levels of isolation in the region. Scientists, as well as transportation and resource 

management agencies, have increased their concern about road impacts on wildlife. To identify needs of 

future research and managements, we reviewed 29 road-ecology-related, peer-reviewed publications and 

governmental research in Arizona and compiled geography, focal species, and topic. A taxonomic bias 

toward large mammals (72%) is evident. Study areas are concentrated along highways and state routes 

(76%). Despite a prevalence of studies on wildlife road crossing, most research focuses on distribution and 

movements, whereas impacts at the population and community level are rarely described. 

Introduction

	 Roads are conspicuous and pervasive features of landscapes

and represent one of the most significant anthropogenic impacts

on natural areas and wildlife (Forman and Alexander 1998). Over

20% of the land in the United States is affected by roads and traffic (Forman 2000). Road construction causes destruction of habitat

and habitat loss directly and facilitates deforestation and landscape

fragmentation (Coffin 2007). Roads and traffic can cause mortality,

impede and altermovements of animals, influence population density

(Fuentes-Montemayor and others 2009;Roedenbeck andVoser 2008;

Rytwinski andFahrig2007;TrombulakandFrissell2000), andchange

community structure (Bissonette and Rosa 2009; Goosem 2000).

	 TheMadreanArchipelagoisdefinedbynaturallevelsoffragmentation due to geography; however, increase of human population and

transportation needs threaten to exacerbate levels of isolation in the

region (ADOT 2006). The human population has increased 24.6% in

Arizona,from 5.1 million in 2000 to 6.4 million in 2010 (U.S.Census

Bureau 2012).The road systeminArizona has expanded dramatically

fromtworough roadsinthe1800stoaround 92,800 kmofroads nowadays (ADOT 2012). With the increase in wildlife-vehicle collisions

in Arizona, transportation and resource management agencies have

elevated their concern about road impacts on wildlife and recognize

the need to develop effective mitigation (Ruediger and others 2005).

In this paper, we search road-ecology-related, peer-reviewed publications and governmental research reports and determine geography as

well asspecies and study focusto assess diversity and identify gapsin

publications and research projectsrelated to road impacts on wildlife

in Arizona.

Materials and Methods

Literature Search

	 We used the Web of Science literature search tool that includes

publications from 1945 to April 22, 2012 to search publications related to roads and wildlife inArizona.We selected the “Topic” search

option and used search terms “road and Arizona” and “highway and

Arizona.” We browsed titles and abstracts in the search results and

included publications related to wildlife in our analysis. For governmental research reports, we focused on research projects conducted

byArizonaDepartment ofTransportation (ADOT) andArizonaGame

and Fish Deportment (AZGFD). To search project reports about road

impactsonwildlife,webrowsedADOTresearchprojects(SPRreports)

from1968 to 2012 in thewebsite (http://www.azdot.gov/TPD/ATRC/

Publications/project_reports/index.asp), AZGFD technical reports

from 1990 to 1999 (http://www.azgfd.gov/w_c/Technical_Reports.

shtml), and AZGFD wildlife and conservation research webpage

(http://www.azgfd.gov/w_c/research.shtml). We also used the following internet resources to search other research reports not under

ADOT and AGFD, including Wildlife and Roads Search Engine

(http://www.wildlifeandroads.org/search/), Transportation Research

Board(http://www.trb.org/Main/Home.aspx),TRIDdatabase (http://

trid.trb.org/), and Google Scholar (http://scholar.google.com/). We

realize that some peer-reviewed articles and governmental reports

might be missed because of lack of congruence between keywords

that we used and publications and the availability of governmental

works to public.
USDA Forest Service Proceedings RMRS-P-67. 2013 435

Effects of Roads on Wildlife in Arizona: How Far Have We Traveled? Chen and Koprowski

Variables

	 We recorded year of publication, focal species, research location,

and the main topic for each peer-reviewed literature and governmental research project. To analyze the taxonomy of the focal species,

we recorded the number of publications for each vertebrate animal

class. For publications with mammals as focal species, we recorded

the number of publications for each Order. We categorized the focal

mammalian species as small mammals if body mass was less than 5

kg (Merritt 2010).We used themidpoint of adult bodymasses;female

bodymasswasusedinsexuallydimorphic species(Hoffmeister1986).

Results and Discussions 

Do We Have Progress?

	 We found a total of 30 studies related to road impacts on wildlife, with 10 peer-reviewed articles, and 20 governmental research

projects conducted by ADOT, AZGFD and U.S. Geological Survey.

We excluded projects that are in-progress or unpublished because

information on research is not consistently accessible. With rapid

development of the subdiscipline of road ecology since 2000, the

number of publications has increased considerably (1900%) in the

past 16 years, from 1 publication by Rosen (1994) in 1994-2000 to

19 publications in 2006-2011 (fig. 1). The increased interest from

governmental agencies in the integration of scientific research with

decision making on transportation planning had positive impacts

on the accumulation of knowledge of road-wildlife interactions and

likely enhanced the publication of peer-review literature. 

What Species Are Underrepresented? Does 

the Size Have Influence?

	 Among 30 studies, the most common taxon of study is the mammals (77%; fig. 2) with few studies on reptiles (10%), birds (3%),

or general survey on multiple taxonomic groups (10%). No case

study examines amphibians; however, road kill is a major source of

amphibian mortality and may contribute to global decline of amphibians (Glista and others 2008). Our effort to understand road effects

on mammals in Arizona does not extend equally to all Orders. Order

Artiodactyla (ungulates) is the most frequently studied group and

elk (Cervus elaphus) and desert bighorn sheep (Ovis canadensis)

are the two most common studied species. When we look at the

number of studies against the proportion of total species for each

order of mammals in Arizona, a taxonomic bias toward ungulates is

evident (fig. 3). Ungulates represent 4% of total mammalian species

in Arizona, but were the subject of 96% of the studies. Compared

to large and medium mammals, small mammals, which constitute

85% of the state's mammalian species, received a disproportionately

small amount of attention in these studies (fig. 4). Of course, vehicle

collisions or evasive driving maneuvers focused on small mammals

Figure 1—Number of publications related to road impacts on wildlife 

in Arizona by year from 1994 to 2011.

Figure 2—Number of studies related to road impacts on wildlife in 

Arizona by vertebrate class from 1994 to 2011. 

Figure 3—Relative proportions of Arizona mammalian species by order 

compared with representation of those orders in studies related to road 

impacts on wildlife from 1994 to 2011. 
436 USDA Forest Service Proceedings RMRS-P-67. 2013

Chen and Koprowski Effects of Roads on Wildlife in Arizona: How Far Have We Traveled?

Figure 4—Relative proportions of Arizona mammallian species by size 

compared with representation of those size classes in studies related to 

road impacts on wildlife from 1994 to 2011.

do not generally cause property damage or injury to humans, factors

that, in part, drive this disproportionate distribution of publications.

Where Are We On the Road? 

	 Current road and wildlife related research in Arizona mainly focuses on the barrier effect of roads on animal movements and efforts

to improve motorists’ safety. Road type is biased in research location with most efforts focused on highways and state routes (77%).

Research topics are aimed at improved road permeability, reduced

wildlife-vehicle collisions, and evaluating effectiveness of wildlife

passages. Arizona has taken a leadership role in mitigation measures

to minimize barrier effects of roads and to restore connectivity by

designing and installing wildlife underpasses, overpasses, wildlifeproof fencing, and alert systems along highways and state routes

(Reuer 2007). Besides wildlife passages, governmental agencies

and scientists also continue to investigate the efficacy of currently

installed structures(for example, culverts) asroad crossing structures

(Mikele and Michael 2007).With this amount of effort, the frequency

of wildlife-vehicle collisions has declined and highway permeability

for elk has been improved (Dodd and others 2007). 

Where Is the Gap?

	 Our analyses suggest that a gap in knowledge of road impacts on

wildlife existsinArizona.Mostimportantly,we knowvery little about

the impacts of roads on wildlife in Arizona and in unique biomes

such as the Sonoran Desert. If we acknowledge a general dearth of

literature on road impacts, we can examine if there are important

areas where additional studies are required and prioritize our needs.

We have already addressed the paucity of studies on groups beyond

large mammals. Large mammals are important focalspecies because

these animals are highly vulnerable to roads in part because they are

more likely to encounterroads due to extendedmovement and ranges,

and populations are more susceptible to road mortality because of

low reproductive rates and low natural density (Fahrig andRytwinski

2009). However, negative effects of roads occur across a wide range

of vertebrates (Laurance and others 2009; Wisdom and others 2000),

and abundant evidence suggests that response to roads and traffic

likely vary considerably acrossspecies(Goosem 2001; Laurance and

others 2004;Taylor and Goldingay 2010). For example,roadsrestrict

movements of forest-dependent species of birds but not frugivorous

and edge and gap species(Laurance and others 2004). Whereaslarge

mammalstend to avoid roads, response ofsmall mammalsto roadsis

more complicated (Fahrig andRytwinski 2009).Despite the extensive

size of forest road networks (Coghlan and Sown 1998), forest roads

are relatively ignored. Several studies have demonstrated that even

narrow roads less than 10 m wide can have barrier effects (Forman

and Alexander 1998; Swihart and Slade 1984; Wilson and others

2007).Environmental changes associatedwithedges createdbyforest

roads may impact species composition within the forest ecosystem,

especially forspecies,such astree squirrels, that are sensitive to forest

fragmentation (Koprowski 2005; Murcia 1995).

	 Besides a taxonomic and geographic bias toward large mammals

and highways in current studies, we need to develop research questions at different levels and scales. Despite several callsfor needs and

increased attention to research at population and community levels

(Fahrig and Rytwinski 2009; van der Ree and others 2011; Underhill

andAnhold (2000), these kinds of research are scarce inArizona. We

have gained important knowledge of barrier effects on individual

animal movements, but we know less about effects of roads on populations. Does the magnitude of population fragmentation caused by

barrier effects of roads affect population persistence? How do roads

affect social structure and reproductive success within populations?

Most road ecology studies focus on single species, and few assess

community level impacts or address species interactions near roads.

The danger of this is that we might miss important pieces of a complex system. For example, abundance of rodents often increased at

areas near roads, potentially due to the negative effects of roads on

predator populations, which cannot be known if we only investigate

a single taxon or closely related species (Bissonette and Rosa 2009;

Rytwinski and Fahrig 2007). We tend to have focused on patterns but

do not fully understand the causes and mechanisms. For example, we

know that roads have barrier effects on severalspeciessuch as desert

bighorn sheep, pronghorn (Antilocapra americana), and species of

snakes, but do not know why (Dodd and others 2010;Jones and others

2011; McKinney and Smith 2007). Do animals avoid roads because

of a gap in cover, or environmental changes along road edges, or

traffic disturbance? Studies that address the relative importance of

different mechanisms of the effects of roads on wildlife are needed

(Roedenbeck and others 2007).

Our Roads Ahead

TheMadreanArchipelagois a regionthat exhibitshighlevelsofdiversity and is fortunately less disturbed compared to many other places

in theUnitedStates.Although the transportation systemhas expanded

in recent decades, road density remains relatively low (World Bank

2008), so that ample opportunities exist to minimize road impacts in

thisregion.One substantial challenge inmanagement ofroad network

systems is that no attempt has been made to synthesize piecemeal

information fromindividualstudiesinto a substantial, comprehensive

picture about how road networks function in broader scale within

the Southwest (Gucinski and others 2001). The need for increased

cooperationbetweengovernmentaldepartments and agenciesis clear.

Comprehensive planning that would minimize road effects requires

collaborationamongacademia,public interestgroups, andlocal,state,

and federal agencies. We encourage enhanced multi-disciplinary,

inter-agency supported events such as the International Conference
USDA Forest Service Proceedings RMRS-P-67. 2013 437

Effects of Roads on Wildlife in Arizona: How Far Have We Traveled? Chen and Koprowski

on Ecology and Transportation (ICOET) and Infra Eco Network

Europe Meetings (IENE) as well as projects that address large scale

landscape management. We believe that large scale efforts such as

The Wildlands Projects (McDonnell 2002) and Arizona’s Wildlife

Linkages Assessment (ADOT 2006) as well as increased interest by

university scientists and the history of a firm commitment to collaborative research among agencies provide the scaffolding for such a

region-wide approach in Arizona. With continued expansion of the

human population predicted for Arizona on the order of 7.4 million

people by 2020 (ADOT 2006), as well as long term projections of

significant redistribution and fragmentation due to climate change

(Opdam andWascher 2004;Weiss and Overpeck 2005), construction

of a comprehensive and collaborative long-term plan is necessary.

Acknowledgments

	 We thank people who have built and contributed to the internet

resources on which this paper is based. Michelle Crabb and Scott

Sprague kindly shared information about their research; Chia Chun

Tsai and Hsin Ju Wu reviewed and provided helpful comments on

the manuscript.

References

ADOT. 2006.Arizona’s wildlife linkages assessment.Arizona Department of

Transportation, Phoenix, AZ. Online: http://www.azdot.gov/inside_adot/

OES/AZ_Wildlife_Linkages/assessment.asp [April 26, 2012].

ADOT. 2012. Arizona Department of Transportation, Phoenix, AZ. [Online].

Available:http://www.azdot.gov/Highways/AboutITD.asp[April26,2012].

Bissonette, J. A.; and Rosa S.A. 2009. Road zone effects in small-mammal

communities. Ecology and Society. 14 (1). Online: http://www.ecologyandsociety.org/vol14/iss1/art27/ [April 26, 2012].

Bouchard,J.A.;FordT.;EigenbrodF.E.;FahrigL.2009.Behavioralresponses

of northern leopard frogs (Rana pipiens) to roads and traffic: implications

for populationpersistence.EcologyandSociety.14(2).Online:http://www.

ecologyandsociety.org/vol14/iss2art23 [April 26, 2012].

Coffin,A. W. 2007. From roadkill to road ecology:Areview of the ecological

effects of roads. Journal of Transport Geography. 15(5):396-406.

Coghlan. G.; and Sown R. 1998. National forest road system and use (Draft

Report, 1-30-98, Engineering Staff). Washington, DC: U.S. Department

of Agriculture, Forest Service.

Dodd, N. L.; Gagnon J. W.; Boe S.; Manzo A.; Schweinsburg R. E. 2007.

Evaluationofmeasurestominimizewildlife-vehicle collisions andmaintain

wildlifepermeabilityacrosshighways:StateRoute260,Arizona,USA.Final

project report 540. Phoenix, AZ: Arizona Department of Transportation,

Arizona Transportation Research Center.

Dodd, N. L.; Gagnon J. W.; Sprague S.; Boe S.; Schweinsburg R. E. 2010.

Assessment of pronghorn movements and strategies to promote highway

permeability: U.S. Highway 89. Final projectreport 619. Phoenix,AZ:ArizonaDepartmentofTransportation,ArizonaTransportationResearchCenter.

Fahrig, L.; and Rytwinski, T. 2009. Effects of roads on animal abundance: an

empirical review and synthesis. Ecology and Society. 14(1). Online: http://

www.ecologyandsociety.org/vol14/iss1/art21/ [April 26, 2012].

Forman, R. T. T. 2000. Estimate of the area affected ecologically by the road

system in the United States. Conservation Biology. 14:31-35.

Forman, R. T. T.; andAlexander, L. E. 1998. Roads and their major ecological

effects. Annual Review of Ecology and Systematics. 29: 207-231.

Fuentes-Montemayor, E.; Cuaron, A. D.; Vazquez-Dominguez, E.; BenitezMalvido,J.;Valenzuela-Galvan,D.;Andresen,E. 2009.Living on the edge:

roads and edge effects on small mammal populations. Journal of Animal

Ecology. 78: 857-865.

Glista, D. J.; DeVault, T. L.; DeWoody, J. A. 2008. Vertebrate road mortality predominatly impacts amphibians. Herpetological Conservation and

Biology. 3(1): 77-87.

Goosem, M. 2000. Effects of tropical rainforest roads on small mammals:

edge changes in community composition. Wildlife Research. 27: 151-163.

Goosem, M. 2001. Effects of tropical rainforest roads on small mammals:

inhibition of crossing movements. Wildlife Research. 28(4): 351-364.

Gucinski,H.; Furniss, M.J.; Ziemer R. R.; M.H. Brookes. 2001. Forest roads:

a synthesis of scientific information. Gen. Tech. Rep. PNW-GTR-509.

Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific

Northwest Research Station. 103 p.

Hoffmeister, D. F. 1986. Mammals ofArizona. Tucson:University ofArizona

Press. 602 p.

Jones, T. R.; Babb, R. D.; Hensley, F. R.; LiWanPo, C.; Sullivan, B. K. 2011.

Sonran desert snake communities at two sites: Concordance and effects of

increasedroadtraffic.HerpetologicalConservationandBiology.6(1):61-71.

Koprowski, J. L. 2005. The response of tree squirrels to fragmentation: A

review and synthesis. Animal Conservation. 8: 369-376.

Laurance, S. G. W.; Stouffer, P. C.; Laurance, W. E. 2004. Effects of road

clearings on movement patterns of understory rainforest birds in central

Amazonia. Conservation Biology. 18(4): 1099-1109.

Laurance, W. F.; Goosem, M.; Laurance, S. G. W. 2009. Impacts of roads

and linear clearings on tropical forests. Trends in Ecology and Evolution.

24(12): 659-669.

McDonnell, T. 2002. Technical review of the wildlands projects and how

it is affecting the management of state, federal and private lands in the

United States. Sequim, Washington: North Western Research Institute.

Online:http://nwri.org/2011/07/technical-review-of-the-wildlands-project2002/#more-93. [April 29, 2012].

McKinney,T.; andSmith,T.2007.US93bighorn sheep study:Distributionand

trans-highwaymovementsofdesertbighornsheepinnorthwesternArizona.

Final Report 576. Phoenix, AZ: Arizona Department of Transportation,

Arizona Transportation Research Center.

Merritt, J.F. 2010. The biology of small mammals. Baltimore: Johns Hopkins

University Press. 336 p.

Mikele, L. P.; and Michael, F. I. 2007. Use of simulated highway underpass

crossingstructuresbyflat-tailedhornedlizards(Phrynosoma Mcallii).Final

project report 594. Phoenix, AZ: Arizona Department of Transportation,

Arizona Transportation Research Center.

Marcia, C. 1995. Edge effects in fragmented forests: Implications for conservation. Trends in Ecology and Evolution. 10(2): 58-62.

Opam, P.; Wascher, D. 2004. Climate change meets habitat fragmentation:

Linking landscape and biogeographical scale levels in research and conservation. Biological Conservation. 117(3): 285-297.

Reuer, C. 2007. Synthesis of animal-vehicle collision mitigation measures.

Final project report 612. Phoenix, AZ: Arizona Department of Transportation, Arizona Transportation Research Center.

Roedenbeck, I.A.; Fahrig, L.; Findlay, C. S.; Houlahan,J. E.;Jaeger,J.A. G.;

Klar, N.; and others. 2007. The Rauischholzhausen agenda for road ecology. Ecology and Society. 12(1). Online: http://www.ecologyandsociety.

org/vol12/iss1/art11/ [April 29, 2012].

Roedenbeck, I.A.; and Voser, P. 2008. Effects of roads on spatial distribution,

abundance and mortality of brown hare (Lepus europaeus) in Switzerland.

European Journal of Wildlife Research. 54: 425-437.

Rosen, P. C.; and Lowe, C. H. 1994. Highway mortality of snakes in the Sonoran desert ofsouthernArizona.BiologicalConservation. 68(2): 143-148.

RuedigerW.;Wall,K.;Wall,R.2005.EffectsofHighwaysonElkHabitatinthe

Western United States and Proposed MitigationApproaches: International

Conference on Ecology andTransportation; 2005August 29- September 2;

San Diego, CA. Proceedings. Raleigh, NC: Center for Transportation and

the Environment, North Carolina State University: 269-278.

Rytwinski, T.; and Fahrig, L. 2007. Effect of road density on abundance of

white-footed mice. Landscape Ecology. 22(10): 1501-1512.

Swihart, R. K.; and Slade, N. A. 1984. Road crossing in Sigmodon hispidus

and Microtus ochrogaster. Journal of Mammalogy. 65(2): 357-360.

Taylor, B. D.; and Goldingay, R. L. 2010. Roads and wildlife: impacts,

mitigation and implicationsfor wildlife management inAustralia. Wildlife

Research. 37(4): 320-331.

Trombulak,S.C.; andFrissell,C.A.2000.Reviewof ecological effectsofroads

onterrestrial andaquatic communities.ConservationBiology.14(1):18-30.
438 USDA Forest Service Proceedings RMRS-P-67. 2013

Chen and Koprowski Effects of Roads on Wildlife in Arizona: How Far Have We Traveled?

Underhill, J.E.; and Angold, P.E. 2000. Effects of roads on wildlife in an

intensely modified landscape. Environmental Review. 8: 21-39.

U.S. Census Bureau. 2012. State and County Quick Facts. U.S.A. Online:

http://quickfacts.census.gov/qfd/states/04000.html.[ April 28, 2012]

van der Ree, R.; Jaeger, J. A. G.; van der Grift, E. A.; Clevenger, A. P. 2011.

Effects of roads and traffic on wildlife populations and landscape function:

road ecology is moving toward largerscales. Ecology and Society. Online:

http://www.ecologyandsociety.org/vol16/iss1/art48/.[ April 28, 2012]

Weiss,J.L.;Overpeck,J.T. 2005.Isthe Sonoran Desert losing its cool?Global

Change Biology.11(12): 2065-2077.

Wilson, R. F.; Marsh, H.; Winter, J. 2007. Importance of canopy connectivity

for home range and movements of the rainforest arboreal ringtail possum

(Hemibelideus lemuroides). Wildlife Research. 34(3): 177-184.

Wisdom, M.J.; Holthausen, R.S.; Wales, B. K. 2000. Source habitats for terrestrial vertebrates of focus in the interior Columbia basin: Broad-scale

trends and management implications. Gen. Tech. Repo. PNW GTR-485.

Portland, OR: USDA Forest Service, Pacific Northwest Research Station.

World Bank. 2008. World Bank Indicators – United States – Transportation.

Online: http://www.worldbank.org/ [April 28, 2012].

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 439

Tools for Research
440 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 441

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Wildlife Survey and Monitoring in the Sky Island Region 

with an Emphasis on Neotropical Felids

Sergio Avila-Villegas and Jessica Lamberton-Moreno

Sky Island Alliance, Tucson, Arizona 

Abstract—The Sky Island region of southwestern United States and northwestern Mexico consists of isolated 

mountain ranges separated by deserts and grasslands. It mixes elements from five major ecosystems: the Rocky 

Mountains, Sierra Madre Occidental, the Sonoran and Chihuahuan deserts and the Neotropics. Here some 

Neotropical species reach their northern ranges, such as jaguars (Panthera onca) and ocelots (Leopardus 

pardalis). Sky Island Alliance seeks to build cooperative relationships with landowners in Sonora and public 

land managers in Arizona to facilitate scientific research, encourage large predator conservation, and establish corridors connecting patches of continuous habitat throughout the region. We have conducted wildlife 

surveys with remote cameras in northern Sonora since 2007 and in Arizona since 2009. Our photographic 

records cover 24 species of wild mammals, including jaguar (Panthera onca), ocelot(Leopardus pardalis), 

puma (Puma concolor) and bobcat (Lynx rufus). In 2009 we documented Arizona’s first live ocelot, and 

recently we documented two individual jaguars 30 miles south of the border in Sonora. The project’s results 

have increased our knowledge of borderland wildlife, especially the Sonoran ocelot, and supported the 

certification of a 10,000-acre private ranch as a wildlife preserve in northern Sonora.

Introduction

	 In2005,SkyIslandAlliance initiateda studyoncross-borderpermeability in northern Sonora, Mexico, to establish corridors connecting

habitat throughout the region, and create a network of conservation

ranches in northern Sonora (Avila 2009).

	 Neotropical felids have been recorded in southern Arizona and

northern Sonora (Brown and Lopez-Gonzalez 2001; Grigione and

others 2007; McCain and Childs 2008; Rosas-Rosas and others

2010). Brown and Lopez-Gonzalez (2001) provide a comprehensive

account of these Neotropical felids throughout the Sky Island region

up until the end of the 1900s. Both Neotropical species are protected

in the United States and Mexico. Currently the U.S. Fish andWildlife

Service (USFWS) leads recovery efforts for both species. Because

of their low density and solitary nature, little is known about their

population size, range, or habitat requirements in the United States

(Grigione and others 2007).

	 At least six male jaguars have been recorded north of the border

since 1996. There were two different sightings in 1996: one in the

Baboquivari Mountains (Childs 1998), and another in the Peloncillo

Mountains (Glenn 1996). McCain and Childs (2008) monitored at

least two jaguars on the Atascosa, Tumacacori, and Baboquivari

Mountains. No home range studies have been conducted for jaguars

in southwestern United States; however McCain and Childs (2008)

used camera-trapsto provide the only estimate of one northern jaguar,

a minimum observed range of 525 mi2.

	 There are two subspecies of ocelots found in the United States:

the Sonoran subspecies (L. p. sonoriensis), described by Goldman

(1943) with four specimens from southernmost Sonora, and the

better-studied Texas subspecies (L. p. albescens). Populations of the

Sonoran ocelot are isolated from theTexas ocelot by the Sierra Madre

highlands (Grigione and others 2007). The population of Texas ocelots is found almost 300 miles south in latitude of the northernmost

Sonoran ocelots.

	 Lopez-Gonzalez and others (2003) conducted surveys throughout

Sonora, collecting36recordsofocelotsfromranchers, cowboys, cattle

associations, and outfitters. They plotted record localities on habitat

maps finding that 75% were associated with subtropical thornscrub,

tropical deciduous forest, and tropical thornscrub; 92% of records

were within an elevation range of 820 to 3773 ft. The most northerly

record of a breeding population was 30°30’ in latitude. The authors

found only one record of a kitten, in southern Sonora, and a few

ocelots were recorded in oak woodlands, all males (Lopez-Gonzalez

and others 2003).

Objectives

•	 To build cooperative relationships with private landowners in

Sonora andArizona to facilitate research and establish corridors

connecting patches of continuous habitat throughout the region.

•	 To survey and inventory land mammalspeciesin the Sky Island

region.

•	 To increase the knowledge about jaguar and ocelot in the Sky

Island region.
442 USDA Forest Service Proceedings RMRS-P-67. 2013

Avila-Villegas and Lamberton-Moreno Wildlife Survey and Monitoring in the Sky Island Region . . .

Methodology

Study Area

	 The Sky Island region of southwestern United States and northwestern Mexico consists of isolated mountain ranges separated by

deserts and grasslands.Itmixes elementsfromfivemajor ecosystems:

the Rocky Mountains, Sierra Madre Occidental, the Sonoran and

Chihuahuan deserts, and the Neotropics. Some Neotropical species,

such as the jaguar, ocelot, and coati (Nasua narica), reach their

northernmost ranges here (fig. 1).

	 We initiated landowner outreach and surveysin northern Sonora in

2005 to establish partnershipsin the region.We used habitatmodelsto

ground-truth thepredictedcorridorsforjaguars andocelots(Boydston

andLopez-Gonzalez2005;Hattenand others2003;Menke andHayes

2003; Menke 2004). We placed a total of 48 remote camerasin select

locations in Nogales, Imuris, and Agua Prieta Municipios in Sonora,

and Santa Cruz and Cochise Counties in Arizona (fig. 2). Figure 1—Biotic influences in the Sky Islands.

Figure 2—Sky Island Ranges. Camera locations include Sierra La Esmeralda, Cibuta and 

Azul mountain ranges, the Cocospera River and San Bernardino Valley in Sonora, and 

the Atascosa, Patagonia, Whetstone, Dragoon and Mule Mountains in Arizona (map by 

Alex Smith, 6/20/2012).
USDA Forest Service Proceedings RMRS-P-67. 2013 443

Wildlife Survey and Monitoring in the Sky Island Region . . . Avila-Villegas and Lamberton-Moreno

	 We adopted a standardized remote camera protocol described in

Chavez and Ceballos(2006). Remote cameras are non-invasive tools

useful for the study of rare, protected or elusive animals (Karanth

and Nichols 1998; Medellin and others 2006). We used DeerCam,

Cuddeback and Covert DLC cameras that were set to display time

and date on the photographs (table1).

	 Given the low density of jaguars and ocelots in their northern

extent (Brown and Lopez-Gonzalez 2001), camera sites were chosen

to maximize the probabilities for photographing felids, instead of a

randomized sampling design. We collected ocelot photographs from

February 2007 to April 2011, and plotted the relative frequency of

events over annual and daily cycles to estimate activity patterns.

Results

	 Our photographic records include 24 species of wild mammals

in five orders: Marsupialia, Rodentia, Lagomorpha, Carnivora and

Artiodactyla.

Sonoran Ocelot

	 We collected69recordsofliveocelots andfourofkilledocelots.Our

remote camera photographs (60 photographs, 54 events) were taken

at elevation ranges between 4185 and 5330 ft. in Madrean evergreen

woodland. A majority of photographs was taken in Sonora’s Sierra

Azul 30 milessouth of the international border. Only one photograph

was taken in Arizona, in the Whetstone Mountains, almost 40 miles

north of the border. These records confirm the northernmost breeding population of Sonoran subspecies at 30°50’ of latitude—the first

reliable ocelot record in Arizona in 45 years—and an ocelot walking

on fresh snow (fig. 3). Our “live” ocelot documentations include at

least two males, one female, one kitten, and two of undetermined

gender (table 2).

	 Analyses of the relative frequencies of 54 photographic events on

24-hr cycles confirm the ocelots’ nocturnal activity patterns, with

peaks of activity between 4:00 and 5:00 and between 20:00 and

0:00. The nocturnal activity patterns of ocelots were similar to those

in Texas where activity peaked between 20:00 and 05:00 (Grigione

and Mrykalo 2004;Yates 2008). Ocelots were mostly active from late

winter to early foresummer (December to May; Yates 2008). Ocelot

daily and annual activity patterns are shown in figs. 4 and 5.

Northern Jaguar

	 We found records of four jaguarsin the United Statesin the 2000s,

three confirmed with photographs. A jaguar named “Macho B” was

Table 1—Camera Locations by County or Municipio.

Municipio or County, Number Landownership

State of cameras (Private/public)

Nogales, Sonora 10 Private

Agua Prieta, Sonora 10 Private

Imuris, Sonora 15 Private

Santa Cruz, AZ 5 Private/Public

Cochise Co., AZ 8 Public

Figure 3—Sonoran ocelot. Sonoran ocelot walking on fresh snow in the 

Sierra Azul, February 2008.

Table 2—Ocelot records in the Sky Islands in the 2000s.

Number of records Description Municipality, State (Year)

1 Live animal in cage (photo) Imuris/Magdalena, Sonora (2007)

2 Poached animals (carcass) Imuris, Sonora (2007)

2 Road kill Highway 15, Sonora (photo) Imuris, Sonora (2008)

Road kill Highway 60, Arizona (photo) Superior, Arizona (2010)

3 SIA tracks sets Imuris, Sonora (2007, 08, 09)

65 photos SIA’s 53 photographic events Imuris, Sonora (2007-2011)

SIA’s 1 photographic event Cochise County, Arizona (2009)

AZGFD reported sightings Cochise County, Arizona (2011, 12)
444 USDA Forest Service Proceedings RMRS-P-67. 2013

Avila-Villegas and Lamberton-Moreno Wildlife Survey and Monitoring in the Sky Island Region . . .

Figure 5—Ocelot annual activity patterns. Ocelot relative frequencies of 52 photographic events over 12-month 

period.

Figure 4—Ocelot daily activity patterns. Ocelot relative frequencies of 54 photographic events over 24-hr cycles.
USDA Forest Service Proceedings RMRS-P-67. 2013 445

Wildlife Survey and Monitoring in the Sky Island Region . . . Avila-Villegas and Lamberton-Moreno

monitored in Arizona from 1996 (McCain and Childs 2008) until his

death in 2009.Anotherjaguarwas photographed in theAnimasValley

in 2005 (Warner Glenn, personal communication). In June 2011, a

U.S.Border Patrol pilotreported a jaguarin the SantaRita Mountains

approximately 40 miles north of the border. In November 2011, the

Arizona Game and Fish Department (AZGFD) confirmed a jaguar

sighting in the Whetstone Mountains, approximately 40 miles north

of the border.

	 During this study we photographed two different jaguars (fig . 6)

in 2010 and 2011 in the Sierra Azul, and identified two track sets

in Sierra Cibuta (2005) and Sierra Azul (2009). Our photographic

records were collected in Madrean evergreen woodland at elevation

between 4430 and 4590 ft.All photographs were taken between 18:00

and 6:00 hrs. The small sample size of photographs might not truly

indicate activity patterns of northern jaguars. Jaguar records in the

Sky Islands in the 2000s are shown in table 3.

Discussion

	 Landowner involvement is critical for long term conservation in

private lands of northern Sonora.Haines and others(2006) emphasize

the importance of protecting private lands for ocelot conservation.

Landowner outreach is an investment that pays off with landowner

buy-in, access,localknowledge, andsharingof anecdotalinformation.

As a result of our outreach and research collaboration in Sonora, a

10,000-acre private ranch called El Aribabi received a Certificate of

Voluntary LandConservation by Mexican agencyCONANPin 2011,

thus constituting the most recent wildlife reserve in northern Mexico.

	 Ocelothabitat characteristicssuchas elevation,vegetationtype, and

vegetation cover found in this study contrast with those described in

Lopez-Gonzalez and others(2003).The authorsfound ocelots associated with tropical or subtropical habitats, mostly in the southern half

of Sonora. The localities where we documented ocelots in northern

Sonora are typical Sky Island habitat.We confirmed the northernmost

breeding population of Sonoran ocelotsin the SierraAzul Mountains

in Madrean evergreen woodlands. The elevation ranges and latitude

at 30°50’ are above those recorded by Lopez-Gonzalez and others

(2003). Additionally we found the first reliable ocelot record in Arizona in 45 years in the Whetstone Mountains at 5330 ft. of elevation

(Grigione and others 2007).

	 The location of a breeding population 30 miles south of the international border, suggests that ocelots could travel northward and

currentlyreside insouthernArizona.The estimateddispersaldistances

for Texas ocelots (3 to 20 miles; Caso 1994; Crawshaw 1995) are

less than those between the breeding population in northern Sonora

and sighting locations in Arizona (approximately 50 miles).

	 The AZGFD reported a road-killed ocelot near Superior, Arizona,

and confirmed it to be of wild origin (De Young and Holbrook 2010).

This animal might represent the northernmost record for the species,

Figure 6—Northern jaguar. Northern jaguar photographed in Sierra 

Azul, January 2010.

Table 3—Jaguar records in the Sky Islands in the 2000s.

Number of Municipality, State 

records Description (Year)

11 photos Warner Glenn (photo) Animas Valley, NM (2005)

SIA’s 9 photographic events Imuris, Sonora (2010-2011)

 (2 individuals)

AZGFD (photo) Cochise Co., Arizona (2011)

Undetermined ~100 photos) McCain and Childs (photos) Santa Cruz and Pima County, Arizona

 (1996-2009)

2 SIA track set Nogales, Sonora (2005)

 SIA track set Imuris, Sonora (2009)

1 AZGFD/Border Patrol report Pima County, Arizona (2011)
446 USDA Forest Service Proceedings RMRS-P-67. 2013

Avila-Villegas and Lamberton-Moreno Wildlife Survey and Monitoring in the Sky Island Region . . .

north of the Sky Island region. We estimate the distance between Superior to the border, to the Huachuca or to the Whetstone Mountains

to be over 120 miles. It is highly probable that ocelots in northern

Sonora and southern Arizona—only 70 miles apart—are connected

by migration corridors and potentially represent an established transboundary population. However, in the case ofthe documentation near

Superior, travel from northern Sonora seems unlikely.

	 Our data on jaguars is scarce; however, it includes photographs of

two different males and a track set potentially expanding the known

range of Macho B. We believe our data are inconclusive in terms of

daily or seasonal activity patterns.

	 In 5 years of monitoring public and private lands along the border, we did not find any anecdotal or reliable records of jaguarondi

(Herpailurus yaguarondi).

Conclusion

	 The lack of scientific information on Sonoran ocelots, other than

historic localities and habitat associations, is evident in the USFWS

Draft Recovery Plan. Resultsfrom thisstudy increase the knowledge

of Sonoran ocelots. The localities where we documented ocelots in

Sonora are typical of Madrean Sky Islands and differ from those

documented before.

	 Grigione and others(2007)suggest that conservation of peripheral

populations,such as ocelot and the jaguar’s northernmostrange, plays

a role in maintaining the genetic viability of a species. It is essential

that source populations of jaguars and ocelots be located and travel

routes be identified.Bothmontane corridors and riparian areas appear

to be important travel routes for Neotropical felids.

	 The importance of trans-boundary conservation cannot be overemphasized for felids and other species. It is critically important to

consider jaguar and ocelot’srecent presence in mountain ranges near

the border and the connecting linkages in order to protect enough

habitatforthe establishmentoffuturepopulations(Grigione andothers

2007).Themost distant portions ofspecies’distributions are often the

last refuge for their survival (Lopez Gonzalez and others 2003). The

northern extent of the ocelot’s range in southeastern Arizona could

prove to contain enough suitable habitat, connective linkages, and

prey species to support a breeding population.

	 The Sonoran ocelot has been given the least amount of attention

in scientific research and conservation planning, plainly contrasting

with information on the Texas subspecies. This lack of information

couldleadtoinappropriatedecisionsforrecovery, asdifferenttypesof

habitat, elevation, and vegetation cover are found in its northernmost

distribution ranges. We believe there are clear differences between

the two ocelot subspecies and their needs, with the northernmost

population of Texas ocelotsfound in latitudes almost 300 milessouth

of the Sonoran ocelots. Due to unique topographic, bio-geographic

and climatic features that occur in northern Sonora and southernArizona, the Sky Island region also represents a distributional extreme

with important genetic and adaptive resources that can characterize

peripheral populations (Grigione and others 2007).

	 To support the recovery of endangered tropical felines in the Sky

Islandregion,migrationcorridorsthatlinkkeyhabitat coresinMexico

and the United States must be identified and protected. In southern

Arizona, large tracts of protected public lands already exist, including NationalWildlifeRefuges, National Monuments,Wilderness and

National Conservation Areas. Opportunities for habitat protection

south ofthe border exist, with Sonora being the second largeststate in

Mexico, andhavinglowhumanpopulationdensities(Lopez-Gonzalez

and others 2003).Relatively intact patches of continuous habitatmake

this area a stronghold for the ocelot.

	 Bordersecurity activitiesrepresent the greatest threat to jaguar and

ocelot recovery in the United States. Habitat connectivity across the

border remains highly threatened by construction of an impermeable

barrier anditsrelatedinfrastructure alongtheborder.The construction

of border infrastructure negatively impacts species that live in the

region, endangered or otherwise. Division of otherwise continuous

habitat by a physical barrier blocksthe natural movements ofspecies

whose distribution in the United States is already limited, confining

them to isolation and potential extinction (Córdova and De la Parra

2007).

References

Avila, S. 2009. The Cuatro Gatos Project: wildlife research and conservation

in northwestern Mexico and implications of the border fence. Wild Felid

Monitor. Winter 2009. Vol. 2(1).

Boydston, E. and C. Lopez-Gonzalez. 2005. Sexual differentiation in the

distributionpotentialofthenorthernjaguars(Panthera onca).Proc.RMRSP-36. U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station.

Brown, D. and C. Lopez-Gonzalez. 2001. Borderland Jaguars: tigres de la

frontera. The University of Utah Press. 170 p.

Chavez, C., and G. Ceballos. 2006. Memorias del Primer Simposio. El jaguar

Mexicano en elSigloXXI:SituaciónActual yManejo.CONABIO-Alianza

WWF Telcel-Universidad Nacional Autónoma de México. México, D.F.

Childs, J. 1998. Tracking the felids of the borderlands. Printing Corner Press.

El Paso, TX. 77 p.

Córdova, A. and C. De la Parra. Eds. 2007. A Barrier to our Shared Environment: The Border Fence between the United States and Mexico. First Edition. Secretariat of Environment and Natural Resources, National Institute

of Ecology, El Colegio de la Frontera Norte, Southwest Consortium for

Environmental Research & Policy. Mexico.

De Young, R. and J. Holbrook. (2010). Analysis and interpretation of ocelot

material lineages from road-killed ocelots in Texas and Arizona. A report

to the US Fish and Wildlife service and Texas Parks and Wildlife Department. Texas A&M University, Kingsville.

Glenn, W. 1996. Eyes of fire: encounter with a borderlands jaguar. El Paso,

TX: Printing Corner Press. 28 p.

Grigione, M. and R. Mrykalo. 2004. Effects of artificial night lighting on

endangered ocelots (Leopardus paradalis) and nocturnal prey along the

UnitedStates-Mexicoborder:Aliterature reviewandhypothesesofpotential

impacts. Urban Ecosystems, 7: 65–77.

Grigione, M., Scoville, A., Scoville, G., and K. Crooks. 2007. Neotropical

cats in southeast Arizona and surrounding areas: past and present status of

jaguars, ocelots and jaguarundis. Mastozoología Neotropical, 14(2):189-

199, Mendoza

Haines, A., Janecka, J., Tewes, M., Grassman, L. and P. Morton. (2006). The

importance of private lands for ocelot Leopardus pardalis conservation in

the United States. Oryx Vol. 40(1).

Hatten, J. R., Averill-Murray, A., and W. E., Van Pelt. 2003. Characterizing

and mapping potential jaguar habitat in Arizona. 203, 1-28. Phoenix, AZ,

Nongame and Endangered Wildlife Program, Arizona Game and Fish

Department.

Karanth K. U. and J. D. Nichols. 1998. Estimation of tiger densities in India

using photographic captures and recaptures. Ecology, 79:2852-2862

Lopez-Gonzalez, C., Brown, D., and J.P. Gallo Reynoso. 2003. The ocelot

Leopardus pardalis in north-western Mexico: ecology, distribution and

conservation status. Oryx. Vol. 37. No.3.
USDA Forest Service Proceedings RMRS-P-67. 2013 447

Wildlife Survey and Monitoring in the Sky Island Region . . . Avila-Villegas and Lamberton-Moreno

McCain, E., and J. Childs. 2008. Evidence of resident jaguar (Panthera onca)

in the southwestern United States and the implications for conservation. J.

of Mammalogy. 89: 1-10.

Medellín, R., Azuara, D., Maffei, L., Zarza, H., Bárcenas, H., Cruz, E., Legaria, R., Ramos-Fernández, G., and S.Avila. (2006). Capítulo III. Censos 

y Monitoreo. In: Chavez, C., and G. Ceballos. 2006. Memorias del Primer 

Simposio. El jaguar Mexicano en el Siglo XXI: Situación Actual y Manejo”.

CONABIO-Alianza WWF Telcel-Universidad Nacional Autónoma de

México. México, D.F.

Menke, K. 2004. Priority Conservation Areas in the U.S.-Mexico Border

Region for North American Tropical Cats: the Jaguar, Jaguarundi, and

Ocelot. University of New Mexico. In: Vacariu, K., and J. Neeley. 2005.

Ecological Considerations for Border Security Operations: Outcomes and

Recommendations of the Border Ecological Symposium. Tucson, Az.

Menke, K. A. and Hayes, C. L. 2003. Evaluation of relative suitability of

potential jaguar habitat in New Mexico. 1-30. New Mexico Game and

Fish Department.

Rosas-Rosas, O., Bender, L.C., and R. Valdes. 2010. Habitat correlates of

jaguar kill-sites of cattle in northeastern Sonora, Mexico. Human-Wildlife

interactions 4(1).

Silver, S. C., Ostro, L. E. T., Marsh, L. K, Maffei, L., Noss, A. J., Kelly, M.

J., Wallace, R. B., Gomez, H., and G. Ayala. 2004. The use of camera traps

for estimating jaguar Panthera onca abundance and density using capture/

recapture analysis. Oryx 30(2): 148-154.

Silver, S. C. 2004. Assessing jaguar abundance using remotely triggered

cameras. 1-25. Wildlife Conservation Society, Global Carnivore Program.

Yates, J. 2008. Using remote cameras to assess predator-prey interactions of

wild felids in the Mexican sky islands. University of Georgia.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
448 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Analysis of the Seasonal Activity Rate of Sympatric 

Carnivores and their Prey in Saguaro National Park

Mary Beth Benton

Saguaro National Park, Tucson, Arizona

Siria A. Cerda-Navarro, Katie R. Keck, and Brittany N. McKnight

	 University of Arizona, Tucson, Arizona

Abstract—We investigated the monthly activity rates of bobcats (Lynx rufus), gray foxes (Urocyon cinereoargenteus) and four of their potential prey species, antelope jackrabbits (Lepus alleni), black-tailed jackrabbits (Lepus 

californicus), desert cottontails (Sylvilagus audubonii) and eastern cottontails (Sylvilagus floridanus), in Saguaro 

National Park using remote camera trap data. We analyzed images taken over an 11-month period and over 

10,000 camera nights and compared the time of day, month, and location of photographs of bobcats, gray 

foxes and four lagomorph species. Our findings suggest that bobcats and gray foxes are successful sympatric 

species for much of the year, but during the autumn and winter seasons gray fox activity increased as a result of 

reduced food availability. Lagomorph activity also changed in the winter, when both jackrabbit species and desert 

cottontail rabbit activity decreased, while eastern cottontail activity increased, in a manner that was quite similar 

to that of gray foxes. Lagomorph activity may be influenced by the activity of their predators. Knowledge of how 

mammals utilize the landscape of Saguaro National Park will enable wildlife managers to implement more effective 

conservation and management strategies. 

Introduction

Sympatry among animals has become more prevalent due to habitat 

loss, urbanization, and expansion of natural distributions (Chamberlain 

and Leopold 2005). Sympatry often results in competition among 

animals due to the use of shared resources. Niche segregation of 

resources can occur in areas where sympatry is pronounced. Fedriani

andothers(1999)foundthatthree sympatric carnivores—Iberian lynx

(Lynx pardinus), Eurasian badger (Meles meles), and red fox (Vulpes 

vulpes)—altered their activity patterns to avoid overlap in resource

utilization. The potential for competition among species that overlap 

in geographic range is increased when species are comparable in 

body size and engage in similar feeding habits (Barrientos and Virgós 

2006). In the Sky Island region of southern Arizona, two sympatric 

carnivore species—bobcat (Lynx rufus) and gray fox (Urocyon cinereoargenteus)—consume similar prey species such as lagomorphs 

and ground dwelling rodents (Fritzell and Haroldson 1982; Larivière 

and Walton 1997). Bobcats and gray foxes inhabit relatively similar 

environments. In Arizona, these two species can be found in mixed 

forests, rocky outcrops, and brush areas from elevations up to 3,000 

m or more (Tekiela 2008). 

One such area in the desert southwest where bobcats and gray foxes 

co-occur is Saguaro National Park in Tucson, Arizona. Both species 

are prevalent within the park, where Benton and others (unpublished 

data) found that their temporal and spatial distributions overlap 

extensively. Riley (2006) argues that bobcats are more affected by

proximity to urbanization than gray foxes.Gray foxes have often been

documented entering developed areas and their home rangestypically

occupy urban locations whereas bobcatsrarely enter developed areas

and have been shown to avoid crossing paved roads (Riley 2006).

These patterns are likely caused by differences in diet and the social

system of the species. Although gray foxes and bobcats tend to prey

on small rodents, bobcats are strictly carnivorous whereas gray foxes

are omnivorous (Riley 2006). Riley (2006) states that this more flexible use of the landscape allows gray foxes easier access to increased

resources and habitat whereas bobcats’ sensitivity has put them at

risk to decreased viability in urban-protected zone interfaces. The

trend described by Riley (2006) does not apply in Tucson, Arizona,

where bobcats are found not only in the wilderness areas of Saguaro

National Park but also in adjacent neighborhoods and deeper into the

suburban sprawl of the community (Haynes and others 2010). The 

large population of mountain lions in Saguaro National Park may be 

influencing the movement of bobcats into urban areas and could be 

one indication as to why so few bobcats are seen on wildlife cameras. 

While coyotes have been shown to kill gray foxes (Neale and Sacks 

2001), the interactions between these canid species may not be as 

severe as the interactions between felid species in Saguaro National 

Park. The interspecific relationships of apex predators and mesopredators impact the behavior of animals at every trophic level; therefore, 

it is important to understand these patterns in order to appropriately 

manage and conserve mammalian wildlife. 

Antelope jackrabbits(Lepus alleni),black-tailedjackrabbits(Lepus 

californicus), desert cottontails (Sylvilagus audubonii), and eastern

cottontails(Sylvilagus floridanus)arecommonlyoccurringlagomorphs

found in Saguaro National Park.Although the park has not collected a
USDA Forest Service Proceedings RMRS-P-67. 2013 449

Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey . . . Benton, Cerda-Navarro, Keck, and McKnight

specimen to confirm the identity of the eastern cottontail, thisspecies

ismost likely to be the cottontail that is often observed at higher elevationsin the park (Swann 2011).The temporal and spatial distributions

of these species demonstrate their sympatry and overlaps in activity

with bobcats and gray foxes (Benton and others, unpublished data).

The lagomorphs of Saguaro National Park are herbivorous species

whose diet consists of green plants and some cactus fruits in spring

and summer and other woody plants and berries in winter (Tekiela

2008). These lagomorphs inhabit similar areas ranging from deserts

and grasslands to woodland thickets (Tekiela 2008).

The Sonoran desert experiences five seasons every year. Foresummer 

in May and June, when this study began, brings high temperatures, low 

humidity, and typically no rain. Most plants and animals are dormant 

during this time, due to the lack of available surface water (Phillips 

and Comus 2000). The summer monsoon season—early-July through 

mid-September—is distinguished by high temperature, high humidity, and frequent thunderstorms and corresponds with the second and 

main growing and flowering season for much of the plant community 

(Phillips and Comus 2000). Autumn—October and November—is 

typically warm, with low humidity and very little rain and triggers the 

growing season of the annual plant community (Phillips and Comus 

2000). Winter—December and January—is characterized by mild 

temperatures and sunny days and gentle rains (Phillips and Comus 

2000). Spring—early February through April—when this study was 

completed, is characterized by mild temperatures and little rainfall, 

and accounts for the first flowering season of the year (Phillips and 

Comus 2000). The differences in the availability of flowers, grasses, 

and fruits to gray foxes and lagomorphs during each season in the 

Sonoran desert may influence the rate of activity of these species as 

well as the rate of activity of their conspecifics and predators.

The goal of this study was to determine whether bobcat and gray

fox activity in Saguaro National Park differs during certain periods of

the year based on dietary differences.Additionally,wewere interested

in how bobcat and gray fox activity affects the seasonal activity of

lagomorphs. We tested several hypotheses using images from wildlife cameras that were captured by a Saguaro National Park wildlife

camera program conducted from May 2011 through March 2012. We

predicted that the activity patterns of bobcats and gray foxes would

be similar during the foresummer, summer monsoon, and autumn

seasons due to theirsympatric nature. However, during the winter we

predicted that gray fox activity would shift when supplementary food

resources,such asfruits, were not available.In addition, we predicted

that lagomorph activity would increase during the foresummer,summermonsoon, and autumn when gray foxes have supplementary food

sources, and decrease in late autumn and winter when fox predation

intensifies. 

Methods

Study Area

	 Saguaro National Park is approximately 37,000 ha and is divided

into two districts: Rincon Mountain District and Tucson Mountain

District (fig. 1). Rincon Mountain District (RMD) lies 32 km east of

the center of Tucson and is the larger of the two districts. Elevation

ranges from 664 m to 2,641 m and the dominant plant communities

include thorn-scrub, chaparral, grasslands, and temperate coniferous

forests.TucsonMountainDistrict(TMD)lies 24 kmwest ofthe center

of Tucson and ranges in elevation from 668 m to 1,498 m. Dominant

Figure 1— Map of Saguaro National Park in Tucson, Arizona. The distance between the Rincon Mountain District (east) and the Tucson 

Mountain District (west) is 24 km.
450 USDA Forest Service Proceedings RMRS-P-67. 2013

Benton, Cerda-Navarro, Keck, and McKnight Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey . . .

plant communitiesinclude those that are characteristic ofthe Sonoran

desert ecosystem.

Data Collection

	 Saguaro National Park began a year-long study in May 2011 using

wildlife camerasto determine speciesrichness and species occupancy

within the park.SaguaroNationalParkmade these data available to us

for use in better understanding the monthly activity rates of bobcats,

gray foxes, and lagomorphs.The park usedCuddeback Capture units,

white flash heat, and motion sensor digital wildlife cameras under a

study design where both districts of the park were divided into 1-km2

plots using a grid feature in the Hawthstoolset ofArcMap, a geospatial processing program. Plots were randomly located and distributed

among four elevation gradients: 0-914 m, 914-1,524 m, 1,524-2,134

m, and greaterthan 2,134 m.Any plot containing more than 50% land

outside the park boundary wasremoved from the data set.The steeper

Rincon Mountain District (RMD) was divided into four strata, while

the Tucson Mountain District (TMD) was divided into two strata.

Fifteen plots were chosen in each stratum using a random selection

tool in Hawthstools. Camera locations were randomly selected using

ArcMap as well. On a rotational basis, two plots in each stratum had

four camerasset, no lessthan 200 m apart,for periods of 6 weeks.The

strategy to keep the camera points random, without human bias, was

to place them within 20 m of the randomly selected coordinate. The

study resulted in a total of 10,338 camera nights throughout Saguaro

National Park, including 3,527 nights in strata 1; 3,361 nights in

strata 2; 1,652 nights in strata 3; and 1,798 nights in strata 4 (66.4%

of these nights were in the Rincon Mountain District and 33.6% in

the Tucson Mountain District). For personnel and camera safety, we

occasionally used alternative random points (e.g. if the point fell on

the side of a cliff). Cameras were set using metal stakes and nylon

straps, and were placed as close to the random points as possible in

an open area in hopes of increasing the probability of detection of

an animal if it was present. The cameras were programmed to sense

heat and motion at a 14° angle from the IR sensor, up to a distance

of 10 m. The date and time were programmed on each camera and a

30-second timerwas engaged,meaning thatwhen the cameradetected

movement and a difference in heat gradient, a photograph would be

captured every 30 seconds.

	 Approximately every 2 weeks, resource management staff and

volunteers checked each camera to collect images, solve troubleshooting issues, replace batteries, and assure camera placement was

still intact. Photos were extracted from SD cards and downloaded to

an MP3 player for backup. These data were then brought back to the

office to save images and ensure data security, enter data, and analyze

photos. The date, time, location, species, and number of individuals

associated with each photo were recorded into the park’s wildlife

camera database. The captured images were analyzed using graphs

to observe patterns and trends.

	 This study was solely descriptive as we searched for patterns in

activity between bobcats, gray foxes, and their potential prey animals.

First, we summarized monthly numbers of photographs of bobcats,

gray foxes, and lagomorphs. Total capture numbers for each species

were not equivalent; therefore, we standardized monthly activity

rates using proportions of photographs. Figure 2 illustrates a total

of 257 gray fox images, compared to 67 bobcat images. Figure 3

demonstrates a total of 959 desert cottontail images; compared to 62

eastern cottontail images, 342 antelope jackrabbit images, and 495

black-tailed jackrabbit images.Eachmonth’s number of captures was

divided into the total number of capturesfor thatspeciesto create the

y-axis proportion of photographs. This method allowed for efficient

visual representation of patterns between our focalspecies. Graphing

the monthly activity of bobcats, gray foxes, and lagomorphs allowed

us to explore potential explanations for differences in mesopredator

activity during specific seasons or months.

Figure 2— Monthly activity rate of bobcats and gray foxes in Saguaro National Park as determined by proportion of photographs. 
USDA Forest Service Proceedings RMRS-P-67. 2013 451

Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey . . . Benton, Cerda-Navarro, Keck, and McKnight

Figure 3—Monthly activity rate of lagomorphs in Saguaro National Park as determined by proportion of photographs. 

Results

	 The total number of images of bobcats, gray foxes, and four lagomorph species captured over an 11-month period in Saguaro National

Park were organized in table 1. As demonstrated in figure 2, activity

of gray foxes was relatively low beginning in the foresummer when

the study began and remained low throughout the summer monsoon

season.Activityratesofgrayfoxesincreaseddramaticallyinthemonth

of October, but decreased from November through March. Figure

2 also illustrates that bobcat activity was high from May through

September, but decreased dramatically in October and November,

before peaking again in January and decreasing between January and

March. Figure 3 shows that black-tailed jackrabbit activity patterns

mimic those of bobcats very closely. Black-tailed jackrabbit activity

was high from May through September, before decreasing dramatically in October, then peaking again in December and decreasing

from December through March. Desert cottontail rabbit activity was

also high during the foresummer and part of the summer monsoon

Table 1—Monthly activity rates of bobcats, gray foxes, desert cottontails, eastern cottontails, antelope jackrabbits, and black-tailed 

jackrabbits in Saguaro National Park as determined by proportion of photographs taken between May 2011 and March 2012

Month Bobcat Gray Fox Desert Cottontail Eastern Cottontail Antelope Jackrabbit Black-tailed Jackrabbit

May 0 6 47 0 20 15

June 4 16 115 0 19 33

July 7 20 109 5 16 52

August 9 22 152 9 15 70

September 11 22 65 4 15 79

October 5 56 67 8 4 33

November 3 39 70 22 27 60

December 12 18 85 0 60 73

January 8 29 115 4 45 36

February 6 22 96 10 73 33

March 2 7 38 0 48 11

Total 67 257 959 62 342 495
452 USDA Forest Service Proceedings RMRS-P-67. 2013

Benton, Cerda-Navarro, Keck, and McKnight Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey . . .

season, before decreasing in September. Unlike black-tailed jackrabbit activity, desert cottontail activity did not increase again in the

winter, but rather it remained relatively low throughout the autumn,

winter, and spring seasons. Antelope jackrabbit activity was low

and constant throughout the foresummer and summer monsoon and

their activity was nearly undetectable in October before increasing

between the months of October and March. During the foresummer,

easterncottontail activitywasundetectable.InJulyandAugust eastern

cottontail activity increased, before going down in September and

spiking substantially in November. Immediately following the spike

in activity in November, eastern cottontail activity was undetectable

in December. From January to February eastern cottontail activity

increased before it was undetected again in March.

Discussion

By capturing images of bobcats and gray foxes in the same areas

at approximately the same times, our results indicate that bobcats

and gray foxes are sympatric carnivores in Saguaro National Park

for a large portion of the year; however, as seen in figure 2, gray fox

activity increased dramatically from September to November before

dropping off in December. We predicted this shift in activity was

the result of the differences in the dietary constraints of bobcats and

gray foxes. These results support our hypothesis that gray foxes alter

their activity when utilizing alternate food sources such as grasses

and fruits to supplement their diet of prey animals. Gray fox activity

is lower in warmer months of the year because they have additional

food sources such as manzanita fruit (Neale and Sacks 2001). Scat

analyses suggest that fruit comprises 45-60% of gray fox diet in the

foresummer, summer monsoon, and autumn months of the year and

only about 15% oftheir diet in the winter and spring (Neale and Sacks

2001). Rodents and lagomorphs make up 15-30% of gray fox diet in

the warmer months of the year and 45-50% of their diet in the winter

and spring. Bobcat activity remains relatively constant throughout

the year because their diet is limited to the consumption of meat; 50-

60% of bobcat diet consists of rodents and lagomorphsin the summer

monsoon and autumn, but during the winterthe percentage ofrodents

in the diet decreases and lagomorphs increases from around 5% in

the autumn to 15% in the winter (Neale and Sacks 2001). Prey selection in bobcats may shift as temperatures decrease due to a reduced

amount of desert squirrel and rodent activity during colder months

when they store food and remain in their burrows throughout the

winter (Phillips and Comus 2000).

	 We found some support for our hypothesisthat lagomorph activity

would decrease in late autumn and winter(fig. 3) when gray fox activity increased.Although black-tailed jackrabbits and desert cottontails

exhibited generally similar patterns of behavior across the 11-month

study, desert cottontail activity was high in the summer monsoon,

low during October-December, and then increased slightly in January

before decreasing in the spring. Black-tailed jackrabbit activity was

high in the foresummer and summer monsoon, but dropped in the

autumn and increased during the winter, before it decreased again in

the spring.Antelope jackrabbit activity waslow and constant through

foresummer and summer. Beginning in October, antelope jackrabbit

activity began to increase and between October and December 2011,

the amount of activity doubled. Thisincrease wasshort-lived, before

it dropped to a more stable amount in the spring. We hypothesized

that the change in lagomorph activity during the autumn months may

be the result of an increase in gray fox predation (fig 4). The desert

cottontail may have been more impacted by the change in gray fox

activityduringautumncomparedtoantelope andblack-tailedjackrabbits. The desert cottontail was the only lagomorph species found at

low elevation that did not return to the same high activity rate seen

in the warmer months of the year following its drop in activity in

the autumn. Unlike the antelope and black-tailed jackrabbits, desert

cottontail activity remained relatively constant at a low rate from

October through March. Eastern cottontail activity was undetected

in the spring, a pattern that was unique among the lagomorphs analyzed. A gradual increase in activity was then seen from late summer

through the fall. A marked increase in eastern cottontail activity was

seen only in November 2011. Later in the winter, activity remained

relatively high but was more constant. The difference in detection

among lagomorphs may be the result of decreased food availability

in the forest due to cold temperatures and snow cover. Lagomorph

activity in Saguaro National Park changes seasonally, which may

be due to surface water availability and food supply. The bimodal

periodicity of rain in the Sonoran desert often influences mammalian

behavior because wateris a limiting resource in this arid environment

(Phillips and Comus 2000). It is important to consider this variable

whenanalyzingdetectionratesofgrayfoxes,bobcats, andlagomorphs

in Saguaro National Park.

	 In order to ensure that our data analysis was not overly simplistic,

we investigated alternative hypotheses for the increase in activity

seen in gray foxes between September and December. One alternative explanation is simply sample size: in October 2011 Saguaro

National Park hosted the 2011 National Geographic BioBlitz event.

This event drew thousands of visitors to both districts of the park,

including the back country areas, and many of the wildlife cameras

were deployed for educational purposes; resulting in a smaller number of camera nights. This decrease in the number of camera nights

overall may have created spurious proportional relationships in the

data. The abundances of bobcats and gray foxes differ between the

two districts of the park due to differences in habitat type and overall

available area. Because this study analyzed the activity of species in

both districts ofthe park, wemust accountforthese differences.TMD

has half the elevation gradient as RMD, and as a result, TMD has

fewer strata. Each month TMD had fewer camera nights than RMD

and in some months there were half as many at TMD compared to

RMD. The difference in the number of camera nights is the result of

the reduced amount of area available to survey in TMD. The amount

of gray fox activity at TMD is higher than RMD even though we had

fewer camera nights in TMD. Approximately 53.3% of gray fox images were captured in TMD. The distribution may be influenced by

Figure 4— Gray fox with a desert cottontail in its mouth. 
USDA Forest Service Proceedings RMRS-P-67. 2013 453

Analysis of the Seasonal Activity Rate of Sympatric Carnivores and their Prey . . . Benton, Cerda-Navarro, Keck, and McKnight

the difference in the amount of sampling conducted in each district,

and gray fox activity could potentially be much higher each month

than previously thought. Future studies using camera data at Saguaro

National Park should account for the differences between districts by

analyzing them as separate entities or by standardizing the number

of camera nights in each district

	 Not only did the BioBlitz alter the October 2011 data collection,

it may have altered wildlife behavior. The increased human activity

of the BioBlitz event may have influenced the behavior of gray foxes

within the park and forced them to relocate to more isolated areas

where cameras were more likely placed. Based on our photographic

results, October 2011 saw the highest amount of gray fox activity and

it may have been human influenced. The mere presence of a wildlife

camera may also cause gray foxes to alter their behavior. Remote

camera trapping did not allow us to identify individuals. Because we

can only count activity at a camera, we do not know if the camera

is recording the activity of one or 100 individuals. For example, a

single animal could be utilizing the same portion of a wash due to

its proximity to the animal’s den. Park employees have noticed over

the years that gray foxes appear to be more curious about wildlife

cameras than many other species. One example of this is on a plot

in TMD where two gray foxes loitered extensively near a specific

camera location,which resulted in numerous photographs ofthe same

individuals.

Conclusions 

	 Our study contributed to the understanding of the coexistence of

two sympatric carnivores and some of their prey sources in a Sky

Island ecosystem. More data collection and analysis would be possible by marking individuals or analyzing bobcat patterns in order to

identify individuals to better understand the population dynamics of

eachspecies.Due tothedelicate andever-changingcompositionofthe

ecosystem at Saguaro National Park, this study should be continued

over many years.An annualstudy would help to determine if gray fox

activity does in fact change in response to differences in food availability at different times of the year or if the activity is more greatly

influenced by external factors like annual precipitation. In addition

to an annualstudy, a more comprehensive analysis of the rodents and

lagomorphs should be conducted in order to understand their ecology directly and not just through their predators. Park biologists and

technicians can utilize this data to study the interspecific interactions

of mammals in Saguaro National Park and make informed decisions

about the management of wildlife species within the park.

Acknowledgments

	 Wewould like to thank the staff and volunteers of SaguaroNational

Park, specifically the Resource Management Department and Don

Swann,Nicholas Perkins, andKristinaRatzlaff.Wewould also like to

thank SergioAvila and the staff and volunteers at Sky IslandAlliance

for their assistance with this project. Without their countless hours of

advice and support this project would not have been possible.

References

Barrientos, R. and E. Virgós. 2006. Reduction of potential food interference

in two sympatric carnivores by sequential use of shared resources. Acta

Oecologica. 30:107-116.

Chamberlain, M. J. and B. D. Leopold. 2005. Overlap in space use among

bobcats (Lynx rufus), coyotes (Canis latrans) and gray foxes (Urocyon 

cinereoargenteus). The American Midland Naturalist. 153:171-179.

Fedriani, J. M., F. Palomares, and M. Delibes. 1999. Relations among three

sympatric Mediterranean carnivores. Oecologia. 121:138-148.

Fedriani, J. M. and T. K. Fueller. 2000. Competition and intraguild predation

among three sympatric carnivores. Oecologia. 125:258-270.

Fritzell, E. K. and K. J. Haroldson. 1982. Urocyon cinereoargenteus. Mammalian Species. 189:1-8.

Haynes, L., J. Lamberton, C. Craddock, [and others]. 2010. Mountain lions

and bobcats of the Tucson Mountains: Monitoring population status and

landscape connectivity. Pima County Final Report.

Lariviére,S. andL.R.Walton.1997. Lynx rufus.MammalianSpecies.563:1-8.

Neale, J. C.C. and B. N. Sacks. 2001. Food habits and space use of gray

foxes in relation to sympatric coyotes and bobcats. Canadian Journal of

Zoology. 79:1794-1800.

Phillips, S.J. and P. W. Comus. 2000.Anatural history of the Sonoran Desert.

Tucson: Arizona-Sonora Desert Museum. 628 p.

Riley,S.P.D.2006.Spatial ecologyofbobcats andgrayfoxesinurbanandrural

zones of a national park. Journal of Wildlife Management. 70:1425-1435.

Swann, D. E. 2011. Mammals of Rincon Mountain District, Saguaro National

Park. Natural Resource Report NPS/SODN/NRR—2011/437. National

Park Service, Fort Collins, CO.

Tekiela, S. 2008. Mammals of Arizona field guide. Adventure Publications

Inc., Cambridge, MN.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
454 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Wildlife Surveys and Monitoring With the Use of Remote 

Camera Traps in the Greater Oak Flat Watershed Near 

Superior, Arizona

Roger Featherstone

	 Director, Arizona Mining Reform Coalition

Sky Jacobs

	 Wild Sonora, Tucson, Arizona

Sergio Avila-Villegas

	 Sky Island Alliance, Tucson, Arizona

Sandra Doumas

	 University of Arizona, Tucson, Arizona

Abstract—In September 2011, we initiated a 2-year “camera trap” mammal survey in the Greater Oak Flat Watershed 

near Superior, Arizona. Our survey area covers a total of 6,475 ha. The area surveyed is primarily a mixing zone of 

upper Sonoran Desert and interior chaparral, with influences from the Madrean vegetation community. Elevations 

range from 1150 to 1450 m. Ten cameras were deployed in early October of 2011 and information was gathered 

and analyzed from that date to April 2012. We located cameras primarily in riparian and xero-riparian drainages. 

Locations were chosen as logical wildlife corridors to obtain a sampling of wildlife while allowing relatively easy 

access to cameras for data collection. To date, we have identified 13 mammalian species, including bobcat (Lynx 

rufus), ringtail (Bassariscus astutus), and coati (Nasua narica).

Introduction 

	 The study focuses on the Greater Oak Flat Watershed east of the

town of Superior, Arizona, and approximately 100 km east of downtown Phoenix, Arizona. The survey area is bounded on the west by

“Apache Leap,” no more than 1 km north of Highway 60, 1 km east

of GaanCanyon (known as Devil’sCanyon on most maps) on the east

and an arbitrary southern limit. The survey area is primarily public

land managed by the USDA Forest Service, Tonto National Forest

(see fig. 1 for map of the survey area).

	 Pyroclasticweldedtuft,specifically“ApacheLeaptuff,” isthemost

common substrate throughout our study area (USDA Forest Service

2010).These formations create a generally rugged and steep topography with deep canyons and jagged spires and ridges. The watershed

is drained by Gaan Canyon, which flows year-round through most of

the survey area and by QueenCreek. Previous flora and fauna surveys

have shown that Gaan Canyon is botanically diverse and supports

a high diversity of bird species (Jacobs 2009). Eleven special status

bird species exist within 8 km of the project area according to review

tools provided by theArizona Game and Fish Department (AZGFD).

The area surveyed is primarily amixing zone of upper SonoranDesert

and interior chaparral, with some influence of Madrean evergreen

woodland.The elevation rangesfrom approximately 1150 to 1450 m.

	 Interior chaparral vegetation includes manzanita (Arctostaphylos 

pungens), catclaw acacia (Acacia greggii), desert broom (Baccharis 

centennial), and scrub oak (Quercus turbinella) (Spangle 2008).

Other common upland speciesinclude hop bush (Dodonaea viscosa),

birchleaf mountain mahogony (Cercocarpus betuloides), jojoba

(Simmondsia chinensis), wait- a-minute bush (Mimosa biuncifera),

cholla (Opuntia sp.), and agave (Agave sp.) (Jacobs Avi). Vegetation

composition throughout the uplands is significantly influenced by

Arizona Uplands division Sonoran Desert elements as evidenced

by the presence of saguaros (Carnegiea gigantea), which are fairly

common on rocky east- and south-facing slopes.

	 The primary human uses oftheGreaterOak FlatWatershed include

recreation, mining, and cattle grazing (Spangle 2008).Amining company is investigating the area for a large underground copper mine

and is conducting pre-feasibility drilling. Federal legislation has been

introduced to privatize much of the study area to accommodate the

construction of a large underground mine. A portion of the Greater

Oak Flat Watershed was set aside from mining by executive order in

1955 (Federal Register 1955).

	 There has been little to no survey of land mammal species in the

study area to date and no comprehensive study of human recreational

activities in the study area. This study will assist land managers and

decision makers in understanding movement, behavior patterns, and

distributions of species that use the watershed as well as the use and

movement of human recreational activities within the watershed.

	 Remotecamerasareextremelyusefulforthestudyofrare,threatened

or endangered species, and/or elusive or cryptic animals. In recent

years they have been used to evaluate the presence and abundance
USDA Forest Service Proceedings RMRS-P-67. 2013 455

Wildlife Surveys and Monitoring With the Use of Remote Camera Traps . . . Featherstone, Jacobs, Avila-Villegas and Doumas

of jaguars, ocelots, and other mammals (Karanth and Nichols 1998;

Avila 2007; McCain and Childs 2008). Remote cameras are a safe,

non-invasive technique, and in some cases can provide estimations

of wildlife population densities (Silver 2004; Silver and others

2004). In 2007, Sky Island Alliance initiated a remote camera study

on cross-border permeability in northern Sonora, Mexico, with the

goal of identifying wildlife corridors that connect the northernmost

populations of jaguars and ocelots with individuals documented in

southern Arizona; the study quickly documented ocelots in Sonora’s

Sierra Azul (Avila 2007).

Oak Flat and the Ocelot

	 In 2010 the AZGFD reported a road-killed ocelot (Leopardus 

pardalis) found on Highway 60 east of Superior, Arizona, between

Oak Flat Campground and Top of the World, possibly representing

the northernmost record for the species. DNA analysis from the U.S.

Fish & Wildlife Service’s forensic lab shows the ocelot was of wild

origin. However, due to lack of additional samples or sequence data

from Sinaloa or Sonora, the lab was not able to determine whether

the ocelot was of Sonoran origin (De Young and Holbrook 2010).

	 Theocelothasbeenassociatedwithawide rangeofhabitats,including mangrove forests, savannah grasslands, thornscrub, and tropical

forests of all types(Lopez-Gonzalez and others 2003).Recentrecords

by Avila (2007) document ocelots in Madrean evergreen woodland,

in elevations above 1200 m .

	 Lopez-Gonzalez and others (2003) emphasize the importance of

considering the ocelot’s short dispersal distances (5-25 km) when

protecting and connecting patches of suitable ocelot habitat. The

location ofthe road-killed ocelotreported byAZGFD isin the interior

chaparral vegetation community, at an elevation of 1334 m within

the Greater Oak Flat Watershed.

Objectives 

	 Our primary objective wasto conduct non-invasive surveys ofland

mammal species and secondarily to survey human activities in the

Greater Oak Flat Watershed. We were interested in gaining baseline

data due to future potential habitat destruction from construction and

operation of a proposed underground block cave mine in the area.An

underground block cave mine would create a void the size of the ore

body that leads to subsidence and the altering of water flow patterns

(Featherstone 2012) .

	 We conducted a preliminary habitat survey prior to placement of

cameras, which indicated sufficient habitat for good land mammal

diversity. An initial literature search showed that, to date, no intensive mammal surveys have been conducted in the Greater Oak Flat

Watershed.

Methodology 

	 We adopted a standardized remote camera protocolto validate presence/absenceofmammalspeciesinagivenarea (Chavez andCeballos

2006).The selection of camera siteswas chosencarefullytomaximize

probability for photographing land mammals. We established basic

criteria to select camera locations using regional topographic maps,

satellite imagery, and GIS surveying for the following variables:

topography, geographic connection of mountain ranges, elevation,

vegetation type, presence of temporary or permanent water source,

and size of corridors (arroyos).

	 Ten remote cameras are located within a 65-km2 study area. We

are using Cuddeback Attack IR cameras within lockable bear proof

camera safes (Cuddeback 2012). Cameras and safes were supplied

at a discount by Cuddeback and we are grateful for their support.

The cameras are equipped with 4 GB memory cards to assure ample

storage space between camera checks.

Figure 1—Map of survey region, camera trap locations, and location of road-killed 

ocelot, Greater Oak Flat Watershed near Superior, Arizona, October 2011-April, 2012.
456 USDA Forest Service Proceedings RMRS-P-67. 2013

Featherstone, Jacobs, Avila-Villegas and Doumas Wildlife Surveys and Monitoring With the Use of Remote Camera Traps . . . 

	 After deployment in select areas, cameras were revisited every 4

to 5 weeks, based on battery life and memory card space. We did not

use any type of attractants, lure, or bait near our cameras, to avoid

species bias or modification of behavior. Cameras are placed with

minimal disturbance to the surrounding vegetation, although care is

taken to make sure that false triggers from blowing vegetation are

kept to a minimum. All remote cameras were set to display time and

date on photographs helping us better evaluate daily activity patterns.

	 Cameras were set to record a still photograph followed by 30 seconds of video. Several cameras have been moved during the course

of the survey to date. Potential for damage/theft based on proximity

to human activity prompted us to reposition one of our cameras. (A

camera was stolen with only one month of data accumulated.) More

than one camera was moved due to a lack of mammal activity at the

site. As the study matures, we will be better able to gauge whether

to move cameras for optimal observation or to leave them in place

to record seasonal fluctuations in activity.

	 In order to manage the numerous photographs in an organized

manner, we developed a system for labeling remote cameras, their

location, and corresponding memory cards. After each field visit,

we collected information on all wildlife species, time and date of

each photo-event and observations, including gender, approximate

age, health status, number of animals in photo, and behavior, into a

database for the purpose of analyzing data systematically.

	 Both photographs and subsequent 30-second videos were used

to tally species numbers. Redundant counts were reduced, such as

when an animal stayed in the vicinity and was captured multiple

times. However, if good judgment would indicate that more than one

individual was present, all were counted, even if all were not in the

frame at the same time (e.g. an individual runs across the frame at

the beginning of the video and a second individual runs in the same

direction later). Differentspeciestogether(dogs with people or hikers

with trucks) generated two records, each counted separately. Skunks

and deer were not identified to species for the tallies. People on foot

or horseback were counted individually but a vehicle (truck, ATV,

motorcycle) was counted as one, regardless of the number of occupants. Birds were recorded, but not counted because their detection

wasincidental to thissurvey. People and domestic dogsthat could be

identified as the same individuals were counted only once if detected

again within 10 minutes. Those returning later in the day, however,

were counted again.

	 Camera locations were grouped into three types of terrain depending on the slope (flat, wide, and narrow) as calculated from a 10-m

resolution digital elevation map (DEM) averaged within 25 m and

100mbufferssurroundingeachcamera location(fig.2).Flat terrain had

slopes of less than 15 percent at both 25 and 100 m, wide canyons

are less than 15 percent at 25 m but greater than 15 percent at 100

m and narrow canyons are characterized as having greater than 15

percent of slope at both 25 and 100 m.

	 “Effort” varied across 15 camera trap locations because cameras

were moved or removed during the survey period. The number of

camera-days was used to adjust summary statistics for this varying

effort. Cameras were in place for a total of 1750 camera-days.

Results

	 We have identified 13 species of wild mammals, as well as humans

and domestic animals, in 15 camera locations (see table of species in

table 1).We had 418 detections of wild mammals or 0.24/camera-day

of 13 different species (deer were counted as 1 species, striped and

hooded skunks were counted as 1 species, table 1). Mean detections

of wild mammal individuals across 15 locations was 0.414 ± 0.438

(95% CI) detections/camera-day (range 0.01-3.5).

	 Sightings per camera broken down by species are shown in figure

3. The greatest variety and number of species were found in the wide

canyon group with one location showing as many as 4.5 sightings

per day for all species.

	 We grouped species photographed into four categories:mesocarnivores, herbivores, domestic, and human (including trucks andATVs).

Figure 4 showsthe distribution of totalspeciesfrom each category by

terrain type.Mesocarnivoresincluded ringtail,raccoon, coyote, coati,

bobcat,skunks, andgrayfox.Herbivoresincludedsquirrels, cottontail,

Figure 2—Mean percent slope within 25-m and 100-m circular buffers surrounding each camera location, as calculated from 10-m resolution DEM, Greater Oak Flat Watershed near Superior, Arizona.
USDA Forest Service Proceedings RMRS-P-67. 2013 457

Wildlife Surveys and Monitoring With the Use of Remote Camera Traps . . . Featherstone, Jacobs, Avila-Villegas and Doumas

Table 1—Species detected by camera traps in Greater Oak Flat 

Watershed near Superior, Arizona, October 2011-April 2012. 

Common name Genus species

bobcat Lynx rufus

coati Nasua narica

cottontail Sylvilagus audubonii

coyote Canis latrans

deer Odocoileus virginianus 

 (white-tailed), O. hemionus (mule)

gray fox Urocyon cinereoargenteus

javelina Pecari tajacu

striped & hooded skunk Mephitis mephitis (striped), 

 M. macroura (hooded)

spotted skunk Spilogale gracilis

hog-nosed skunk Conepatus mesoleucus

raccoon Procyon lotor

ringtail Bassariscus astutus

rock squirrel Spermophilus variegatus

domestic dog Canis familiaris

domestic cat Felis catus

domestic cattle Bos taurus

javelin, and deer. Domestics included dogs, cats, and cattle. Humans

included those on foot, on horseback, and motorized vehicles.

	 Ourdata indicate thatthemost commonandwidelydistributedfocal

speciesin ourstudy area isthe grey fox (Urocyon cinereoargenteus).

Javelina (Pecari tajacu) was the least recorded with only 1 photograph, which may be due to camera bias(tracks of javelina have been

noted in the area). Preliminary results indicate that mammal density

is highest in wide canyons while both numbers and species richness

are lower in narrow/steep canyons. It also appears that bobcat and

coyote favor roads for travel despite higher human traffic.

	 Data collected to date showthat human activity is highestin thewider

canyonsandisthelowestinnarrowcanyons.Theheaviestusebyhumans

occurred in March, while the least was in December (fig. 5).

Discussion

	 Although thissurvey’s primary purpose isto study land mammals,

the versatility of using camera traps to also record human activities

and movements allows us to expand the scope of the survey to also

include mapping of human recreational activities such as hiking,

rock climbing, and 4-wheel-drive activities. In most cases it is easy

Figure 3—Sittings per camera-day for camera locations, grouped by terrain type. Flat terrain had 

slopes of <15% within 25 m and 100 m, wide canyons had slopes <15% within 25 m and >15% 

within 100 m, and narrow canyons had >15% slope within 25 m and 100 m of the camera location, 

Greater Oak Flat Watershed near Superior, Arizona.
458 USDA Forest Service Proceedings RMRS-P-67. 2013

Featherstone, Jacobs, Avila-Villegas and Doumas Wildlife Surveys and Monitoring With the Use of Remote Camera Traps . . . 

Figure 4—Sitings per camera-day, grouped by mammal 

type and terrain type, Greater Oak Flat Watershed near 

Superior, AZ.

Figure 5—Sittings per camera-day, grouped by month, Greater Oak Flat Watershed 

near Superior, Arizona.

to differentiate whether we are observing a hiker or a rock climber

by the gear that is carried. We have several observations of domestic

cats (Felis catus) that appear to have gone feral, but observations of

domestic dogs (Canis familiaris) show that all have been paired with

humans. Locations with domestic cats were those closest to paved

roads and permanent human activity. Records and analysis of human

activities hopefully will not only allow us to determine whether wild

species are being displaced, but will also create a useful record for

landmanagerstoprovideongoingrecreationalopportunitiesonpublic

land.

	 The data so far show a broad distribution of herbivores and mesocarnivores, but to date is lacking records of large carnivores such as

black bear (Ursus americanus) and mountain lion (Puma concolor).

Previous wildlife surveys have indicated that black bear occur in the

study area (Jacobs and Flesch 2007) and there appears to be suitable,

if not ideal, habitat for mountain lions. Possible reasons for this may

include insufficient time to capture images of less abundant species

or that they have been displaced by increasing human activities such

those associated with drillrigs and industrial mining activities.While

recreation has been ongoing for generations in the study area (Roy

Chavez, personal communication), additional human activities such

as those associated with drill rigs and other mining equipment have

not.

Conclusions

	 The period of thissurvey to date has been the year’s cooler months

(October throughApril). We expect human recreational activity to be

higherinthe coolermonths andtodropoffoverhottersummermonths.

This project has been recording data for a relatively short time frame

of 6 months. Over the course of the study, which is planned to be 2

years or longer, clearer trends and usage patterns should develop. In

addition, a full year of data should solidify seasonal trends.

References

Avila, S. 2009. The Cuatro Gatos Project: Wildlife research and conservation

in northwestern Mexico and implications of the border fence. Wild Felid

Monitor. 2(1).

Chavez, C. and G. Ceballos. 2006. Memorias del Primer Simposio. El jaguar

Mexicano en elSigloXXI:SituaciónActual yManejo.CONABIO-Alianza

WWF Telcel-Universidad Nacional Autónoma de México. México, D.F.

Cuddeback 2012. Cuddeback Attack IR scouting camera http://cuddeback.

com/scouting_camera_products/attackIR.html [April 26, 2012]

De Young, R. and J. Holbrook. (2010). Analysis and interpretation of ocelot

material lineages from road-killed ocelots in Texas and Arizona. A report
USDA Forest Service Proceedings RMRS-P-67. 2013 459

Wildlife Surveys and Monitoring With the Use of Remote Camera Traps . . . Featherstone, Jacobs, Avila-Villegas and Doumas

to the U.S. Fish and Wildlife Service and Texas Parks and Wildlife Department. Texas A&M University, Kingsville.

Featherstone,R. 2012.Impacts of block cavemining toOakFlatCampground,

Apache Leap, and Gaan Canyon. http://www.azminingreform.org/sites/

default/files/docs/Impacts%20of%20Block%20Cave%20Mining.pdf

[May 2, 2012]

FederalRegister(1955). United States Government FederalRegister, October

1, 1955. Page 7337.

Karanth K. U. and J. D. Nichols. 1998. Estimation of tiger densities in India

using photographic captures and recaptures. Ecology. 79:2852-2862.

Jacobs, S. and Flesch,A.July 2007. Vegetation and wildlife survey of Devil’s

Canyon, Tonto National Forest. Online: www.wildsonora.com/reports.

Jacobs, S. May 2009. Vegetation and wildlife survey of Devil’sCanyon (Gaan

Canyon), Tonto National Forest. Online: www.wildsonora.com/reports.

Lopez-Gonzalez, C., Brown, D., and J.P. Gallo Reynoso. 2003. The ocelot

Leopardus pardalis in north-western Mexico: Ecology, distribution and

conservation status. Oryx. 37(3).

McCain, E., and J. Childs. 2008. Evidence of resident jaguar (Panthera onca)

in the southwestern United States and the implications for conservation. J.

of Mammalogy. 89: 1-10.

Silver, S. C., Ostro, L. E. T., Marsh, L. K, [and others]. 2004. The use of

camera traps for estimating jaguar Panthera onca abundance and density

using capture/recapture analysis. Oryx. 30(2): 148-154.

Silver, S. C. 2004. Assessing jaguar abundance using remotely triggered

cameras. Wildlife Conservation Society, Global Carnivore Program: 1-25.

Spangle, S. L. 2008. Biological opinion on Highway 60 widening (HOP-AZ,

STP-060-D(200)A, ADOT TRACS No 60 GI 236 H6140 01C) 4-5.

USDA Forest Service. May 2010. Environmental Assessment, Resolution

Copper Mining Pre-Feasibility Activities Plan of Operation: 3-9.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
460 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 461

Posters
462 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 463

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Medium and Large Mammals in the Sierra La Madera, 

Sonora, Mexico

Erick Oswaldo Bermúdez-Enríquez

Universidad de la Sierra, Moctezuma, Sonora

Rosa Elena Jiménez-Maldonado

Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Cananea, Sonora

Gertrudis Yanes-Arvayo, María de la Paz Montañez-Armenta, and Hugo Silva-Kurumiya

 Universidad de la Sierra, Moctezuma, Sonora

Abstract—Sierra La Madera is a Sky Island mountain range in the Madrean Archipelago. It is in Fracción 

V of the Ajos-Bavispe CONANP Reserve in the Municipios (= Counties) of Cumpas, Granados, Huásabas, 

Moctezuma, and Villa Hidalgo. Medium and large mammals were inventoried using camera traps. Eighteen 

Wild View 2® camera traps were deployed during four sampling periods: August- September, SeptemberNovember, and November-December (two times). The first and second sampling periods were on Ranchos 

La Bellota, La Palmita, and San Fernando in the southern Sierra La Madera. The last two sampling periods 

were on Ranchos La Mesa and El Mezquite, and Brecha CONAFOR in the northern Sierra la Madera, and 

again Brecha CONAFOR. The vegetation sampled was foothills thornscrub, oak woodland and pine-oak 

forest. Eighteen species of mammals in the orders Carnivora, Artiodactyla, Didelphimorphia, Lagomorpha, 

and Rodentia were photographed. Four species of birds, including Aquila chrysaetos (golden eagle) were 

also photographed.

Introduction

Sierra La Madera is a Sky Island mountain range located in the 

transition between the New World tropics and the northern temperate 

zone that is about 29°N in east-central Sonora, which enhances the

diversity of flora and fauna. There are very few faunal studies of any

kind in the Sky Island Region in Sonora. In 2011, Universidad de

la Sierra together with the CONANP Ajos-Bavispe Reserve began

faunal inventories using camera traps in the Sierra La Madera. Here,

we presentthe preliminary documentation ofmediumand largemammals from 2011-2012.

Study Area

The Ajos-Bavispe National Forest Reserve and National Wildlife

Refuge is a Reserve in the Comisión de Áreas Protegidas Naturales

(CONANP)systemin northeastern Sonora, Mexico.It coversmost of 

184,000 hectares in five areas. Fracción I is in Sierra Pilares de Teras

and part of the Sierra del Tigre. Fracción II is in the Sierra Pilares

de Nacozari. Fracción III is in San Diego and part of the Sierra del 

Tigre. Fracción IV includesthe Sierras de Ajos, Buenos Aires, and La

Púrica. Fracción V includes the Sierra La Madera at 29°55’N latitude

109°30’W longitude. (fig. 1). Complex topography and an elevational

range of ca. 1685 m (from 615 m along Río Bavispe at Huásabas to

over 2300 m on the highest peak)result in diverse habitats.The Sierra

la Madera is in the Municipios of Cumpas, Granados, Huásabas,

Montezuma, and Villa Hidalgo, Sonora. Vegetation in the Sierra la 

Madera along an elevational gradient is foothills thornscrub, desert

grassland, oak woodland, and pine-oak and pine forests (INEGI,

1980; Yanes-A. and others 2010).

According to Caire (1978) and other references cited in CONANP

(s/f), the potential mammal fauna in the Ajos-Bavispe Reserve is 93

species, representing 62.4% of the mammals of Sonora (CastilloGámez and others 2010) including white-tailed deer (Odocoileus 

virginianus), mule deer (O. hemionus), puma (Puma concolor), collared peccary (Tayassu tajacu), gray fox (Urocyon cinereoargenteus),

squirrels (Sciurus arizonensis, Spermophilus variegatus), rabbits 

(Lepus spp., Sylvilagus spp), and others.

Materials and Methods

Wildview Extreme 2® Camera traps were placed in habitats during

four periods from August2011 to January2012.Theywerestrategically

located at approximately 1 kilometer intervals on tree trunks 30 and 

70 cm above the ground, with a north-south orientation. Attractants 

(canned sardines or corn moistened with vanilla extract) were placed

at a distance of about 3 m from the cameras.Geographical coordinates 

andelevationswere obtained using a GarminEtrexGarmin® GPS unit.

Dominant plant species were listed to describe the habitat at each site.
464 USDA Forest Service Proceedings RMRS-P-67. 2013

Bermúdez-Enríquez, Jiménez-Maldonado, Yanes-Arvayo, and others Medium and Large Mammals in the Sierra La Madera, Sonora, Mexico

The first and second sampling periods were on Ranchos La Bellota,

La Palmita/Mesa Quemada, and San Fernando in the southern Sierra 

la Madera. The third sampling period was on Ranchos El Mesquite

and La Mesa, and onBrechaCONAFOR. During the fourth sampling 

period, six cameras were returned to Brecha CONAFOR.

Results

Eighteen species of mammals were recorded in orders Carnivora

(five families), Artiodactyla (two families), Rodentia, Lagomorpha,

and Didelphimorphia (one family each). Of mammals photographed,

55% were medium-sized mammals and 45% large mammals (table 1). 

Domesticorwildlife speciesintroduced by man were also documented, 

including cows, horses, burros, goats, and dogs. Fourteen mammal

Table 1—Wildlife species recorded in August-December 2011 by camera traps in the Sierra 

La Madera, Montezuma, Sonora. SP = sampling period. *Species with protection category 

according to NOM-059-SEMARNAT-2010.

Order Family Species SP 1 SP 2 SP 3 SP 4

Carnívora Canidae Urocyon cinereoargenteus X X X X

 Canis latrans X X X

Felidae Puma concolor X X X X

 Leopardus pardalis* X X

 Lynx rufus X X X X

Ursidae Ursus americanus* X

Mephitidae Mephitis macroura X X X

 Conepatus mesoleucus X X X X

 Spilogale putorius X X X X

Procyonidae Nasua narica X X X

 Bassariscus astutus X X

 Procyon lotor X X

Artiodactyla Cervidae Odocoileus virginianus X X X X

Tayassuidae Tayassu tajacu X X X

Didelphimorphia Didelphidae Didelphis virginiana X X

Lagomorpha Leporidae Sylvilagus floridanus X X

Rodentia Sciuridae Sciurus arizonensis X X

Rodentia Sciuridae Spermophilus variegatus X X X X

Figure 1—Study area, Ajos-Bavispe National Forest Reserve and national wildlife Refigie in northeastern Sonora. 

Fracción V includes the Sierra La Madera at 29°55'N lattitude 109°30'W longitude.
USDA Forest Service Proceedings RMRS-P-67. 2013 465

Medium and Large Mammals in the Sierra La Madera, Sonora, Mexico Bermúdez-Enríquez, Jiménez-Maldonado, Yanes-Arvayo, and others

specieswerephotographedduringthesecondsamplingperiod.Asingle

black bear (Ursus americanus) was seen at Rancho San Fernando

Discussion and Conclusions

Fifty-five percent ofthe recordsbelong tomedium-sized mammals. 

The rock squirrel (Spermophilus variegatus) was the most abundant 

species. In the large mammals, the white-tailed deer was the most 

abundant.Medium-sizedmammalsweremost commoninfall-winter,

while large mammals were seen more often in summer-fall.

	 The Sierra La Madera mammal fauna has only 19.4% of the 93 

mammal species potentially occuring in the Ajos-Bavispe Reserve

(Caire, 1978; CONANP, s / f), but small species were not sampled.

The Arizona gray squirrel (Sciurus arizonensis) was seen in the 

third and fourth sampling periods in oak woodland and pine-oak 

forest on Rancho la Mesa and Brecha CONAFOR. Thisis a 110 eastsoutheastrange extension fromArroyo Santo Domingo nearCucurpe

of a central Arizona-northern Sonora regional endemic species. The

Mexican Fox Squirrel (Sciurus nayaritensis) was not photographed

in the present study, but was previously observed on Rancho San

Fernando by Stephen L. Minter in August 2010 and the northern

Sierra La Madera by Dale Turner in September 2003 (see Madrean

Archipelago Biodiversity Assessment [MABA] database, Madrean.

org). This is a Sierra Madre Occidental species reaching its western

range limits.

Three species of mammals (Leopardus pardalis, Sciurus arizonensis, U. americanus) as well as golden eagle (Aquila chrysaetos) 

are protected under NOM-059-SEMARNAT-2010. These results are

preliminary and additional camera trapsin different areas and habitats

are needed to more fully document the mammal fauna of the Sierra

La Madera, which likely contains jaguar (Panthera onca).

References

Caire, William. 1978. The distribution and zoogeography of mammals of

Sonora. PhD. Diss. Albuquerque: New Mexico.

Castillo-Gámez,ReynaA.;Gallo-Reynoso,JuanP.;Egido-Villarreal,Janitzio;

and Caire, William. 2010 Mamíferos. Pg. 421-426 in Molina-Freaner,

Francisco and Van Devender, Thomas R. eds., Diversidad Biológica de

Sonora, Universidad Nacional Autónoma de México, Hermosillo, Sonora.

CONANP. [n.d.] Estudio Previo Justificativo, Área de Protección de Flora

y Fauna Mavavi. Secretaría de Medio Ambiente y Recursos Naturales.

México. 150 p.

Norma Oficial Mexicana. 2010. NOM-059-SEMARNAT-2010. Diario Oficial de la Federación. Segunda sección. Jueves 30 de diciembre de 2010.

Mexico. 78 p.

Yanes-Arvayo, Gertrudis; Montañez-Armenta, María de la Paz ; SilvaKurumiya, Hugo; and Gil-Montaño, Enrique. 2011. Catálogo de Fauna de

Sierra La Madera. Fracción V de la Reserva Ajos Bavispe. Universidad de

la Sierra, Hermosillo, Sonora.

Yanes-Arvayo,Gertrudis;Silva-Kurumiya,Hugo;Montañez-Armenta,María

de la Paz; and Castellanos-Villegas, Alejando E. 2010. Flora de Sierra la

Madera en el estado de Sonora, México. VII Internacional Sobre la Flora

Silvestre en Zonas Áridas. Departamento de Investigaciones Científicas y

Tecnológicas de la Universidad de Sonora. Hermosillo, Sonora. México.

Del 17-19 de marzo.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
466 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Herpetofauna at the Appleton-Whittell Research Ranch

Roger C. Cogan

Appleton-Whittell Research Ranch of the National Audubon Society, Elgin, Arizona

Abstract—A rich diversity of amphibian and reptile species occurs at the Appleton-Whittell Research Ranch, 

an 8000-acre sanctuary for native biota and research facility in the semi-arid grasslands of southeastern 

Arizona, created in 1969 and managed by the National Audubon Society since 1980. Nine species of amphibians and 42 species of reptiles have been identified by staff and researchers within the preserve. Efforts 

are underway to document the current richness of the herpetofauna. Recent surveys in 2010–2012 have 

confirmed continued presence of 26 species. As part of that inventory effort, we located seven overwintering 

sites of rattlesnakes. Our challenge into the future is to adaptively manage the Research Ranch to provide 

sanctuary to the appropriate plant and animal species under a likely changing climate. 

Introduction

	 Sincecattlewereremovedinthe1960scollectingeffortsoverseveral

decades by numerous researchers have identified 9 amphibian and

42 reptile species representing 29 genera within the preserve. There

have been ongoing investigations with several individual species.

However, the herpetofauna as a whole has not been assessed. Efforts

are currently ongoing to locate and document the continued existence

or absence of all herp species that occur within the Research Ranch

boundaries and to discover and track use of wintering sites used by

rattlesnakes.

Methods and Results

	 The Research Ranch management area covering 8000 acres is

primarily semi-arid grassland. Partners include the Bureau of Land

Management, U.S. Forest Service,ResolutionCopperCompany,The

Research Ranch Foundation, and The Nature Conservancy. In addition to its own property, Audubon manages parcels of land owned by

these partners under various contractual agreements. Conservation

and research are key elements of management agreements. 

	 Searchesfor amphibians andreptiles are conductedwhenconditions

are appropriate for herp surface activity; however, most encounters

occurduringtimeswhenstaff andresearchers are travelingorworking

in the field. When sightings occur, they are documented with photographs whenever possible and recorded. Unique and rare sightings

are recorded with GPS for future reference.

Conclusions

	 During the history of the Research Ranch there have been several

surveys for herps and individual species have been investigated (i.e. 

Dodero and Spengler, unpublished checklist; Smith and Chiszar

2002). This is the first attempt to monitor and document presence

or absence of all herp species previously identified at the sanctuary.

Since 2010 surveys have confirmed that 26 species are still present at

the Research Ranch (table 1). Seven rattlesnake wintering sites have

thus far been identified and located. These locations are primarily

located adjacent to wash drainages in rock outcrops or boulder piles.

They are occupied by Western Diamondback Rattlesnakes (Crotalus 

atrox) (fig. 1) and Black-tailed Rattlesnakes (Crotalus molossus).

These sites are also utilized by Sonoran Gopher snake (Pituophis 

catenifer affinis) and Sonoran Whipsnake (Masticophis bilineatus).

Rockrattlesnake (Crotalus lepidus) andMohave rattlesnake (Crotalus 

scutulatus) wintering sites currently have not been located. Efforts

are underway to locate other preferred wintering sites.

Reference

Smith, Hobart M.; Chiszar, David. 2000. The herpetofauna of the Research

Ranch. In: Kennedy, Linda; Seltzer, Stephanie, editors. Audubon Research

Ranch2000.NationalAudubonAppleton-WhittellResearchRanch,ElginAZ..

Table 1—Herpetofauna identified at the Research Ranch.

Amphibians: Nine species, of seven genera

Red-spotted Toada (Anaxyrus punctatus)

Sonoran Desert Toada (Ollotis alvarius)

Couch’s Spadefoot Toad (Scaphiopus couchii)

Chihuahua Spadefoot Toada (Spea multiplicata stagnalis)

Canyon Treefroga (Hyla arenicolor)

Tiger Salamander (Ambystoma mavortium ssp.)

Chiricahua Leopard Frog Lithobates chiricahuensis)

 Believed to be extirpated

Lowland Leopard Frog (Lithobates yavapaiensis)

 Believed to be extirpated

American Bullfroga (Lithobates catesbeiana)

 Invasive non-native
USDA Forest Service Proceedings RMRS-P-67. 2013 467

Herpetofauna at the Appleton-Whittell Research Ranch Cogan

Lizards: Nineteen species, of eight genera

Arizona Striped Whiptail (Aspidoscelis arizonae)

Canyon Spotted Whiptail (Aspidoscelis burti

 stictogrammus)

Chihuahuan Spotted Whiptail (Aspidoscelis exsanguis) 

Gila Spotted Whiptail (Aspidoscelis flagellicaudus) 

Sonoran Spotted Whiptaila (Aspidoscelis sonorae)

Sonoran Tiger Whiptail (Aspidoscelis tigris

 punctilineatus) 

Desert Grassland Whiptaila (Aspidoscelis uniparens) 

Eastern Collared Lizard (Crotaphytus collaris)

Madrean Alligator Lizarda (Elgaria kingii nobilis)

Mountain Skink (Plestiodon callicephalus) 

Great Plains Skink (Plestiodon obsoletus)

Chihuahuan Earless Lizarda (Holbrookia maculata

 flavilenta)

Greater Short-horned Lizarda (Phrynosoma hernandesi)

Round-tailed Horned Lizard (Phrynosoma modestum) 

Regal Horned Lizard (Phrynosoma solare)

Clark’s Spiny Lizarda (Sceloporus clarkii)

Slevin’s Bunchgrass Lizarda (Sceloporus slevini)

Southwestern Fence Lizarda (Sceloporus cowlesi)

Ornate Tree Lizarda (Urosaurus ornatus linearis) 

Snakes: Twenty species, of twelve genera

Western Diamondback

 Rattlesnakea (Crotalus atrox)

Mojave Rattlesnakea (Crotalus scutulatus)

Rock Rattlesnakea (Crotalus lepidus) 

Figure 1—Western Diamondback Rattlesnake (Crotalus atrox).

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.

Black-tailed Rattlesnakea (Crotalus molossus)

Regal Ringneck Snakea (Diadophis punctatus regalis)

Chihuahuan 

 Hook-nosed Snake (Gyalopion canum)

Mexican Hognose Snake (Heterodon nasicus kennerlyi)

Spotted Nightsnake (Hypsiglena torquata

 ochrorhynchus)

Western Black Kingsnake (Lampropeltis getula nigrita)

Desert King Snake (Lampropeltis g. splendida)

Arizona Mountain 

 Kingsnakea (Lampropeltis p. pyromelana)

Sonoran Whipsnakea (Masticophis bilineatus)

Sonoran Coachwhip (Masticophis flagellum

 cingulum)

Sonoran Coral Snake (Micruroides e. euryxanthus)

Sonoran Gopher Snakea (Pituophis catenifer affinis)

Western Patchnose Snake (Salvadora deserticola)

Eastern Patchnose Snakea (Salvadora g. grahamiae)

Western Groundsnakea (Sonora semiannulata)

Western Black-necked

 Garter Snakea (Thamnophis c. cyrtopsis)

Mexican Garter Snake (Thamnophis eques megalops)

Checkered Garter Snakea (Thamnophis m. marcianus) 

Turtles: Two species, of two genera

Sonoran Mud Turtlea (Kinosternon sonoriense)

Desert Box Turtlea (Terrapene ornate luteola)

a

Continued presence visually observed and documented with photographs 2010-

2012.
468 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) 

From a Cave in the Huachuca Mountains, Arizona

Nicholas J. Czaplewski

Oklahoma Museum of Natural History, University of Oklahoma, Norman, Oklahoma

Steve Willsey

Hereford, Arizona

Abstract—In 2008, Steve Willsey discovered the fragmentary cranium of a bear loose on the floor of a cave 

at about 2270 m elevation near the crest of the Huachuca Mountains. In 2009, we revisited the cave to 

examine the specimen with the intention of identifying the species. We photographed and measured the 

main pieces and left them in the cave. The skull is from an adult, probably male, with prominent sagittal 

crest. Bears are highly variable morphologically and their remains are difficult to identify. The morphological 

features and measurements of the Huachuca Mountains cranium are somewhat equivocal, but most available features indicate a brown bear, Ursus cf. arctos. Some parts are encrusted with carbonate and could 

be better examined after collection and preparation as well as comparison with late Pleistocene brown and 

black bears. Based on its state of preservation, the cranium possibly represents a late Pleistocene occurrence, which could be determined by radiometric dating. There is no previous fossil record of U. arctos in 

the Sky Islands, nor in the rest of Arizona, New Mexico, Chihuahua, or Sonora. Despite the lack of other 

fossil records in the region, this late Quaternary occurrence affirms the historic records of the species in the 

Apache Highlands before extirpation. 

Introduction

	 As with the dynamics of diverse indigenous cultures of Apacheria

beforecolonization,anunderstandingofthepresent-daybiodiversityof

theApacheHighlandsregioncanbenefitfromthe temporalperspective

ofrecent geological time (Martin 2005).The Quaternary fossilrecord

may also inform certain management decisions. Historically, grizzly

(brown) bears occurred throughouttheSky Islandsregion and beyond

(see map of historic records in Brown, 1985:42-43; also in Brown

and Davis 1995;Cortés-Calva 2000; Pavlik 2006).Most grizzly bears

were killed out during the second half of the nineteenth century, with

few surviving in the Southwest until the mid-twentieth century. The

latesthistoricalrecordinsouthwesternNorthAmericawas about1976

in northern Sonora (Gallo-Reynoso and others 2008). Black bears

are known widely throughout Apacheria from late Pleistocene to the

present day. We report an as-yet undated Quaternary occurrence of

a probable brown bear, Ursus cf. U. arctos, in a high-elevation cave

in the Huachuca Mountains, Cochise County, Arizona.

Materials and Methods

	 Steve Willsey discovered the fragmentary cranium of a bear loose

on the floor of a cave at about 2270 m elevation near the crest of the

HuachucaMountains,Arizona, in 2008.We revisited the cave in 2009

to examine the specimen with the intention of identifying the species.

We photographed and measured the main pieces and left them in the

cave. The skull has a prominent sagittal crest indicating an adult,

probably male individual. Some parts are encrusted with carbonate

and could be better examined after collection and preparation as well

as direct comparison with late Pleistocene brown and black bears.

Bears are highly variable morphologically and their remains are difficult to identify. In parts of NorthAmerica, many black bears(Ursus 

americanus) were larger in the late Pleistocene and underwent a size

reduction in the Holocene, further complicating the identification of

fossil bears.

	 The cranium is broken into several pieces (fig. 1). The braincase

is mostly intact and forms the largest piece with the ethmoid plate

preserved anteriorly. Both zygomatic arches are broken away at their

bases, the left mastoid, petrosal, and occipital condyle are missing.

The sagitttal crest is long, tall, and strong. A fragment of the right

frontal bone includes the postorbital process. The right maxillapremaxilla is broken into two pieces and includes the anterior root

of the zygomatic arch. Several small pieces also are present. Teeth

I1 and I2 are not preserved, while I3 and C1 are broken off at the

alveolar rims but the root of each is present. Crowns of P4, M1, and

M2 are present. Posterior to the canine, the maxilla and cheek teeth

are encrusted with carbonate; however, at least two empty alveoli for

smallpremolars arepresent, oneposteriortoC1andone anteriortoP4.

P4 is encrusted with carbonate but appears to have a well-developed

median accessory cusp. M2 is broadest anteriorly rather than at its

midpoint.
USDA Forest Service Proceedings RMRS-P-67. 2013 469

 Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) From a Cave . . . Czaplewski and Willsey

Results and Discussion

	 Qualitative morphological characteristics of the cranium and teeth

distinguish the Huachuca Mountains craniumas amember ofthe subfamily Ursinae (brown and black bears) rather than the Tremarctinae

(short-faced and spectacled bears), which are also known as fossils

in the Pleistocene and Pliocene epochs in the Sky Island region.

Measurements of the cranium are provided in table 1. By comparison

with recent brown and black bears from the Southwest (Hoffmeister

1986), the cave fossil shows similarities with both brown and black

bears. In two qualitative characters the fossil matches brown bears:

the median accessory cusp on P4 is normally present in brown bears

Ursus arctos and normally absent in black bears, Ursus americanus;

and the M2 being broadest anteriorly characterizes brown bears,

whereasin black bearsM2 is broadest atthemidpoint.M1 dimensions

of the fossil are consistent with grizzly bearsin quantitative mensural

charactersthathavebeenusedpreviouslytodistinguishthe twospecies

(table 1), whereas M2 dimensions are larger than or consistent with

the largest recent black bears (Miller and others 2009) but near the

average for Sonoran brown bears. Gordon (1977) suggested that the

M1 is greater than 20.4 mm long and 10.5 mm wide in grizzly bears

and less in black bears; the Huachuca Mountains bear’s M1 is 20.8 x

15.6 mm. The Huachuca Mountains bear’s cranium is also larger in

the few available comparative measurements than a late Pleistocene

fossil black bear from Papago Springs Cave, Canelo Hills, Arizona

(Skinner 1942). The morphological features and measurements of

the Huachuca Mountains cranium are somewhat equivocal but most

available featuresindicate a brown bear, Ursus cf. arctos.Based on its

stateofpreservation,the craniumpossiblyrepresents a latePleistocene

occurrence, which could be determined by radiometric dating. There

is no previous fossil record of U. arctos in the Sky Islands, nor in the

rest of Arizona, New Mexico, Chihuahua, or Sonora. If the fossil is

collected, it could be cleaned of carbonate and repaired. It could then

be comparedwithrecentspecimensfromsouthwesternNorthAmerica

of black and brown bears, and also with late Pleistocene black bears

with a view toward further discriminating the species.

	 The Huachuca Mountains fossil is important because it provides

the first late Pleistocene fossil record of a probable brown bear in

Apacheria and in all ofArizona,NewMexico,Chihuahua, andSonora

(Arroyo-Cabrales and others 2002; Harris 1993, 2005, 2012; Mead

and others 2005; Morgan and Lucas 2005; White and others 2010).

Although they might not have been continuously contemporaneous

in the region, this record indicates that three bears, U. cf. arctos, U. 

americanus, and Arctodus simus(giantshort-faced bear; Czaplewski

unpublished data) occurred in the Madrean Archipelago in the late

Pleistocene. Stable isotope analyses indicate that short-faced bears

were highly carnivorous and possibly specialized as scavengers

compared to brown bears (Matheus 1995). Perhaps the presence of

Figure 1—Portions of a Quaternary bear cranium from a cave in the Huachuca Mountains, Arizona. A-B, braincase in right lateral (A) and dorsal (B) 

views. C-D, right half of rostrum with parts of the jugal, palatine, premaxillary, and maxillary including the root of the upper canine, alveoli for small 

premolars, and intact P4-M1-M2, in lateral (C) and palatal (D) views. Much of the teeth and bone surface is encrusted with carbonate.
470 USDA Forest Service Proceedings RMRS-P-67. 2013

Czaplewski and Willsey Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) From a Cave . . .

the carnivorous short-faced bear constrained the body size of grizzly

bears in the Pleistocene.

	 Pavlik (2006) reviewed late Pleistocene records of black bears

in the Huachuca Mountains and Sky Islands as well as many other

aspects of their biology and conservation. He noted that black bears

still occur in the Huachuca Mountains in modest numbers amid increasing pressure from human encroachment, habitat loss, and recent

drought and fire. Although undated, the occurrence of a brown bear

in the Huachuca Mountains at some time during the late Quaternary

affirms the historic records of the species in the Apache Highlands

before it was eradicated and lends support to estimates of the suitability of still-existing habitat there for the species (Povilitis 1995).

We must adopt a cultural ecological moral to share this region with

the bears and other living things, understand that they and we are

integrative forces in the region, and pass on the same responsibilities

to our children in the greater scheme of conserving bears across the

region.

Acknowledgments

	 We acknowledge the help of Brenda Haynes, Ellen Preiss, and

Kyle Davies in the fieldwork and Juan Pablo Gallo Reynoso, Steve

Pavlik, and Sheridan Stone for helpful comments on the manuscript.

References

Anderson, Sydney. 1972. Mammals of Chihuahua: taxonomy and distribution. Bulletin of the American Museum of Natural History 148: 149-410.

Arroyo-Cabrales,Joaquín; Polaco,O.J.; and Johnson,E. 2002.Lamastofauna

del cuaternario tardío enMéxico.Pp. 103-123 inM.MontellanoBallesteros

andJ.Arroyo-Cabrales(eds.)Avancesenlosestudiospaleomastozoológicos.

ColecciónCientífica,SerieArqueología,InstitutoNacionaldeAntropología

e Historia, México, D. F.

Brown, David E. 1985. The Grizzly in the Southwest: Documentary of an

extinction. University of Oklahoma Press, Norman. 274 pp.

Brown, D. E.; and Davis, R. 1995. One hundred years of vicissitude: Terrestrial bird and mammal distribution changes in theAmerican Southwest,

1890-1990. Pp. 231-244 in Biodiversity and management of the Madrean

Archipelago:TheSkyIslandsofsouthwesternUnitedStatesandnorthwestern

Mexico. Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture,

Forest Service, Rocky Mountain Forest and Range Experiment Station,

Fort Collins, CO. 670 pp.

Cortés-Calva,P.2000.FamiliaUrsidae.Pp.707-716inS.T.Álvarez-Castañeda

and J. L. Patton (eds.) Mamíferos del Noroeste de México. Centro de

Investigaciones Biológicas del Noroeste, S. C.

Gallo-Reynoso, Juan Pablo; Van Devender, T. R.; Reina-Guerrero, A. L.;

Egido-Villarreal, J.; and Pfeiler, E. 2008. Probable occurrence of a brown

bear (Ursus arctos) in Sonora, Mexico, in 1976. Southwestern Naturalist

53:256-260.

Gordon, K. 1977. Molar measurements as a taxonomic tool. Journal of Mammalogy 58:247-248.

Harris,Arthur H. 1993. Quaternary vertebrates of New Mexico. New Mexico

Museum of Natural History and Science Bulletin 2:179-197.

Harris, Arthur H. 2005. Caves as unique resources for Pleistocene vertebrate

faunas. New Mexico Museum of Natural History and Science Bulletin

28:249-251.

Harris, Arthur H. 2012. Pleistocene vertebrates of New Mexico and transPecosTexas. Online: www.utep.edu/leb/pleistNM/forword.htm.[accessed

4 April 2012.]

Hoffmeister, Donald F. 1986. Mammals of Arizona. University of Arizona

Press and Arizona Game and Fish Department.

Martin, Paul S. 2005. Ghostly grazers and sky islands. Pp. 26-34 in Connecting mountain islands and desert seas: Biodiversity and management of

the Madrean Archipelago II. Proceedings RMRS-P-36. U.S. Department

of Agriculture, Forest Service, Rocky Mountain Research Station, Fort

Collins, CO. 631 pp.

Matheus, Paul E. 1995. Diet and co-ecology of Pleistocene short-faced bears

and brown bears in eastern Beringia. Quaternary Research 44:447-453.

Table 1—Measurements (in mm) of intact portions of the Huachuca Mountains, Arizona, fossil bear cranium 

compared with some of the same measurements published for recent bears in the Sky Islands region. 

Element Huachuca Ursus Ursus Ursus Ursus

measured Mts. Ursus arctos arctos arctos americanus

 cranium SONa CHIb AZ, NMc AZ, NMd

Foramen magnum diameter [35]e

Breadth across occipital condyles [71]

Braincase breadth 109.3 89 96.7

(95-98)

Maxillary toothrow length 119.7

C1 length 22.0

C1 width 15.6

P4-M2 length 63.5 56 74.07 53.07

(67.2-80.0) (50.1-57.6)

P4 length 13.5 19.2 21.7

 (20.9-21.9)

P4 width 11.4

M1 length 20.8

M1 width 15.6

M2 length 29.1 26.8 36.09 25.31

(33.1-39.9) (22.7-28.6)

M2 width 16.6 15.4 19.62 15.08

(18.2-20.5) (13.8-16.0)

a

One U. arctos of unknown sex from Sonora (SON) provided by Gallo-Reynoso et al. (2008). b

Mean and observed range of 3 male U. arctos from Chihuahua (CHI) provided by Anderson (1972). c

Mean and range of 10 male Ursus arctos from Arizona (AZ) and New Mexico (NM) provided by Hoffmeister (1986:tables 5.72, 5.73). d

Mean and range of 8 male Ursus americanus from Arizona and New Mexico (Hoffmeister 1986:tables 5.72, 5.73). e

Brackets [ ] indicate estimated measurement of broken element.
USDA Forest Service Proceedings RMRS-P-67. 2013 471

 Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) From a Cave . . . Czaplewski and Willsey

Mead,JamesI.;Czaplewski,N.J.; andAgenbroad,L.D. 2005.Rancholabrean

(late Pleistocene) mammals and localities of Arizona. R. D. McCord (ed.)

Vertebrate Paleontology of Arizona. Mesa Southwest Museum Bulletin

11:139-180.

Miller, Edward H.; Mahoney, S. P.; Kennedy, M. L.; and Kennedy, P. K. 2009.

Variation,sexual dimorphism, and allometry inmolarsize ofthe black bear.

Journal of Mammalogy 90(2):491-503.

Morgan, Gary S.; and Lucas, S. G. 2005. Pleistocene vertebrate faunas in

New Mexico from alluvial, fluvial, and lacustrine deposits. New Mexico

Museum of Natural History and Science Bulletin 28:185-248.

Pavlik, Steve. 2006. Ursus in a Sky Island Range: The ecology, history, and

management of black bears in the Huachuca Mountains of southeastern

Arizona. 58 pp. Online: http://blogs.nwic.edu/pavlik/papers/.

Povilitis, Tony. 1995. A nature reserve system for the Gila River-Sky Island

region of Arizona and New Mexico: Some preliminary suggestions. Pp.

534-541 in Biodiversity and management of the Madrean Archipelago:

The Sky Islands of southwestern United States and northwestern Mexico.

Gen. Tech. Rep. RM-GTR-264. U.S. Department of Agriculture, Forest

Service, Rocky Mountain Forest and Range Experiment Station, Fort

Collins, CO. 670 pp.

Skinner, Morris F. 1942. The fauna of Papago Springs Cave, Arizona, and a

study of Stockoceros; with three new antilocaprines from Nebraska and

Arizona.BulletinoftheAmericanMuseumofNaturalHistory80(6):143-220.

White, R. S.; Mead, J. I.; Baez, A.; and Swift, S. L. 2010. Localidades de

vertebrados fósiles del Neógeno (Mioceno, Plioceno y Pleistoceno): una

evaluación preliminar de la biodiversidad del pasado. Pp. 51-72 in F. E.

Molina-FreaneryT.R.VanDevender(eds.)DiversidadBiológicadeSonora.

Universidad Nacional Autónoma de Mexico, México.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
472 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

FireScape: A Program for Whole-Mountain Fire 

Management in the Sky Island Region

Brooke Gebow

The Nature Conservancy, Southeastern Arizona Preserves, Hereford Arizona; USDA Forest Service, Coronado 

National Forest, Tucson Arizona

Christopher Stetson

USDA Forest Service, Coronado National Forest, Tucson Arizona

Donald A. Falk and Corrine Dolan

School of Natural Resources and the Environment, Biological Sciences East, University of Arizona, Tucson 

Arizona

Abstract—The Coronado National Forest’s (CNF) FireScape program works to remove barriers to fire playing 

its natural role on the landscape. A long-term goal is creating landscapes that are able to survive wildfire with 

biodiversity and key ecological processes intact, especially important in the face of a drier, hotter Southwest. 

The FireScape team is nurturing multiple efforts around the Sky Islands—no two projects are alike, but those 

underway share an approach that includes multiple jurisdictions, investigations by University of Arizona 

scientists, public engagement, assessing treatment need at the whole-mountain scale, and creatively removing implementation barriers when funding is scarce. Clearance for treatments in designated Wilderness is a 

need across all projects. The FireScape team has completed a fuels map and an analysis of departure from 

reference condition for southeastern Arizona that covers the CNF and partner lands. Partners have worked 

together to update Landfire data for this 14 million-acre area. These products provide inputs for fire behavior 

and effects analyses to support decision making and outreach. The website www.azfirescape.org is a work in 

progress that displays project information, including maps, reports, and vegetation and fuels data

	 “FireScape” is a Coronado National Forest (CNF) program for

expanding safe, ecologically sound, large-scale,multi-party firemanagement across southeastern Arizona. The leadership team consists

of representatives from the Forest Service, The Nature Conservancy,

the University of Arizona (UA), and other regional land managers.

This paper represents a 2012 progress report for work that has been

underway for more than 10 years and that needs to remain a priority

forlandmanagersin the region.The 2011 fire season in the Southwest

brought a 300% increase in area burned compared with the average

for the previous 10 years—2.20 million acres (vs. 0.67 million) in

Arizona, New Mexico, and West Texas (NIFC 2011). High-severity

effects in dry woodland and forest types have left land managers uncertain about recovery trajectories (Wallow Fire example, Wadleigh

2011).

	 TheCNFcoversanumberofdispersedmountainrangesseparatedby

non-Forest Service land.These Sky Islands present different physical

and biological backgrounds and challenges as well as different sets

of neighbors. Thus they require custom approaches to fire management. Commercial timber operations have limited feasibility on this

landscape, which makes it difficult to compete for treatment funding

that ranks “utilization” significantly. The mountains and valleys of

southeasternArizona,however,have tremendousvalue aswatersheds,

habitat, working lands, and recreation sites. The importance of the

FireScape project is magnified given the 2011 fires.

	 The many goals of FireScape can be condensed into a creative

approach to fire management at a much bigger scale than in the past.

The grasslands, woodlands, and forests of southeastern Arizona

evolved under regimes of frequent low- and mixed-severity fire—on

the order of every 5 to 20 years (Kaib 1998, Swetnam and Baisan

1996). If 1.5 million acres of the CNF used to burn every 15 years,

keeping up means treating 100,000 acres/year. In some (but not all)

years the Forest might come close to half or two-thirds that acreage.

Expanding treatment and getting ahead of high-severity wildfires

not only reduces the loss of healthy systems but also costs less than

wildfire suppression and generates less smoke.

	 A long-term goal is creating landscapes that are able to survive

wildfire with biodiversity intact, which becomes especially important

inthe faceof adrier,hotterSouthwest.Projectsincludemonitoringthat

assesses oursuccess and guides adjustmentsto improve our management.Thiswork comes at a timewhen theCNF is alsomanagingmore

wildfires for resource benefit, a shift from reliance on prescribed fire

and other planned treatments to keep landscapes healthy. FireScape

also emphasizes the need to keep fire out of systems where frequent

fire has not been the norm, such as upland Sonoran desert, Arizona

chaparral, and mountaintop spruce-fir systems.
USDA Forest Service Proceedings RMRS-P-67. 2013 473

FireScape: A Program for Whole-Mountain Fire Management in the Sky Island Region Gebow, Stetson, Falk, and Dolan

	 TheFireScapeprogramisone facetoftheCNF’sworkunderwayfor

more than a decade to remove barriers to fire playing its natural role

on the landscape. Other efforts include the Peloncillo Programmatic

Fire Plan (1999) that laid out procedures for carrying out large-scale

burns in the Peloncillo Mountains while protecting the endangered

New Mexico ridge-nosed rattlesnake. The Pinaleño Ecosystem Restoration Program (2011) specifies careful mountaintop treatments

that will protect sensitive resources and safely allow fire to play a

natural role elsewhere in the range. The CNF is also a party to the

Altar Valley Fire Management Plan (2008) that proposes to restore

grasslands along and adjacent to the forest’s southwestern edge.

	 The FireScape team is currently nurturing efforts at four locations

around the Coronado—no two projects are alike, but those underway

share an approach thatincludesmultiple jurisdictions,UAscientists at

the table, assessing treatment need at the whole-mountain scale, and

completing compliance at large scale and across programs (NEPA,

Endangered Species Act section 7, and cultural resources). Getting

clearance from the Regional Forester for treatments in designated

Forest Service Wildernessis a need across all FireScape projects. We

have completed a fuelsmap and “departure” analysisforsoutheastern

Arizona that covers the CNF and partner lands. Partners worked together to correct and update Landfire data for a 14 million-acre area.

These products provide inputs for fire behavior and effects models

to support decision making and outreach.

	 In 2009, theHuachuca FireScape project completed a three-agency

Environmental Assessment (National Environmental Policy Act

[NEPA] compliance) and received aBiological Opinion (Endangered

Species Act compliance) covering a menu of treatments for 400,000

acresinsoutheasternArizona.CoronadoNationalMemorial(National

Park Service), and Fort Huachuca (U.S. Army) are the other federal

partners, but adjacent Bureau of Land Management (BLM), TNC,

Audubon, and private ranch lands are included in the greater planning area. These partners have worked together for over a decade to

coordinate fire management in an area where wildfires burn freely

across boundaries. Key treatments are large prescribed burns, but also

thinningthegrowingwildland-urbaninterface (WUI).The compliance

phase was completed with no appeals, and fire staff are implementing projects as funding becomes available. The 2011 Monument Fire

burned 30,000 acres of the Huachuca landscape.

	 The500,000-acreCatalina-RinconFireScape includesUAresearch

aimed at predicting fire behavior on landscapes that are topographically and ecologically complex, contain amixture of vegetation types,

and have recently experienced uncharacteristically severe wildfires.

The project highlights differences in how fire has shaped the Santa

Catalina versus the Rincon mountains, which share similar physical

and biological conditions, but different land use histories. Ecological

mapping was completed in 2009 and covers partner lands—CNF,

NPS, State ofArizona, and private.Treating in designatedWilderness

is a challenge for this landscape but is needed to preserve desired

ecological conditions, improve wildlife management opportunities

(particularlydesertbighornsheep),makeprescribedfiremore feasible,

and reduce threats to WUI. Public engagement is a huge component,

given the Tucson’s-backyard location. Formal compliance (NEPA,

ESA section 7, NHPA) is underway; public scoping for NEPA took

place during spring 2011.Catalina-Rincon work also analyzes carbon

cycle effects of fire treatments and coordinates with efforts to halt

the spread of invasive buffelgrass, a species that introduces fire into

non-fire-adapted upland Sonoran desert.

	 TheChiricahua FireScape focuses on restoring natural fire regimes

intheChiricahua,Dragoon, andDosCabezasmountains,plus adjacent

lands. Chiricahua National Monument and Fort Bowie National Historic Site are National Park Service partners, and the BLM, USFWS

Leslie Canyon National Wildlife Refuge, the State of Arizona, and

NaturalResourcesConservationServicehave alsojoinedtheplanning.

A number of grazing permittees who own lands adjacent to agency

holdings have elected to participate. Total acreage for this project is

running at about 500,000. Ecological mapping across jurisdictions

has been completed, and the new fuels layer and departure analysis

allowmodeling fire behavior and effects with and without treatments.

The science component includes a synthesis of and addition to fire

history work for these mountains. Public scoping for NEPAoccurred

in early 2011. The 2011 Horseshoe 2 Fire burned throughout the

Chiricahua range, but its effect on the project is reassessment rather

than postponement.

	 The 135,000-acre Galiuro FireScape covers remote country with

little WUI and concentrates on very large-scale burns. We are making a case on this landscape to use helicopter ignition for prescribed

burning in designated Wilderness. The CNF, BLM, TNC, UA, and

grazing permittees are project parties. Additional fire history studies

are incorporated into the science side of this project. Compliance is

underway, and a 40,000-acre three-party (BLM. TNC, CNF) burn is

in the works.

	 Cross-boundary,large-scalefiremanagement comeswithnumerous

institutional challenges.All-lands conservation is a directive to agenciesthat have few resourcesto devote to such work.Agency turnover

means passion ebbs and flows—loss of key playersslowsthe process

way down. FireScape competes with other agency programsforstaff

attention while wildfires get bigger and display increasingly severe

effects. Funding for planning and implementation is not likely to

increase soon. The FireScape teams carry on, knowing that the work

is needed, and that creative, broad-based efforts ultimately succeed.

The website www.azfirescape.org is a work in progress that displays

project information, including maps, for FireScape.

References

Kaib,J. 1998. Fire history in riparian canyon pine-oak forests and the intervening desert grasslands of the Southwest borderlands: A dendroecological,

historical, and cultural inquiry. Tucson, AZ: The University of Arizona.

234 p. Thesis.

National Interagency Coordination Center (NICC) 2011. 2011 Fire Season Summary. Online: http://www.predictiveservices.nifc.gov/intelligence/2011_Mid-Season_Report_Oct.pdf.

Swetnam, T. W. and C. H. Baisan. 1996. Fire Histories of Montane Forests

in the Madrean Borderlands. P. 15-36 in Ffolliott, P.F., DeBano, L. F.,

Baker, M.B., [and others], tech. cords. Effects of fire on Madrean Province

ecosystems—a symposium proceedings; March 11-15, 1996; Tucson AZ.

Gene. Tech. Rep. RM-GTR-289. Fort Collins, CO: U.S. Department of

Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 227 p.

Wadleigh, L. 2011. Wallow Fire 2011 Large Scale Event Recovery Rapid

Assessment Team Fire/Fuels Report. U.S. Department of Agriculture,

Forest Service, Apache-Sitgreaves National Forests. http://www.fs.usda.

gov/Internet/FSE_DOCUMENTS/stelprdb5333354.pdf.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
474 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Floristic Analysis of Heterogeneous Landscape Patches in 

a Biological Corridor in the El Rodeo-Básora Area near 

Moctezuma, Sonora, Mexico 

Ana Lilia Hernández-Rodríguez, María de la Paz Montañez-Armenta, Gertrudis Yanes-Arvayo, 

and Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora

Abstract—A floristic study was conducted in a heterogeneous landscape near Moctezuma, Sonora. From 

August to December 2011, Ranchos El Rodeo and El Básora were visited three times. The vegetation at about 

900 m elevation is foothills thornscrub, oak woodland, and induced buffelgrass grassland. A total of 120 plant 

taxa in 95 genera and 41 families were documented. Taxa were identified using regional literature, consulting 

with experts, and comparison with specimens in the Universidad de Sonora Herbarium. The families with 

more taxa are Asteraceae, Convolvulaceae, Fabaceae, and Solanaceae. The study area is a transitional zone 

between thornscrub oak woodland, making it an important biological corridor in central Sonora. Loss of 

flora is due to land use practices, especially clearing to introduce grass like buffelgrass (Pennisetum ciliare). 

Non-native plants could displace native taxa like Zinnia zinnioides, Ipomoea cristulata, and I. hederacea.

Introduction

	 The floristic richness of east-central Sonora is due to its location 

in the transition between the New World tropics and the northern

temperate zone at about 29°N, which enhances the diversity of

flora, fauna, vegetation types, and habitats. Large geographic areas

with difficult access have not been explored botanically. This paper

documents the flora and characterizes plant community patches in

a heterogeneous landscape in this area (Wiens 1995), which could

be used as biological corridors between foothills thornscrub, desert

grassland, and oak woodland. Our floristic inventory in Ranchos El

Rodeo-Básora area in the Municipio de Moctezuma is an important

contribution to the flora of Sonora.

Study Area

	 The Ranchos El Rodeo-Básora area is a corridor that connects the

mountains of central Sonora with the Sierra La Madera, a Sky Island

mountain range in the Madrean Archipelago. The study area is along

the Hermosillo-Moctezuma highway about 30 km (by air) southwest

of Moctezuma (between 29°36’39” to 29°39 ́06”N and 109°52 ́50”

to 109°56 ́18”W). The climate of the area is dry tropical with mean

maximumtemperatureof29.8oCduringJune andJuly,meanminimum

of 12 oC in December and January, and mean annual temperature is

of 21.1 oC. Frost only occurs occasionally, and hard freezes like that

of February 2-3, 2011, are very rare. Annual average rainfall is 460

mm(Instituto Nacional deEstadística y Geografía 2010). Subtropical

dry forest, grassland, and open low forest are the vegetation types

in the area (Instituto Nacional de Estadística y Geografía 2010).

Subtropical dry forest corresponds to foothills thornscrub of Felger

and others (2001) and Sinaloan thornscrub of Brown (1994). Open

low forest is also called oak woodland or encinal. Trees and shrubs

such as Acacia constricta, A. occidentalis, Ceiba acuminata, Bursera 

spp., Havardia mexicana, Lysiloma spp., Ipomoea arborescens, and

Prosopis velutina characterize vegetation in the study area (fig. 1).

Methods

	 The study sites were selected by field trips to the area and using

GoogleEarth satellite image (Image©2010 GeoEye.©2010 Google.

©2010 INEGI). Selection criteria were the presence of well-differentiation vegetation patches in a landscape mosaic, and connectivity

between them. Six sampling sites were visited inAugust, September,

and October 2011. A floristic list was constructed using the relevé

method (Múller-Dumbois and Ellenberg 1974). Plants were collected

for identification using regional floristic literature, consultation with

experts, and comparison with specimens in the University of Sonora

Herbarium (USON). Voucherspecimens were deposited into USON.

Collections, observations, and images from the El Rodeo-Básora

flora are available online in the Madrean Archipelago Biodiversity

(MABA) database (Madrean.org).

Results and Discussion

	 A total of 120 taxa in 95 genera, and 41 families were documented in the El Rodeo-Básora area. The families with more taxa

are Asteraceae (22), Fabaceae (16), Convolvulaceae (9), Cactaceae

(7), and Solanaceae (5) (fig. 2). Asteraceae, Fabaceae, and Poaceae
USDA Forest Service Proceedings RMRS-P-67. 2013 475

Floristic Analysis of Heterogeneous Landscape Patches in a Biological Corridor . . . Hernández-Rodríguez, and others

Figure 1—Study area and vegetation.

are the most speciose families in floras in Sonora (Van Devender

and others 2010). Life forms of the taxa were classified according to

Whittaker (1975) and (Krebs 1972): (a) trees—woody plants greater

than threemtall;(b)shrubs—short,woody plantlessthan threemtall;

(c) herbaceous—not woody plant above ground; (d) vines—woody

vines or herbaceous vines; (e) epiphyte—plants that grow on other

plants, but are not parasitic. Herbs with 62 taxa are the most common

life form (fig. 3). The physical structure of the vegetation is defined

by trees, shrubs, and large succulents (especially organpipe cactus

[Stenocereus thurberi]). Trees with 17 taxa are much more diverse

than in desert grassland, but only represent 14.2% of the flora. Some

common plants in the transition between foothills thornscrub and

oak woodland include Ceiba acuminata, Fouquieria macdougalli, F. 

splendens, Ipomoea arborescens, Lysiloma divaricatum, Parkinsonia 

praecox, Prosopis velutina, Quercus chihuahuensis, Q. viminea, and

Sabal uresana. Floristic richnessisthreatened by cattle ranching and

the introduction of exotic forage plants. 

Four species intheElRodeo-Básoraflorahave federalprotection 

status in Mexico according to the NOM-059-SEMARNAT-2010

(DOF 2010). Agave angustifolia (Agavaceae) has a status of Amenazada (= Threatened), while Amoreuxia palmatifida (Bixaceae), 

Crusea hispida (Rubiaceae), and Sabal uresana (Arecaeae) have

special status.

Figure 2—Dominant families in the El Rodeo-Básora area.
476 USDA Forest Service Proceedings RMRS-P-67. 2013

Hernández-Rodríguez, and others Floristic Analysis of Heterogeneous Landscape Patches in a Biological Corridor . . . 

We conclude that the El Rodeo-Básora area is of special interest and

conservation concern because of its mosaic of foothills thornscrub,

desert grassland, and oak woodland, its floristic diversity, its role as

a biological corridor connecting central Sonora landscapesto the Sky

Island region, and the presence of four federally protected species.

Acknowledgments

	 We thank J.Jesús Sánchez-Escalante at the Universidad de Sonora

for help identifying plants and putting voucherspecimensintoUSON.

Figure 3—Dominant life forms of taxa in the El Rodeo-Básora area.

We thank Thomas R. Van Devender and Ana Lilia Reina-Guerrero

for help in identifying plants and a careful review of the manuscript.

References

Diario Oficial de la Federación. 30 de octubre de 2010. Page 28. (Segunda

Sección)1. NORMA Oficial Mexicana NOM-059-SEMARNAT-2010,

Protecciónambiental-EspeciesnativasdeMéxicodeflorayfauna silvestresCategorías de riesgo y especificaciones para su inclusión, exclusión o

cambio-Lista de especies en riesgo.

Brown, David E. 1994. Biotic communities of the southwestern United States

and northwestern Mexico. Salt Lake City: University of Utah. 342 p.

Felger, Richard; Johnson, Matthew B.; Wilson, Michael F. 2001. The trees of

Sonora, Mexico. New York: Oxford University Press. 391 p.

Instituto Nacional de Estadística y Geografía. Consultado el 26 de octubre de

2011. En http: www.inegi.gob.mx 2010.

Krebs,J.Charles. 1972. Ecologìa. Estudio de la Distribución y laAbundancia.

2nd. Edición. México: Editorial OXFORD. pp. 452-455.

Mueller–Dombois, D.; Ellenberg, H. 1974. Aims and methods of vegetation

ecology. New York: John Wiley. 547 p.

Van Devender, Thomas. R.; Felger, Richard F.; Fishbein, Mark; [and others]. 2010. Biodiversidad de las plantas vasculares. Pp. 229-262 in

Molina-Freaner, Francisco; and Van Devender, Thomas R. eds. Diversidad Biológica de Sonora, Hermosillo: Universidad Nacional Autónoma

de México. http://www.skyislandalliance.org/Sonoran%20vascular%20

plants-VanDevender%20et%20al.2010.pdf

Wiens, John A. 1994. Habitat fragmentation: Island v landscape perspectives

on bird conservation. Ibis. 37: 97-104.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 477

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Association Between Nurse Plants and Saguaros 

(Carnegiea gigantea) in the Western Sonora

C. Hinojo-Hinojo, C. Trujillo-López, O. Calva-Pérez, O. Galaz-García, A. E. Castellanos-Villegas

Departamento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Hermosillo, Mexico

Abstract—The objectives in this work were to determine nurse species associated with the saguaro, as well 

as the diversity of plant species in the saguaro patch. We also recorded the presence of termites on saguaro 

green tissues to evaluate their existence and possible causes for their presence. At each site a point-centered 

quarter transect was used, where the data on the closet saguaros were taken for size and height, percent litter 

soil coverage, and termite activity, as well as floristic composition, plant species cover, and the dimension 

of the associated patch of vegetation. Data on 55 saguaros showed the most common nurse species were 

Larrea tridentata, Olneya tesota, Parkinsonia microphylla, and Ambrosia deltoidea. Twenty-one (21) saguaro 

individuals were found distributed in different patches where Olneya tesota and Parkinsonia microphylla 

showed the most species diversity. Termite activity was present in saguaros with little or no litter cover in the 

soil. This is the first work focusing on the causes for the increase of termite invasion of live saguaro tissues.

Introduction

	 Cacti are a diverse family and include about 2,000 species, which

are often endemic and unique elements ofthe arid and semi-arid areas

ofAmerica (Drezner and Lazarus 2008; Godinez-Alvarez and others

2003;Hunt andothers2006).Formost cacti,facilitationisundoubtedly

one of the ecological biotic interactions most widely documented.

Different species of perennial plants tend to favor germination of

seeds and survival of seedlings of various species of cacti, leading

to patterns of patches of strongly clumped species. A well-studied

example is the interaction of saguaro (Carnegiea gigantea) with

perennial trees and shrubs, where they facilitate the establishment of

seedlings of saguaro. However, when the saguaro is an adult it can

affect and compete with the nurse plant. Within the Sonoran Desert

in Sonora, some of the most important nurse plants have been used

for fuel, fencing, and crafts for a long time (Castellanos and others

2010a), despite that not much is known and many ecological aspects

of this interaction still need to be studied. For example, Castellanos

and others (2010b) have documented the interaction of termites, affecting green tissues of living saguaros, and propose this may be an

important cause in the death of saguaros and possible affecting its

population dynamics. Forsuch reasons, our research questions were:

1.Whatistheimportanceofnurseplantstothepopulationsofsaguaros?

2. Do patchiness diminish adverse biotic interactions? and

3. Arethereotherindirecteffectssuchasamountoflitterunderthenurse

plant that contributes to the saguaro not being invaded by termites?

Materials and Methods 

	 Sampled sites were located in the northwestern region of the

State of Sonora, near Puerto Libertad. We selected three sites that

had been previously sampled (Castellanos and others 2010b), in

which populations of saguaro are well represented. The selection of

saguaroswas alongtransects usingthe point-centeredquartermethod,

with a separation of 50 meters between each of them. At each point

we sampled the nearest saguaro in each of the four quadrants. This

method was used in each of the three sampling sites. After that, we

measured saguaro height, coverage ofthe patch, floristic composition

under the canopy of nurse species, and the amount of litter cover in

the soil close to the basal area around the saguaros. To determine the

age of saguaro, we used the formula proposed by Drezner (2002) in

which saguaro growth relates to their age. For the application of the

formula we required the calculation of a saguaro growth factor, which

differs according to the area to be analyzed and which consists in

measuring the population of saguaro during one year. We assumed a

factor of 0.85, that seems to fit most other studies, in order to reduce

the error when applying the formula.

Results and Discussion

Characterization of Saguaro (C. gigantea) 

Patches

	 We found a total of 21 species within different patches associated

with saguaro. The main nurse plants associated to saguaro in our

study region were Olneya tesota, Parkinsonia microphylla, Ambrosia 

deltoidea and Larrea tridentata. The last plant species allowed the

establishment of 11 other species, but presented a large variability

amongallpatches.TherewerepatcheswhereL. tridentata wastheonly

species associated with saguaro, and others where it was associated

with two to three different species. Patches with the largest diversity

of species were under P. microphylla and O. tesota, and were related

to the larger canopy cover from those patches.
478 USDA Forest Service Proceedings RMRS-P-67. 2013

Hinojo-Hinojo, Trujillo-López, Calva-Pérez, and others Association Between Nurse Plants and Saguaros . . .

Description of the Saguaro (C. gigantea) and 

Its Nurse Plants Populations.

	 The three study sites showed a similar population density with

an average of 58.8 plants per hectare. Our data indicated that the

number of saguaros sampled per site was sufficient to estimate the

population density. The saguaro population structure had a progressive pyramid-like shape. This indicates a young saguaro population

with a constant growth, and it is noteworthy that the most abundant

saguaro size range wasfor plants between 0 and 1.5 metersin height,

which meansthat the saguaro populations are dominated by juveniles

in the three study sites.

	 The main plant species found as nurse plants of saguaro were Larrea tridentata, Ambrosia deltoidea, Olneya tesota and Parkinsonia 

microphylla, in decreasing importance. In addition we calculated the

occurrence of facilitation by these fourspecies and found that Larrea 

tridentata had the highest percentage (30.4%) on the three sites.

Percentage Litter Cover in Soil and Calculated 

Age of Saguaros (C. Gigantea) Studied

	 Litter cover in soil surrounding the base of saguaro of different

ages changed or was not present, so there was no specific pattern or

influence of the age of the saguaros with respect to percentage soil

covered with litter even though that could have been influenced by

the nurse plant to which it is associated. However, we found that older

saguaros did not have any litter on the soil, perhaps because they do

not already have a nurse plant that could provide such litter material,

unlike the juvenile saguaros that had higher litter soil cover.

Absence or Presence of Termites at Saguaro 

(C. gigantea) Sites

	 We evaluated the susceptibly to invasion of termites in relation

to the age of saguaros, and found that termite activity was higher

between the age of 30 to 80 years. In the limits of the population

of saguaros (13-118 years) did not show invasion by termites. This

may be because young saguaros are sheltered and protected under

the canopy of the nurse plants.

	 Based on the average from four measurements of percent soil

litter cover, we found that individuals with low percent soil litter

cover (0 to 30%) were more susceptible to invasion in the tissues of

the saguaro by termites. It was further noticed that there was a 1:1

ratio for individuals with low percent soil litter cover, indicating that

termites do not always tend to invade the saguaro that does not have

a noticeable percentage of soil litter cover. Also, we found that the

main nurse plants species (Olneya tesota, Parkinsonia microphylla, 

Ambrosia deltoidea and Larrea tridentata) did not present difference

on the organic matter despite the form of life (tree or shrub) and its

canopy coverage.

Conclusions

	 The main nurse plants for saguaro in our study area were Olneya 

tesota, Parkinsonia microphylla, Ambrosia deltoidea and Larrea 

tridentata, which differ in their role within the studied populations.

Their nurse contribution depends mainly on the size of the patch

and therefore, the diversity of species associated with nurse plants.

We found 21 species primarily associated with the evergreen plants

Olneya tesota andParkinsonia microphylla includingsaguaro.Larrea 

tridentata was a common nurse plant species for saguaro in the three

sites.The ecological and ecophysiologicalrelations of a nurse species

to the saguaro population dynamics are a focus of more studies to

understand the many details about these interactionsthat occurin arid

and semiarid lands.The percentage litter coverin the soil provided by

the nurse plant, and the species under it, seem to work as a defense

mechanism for saguaros against herbivore attack by termites. It was

found that termites do not invade saguaros with larger litter cover

of soil at their base, meaning that the absence of litter is related to

the invasion of termites, although other mechanisms may also be

responsible for termite invasion to living green tissues of saguaro.

References

Búrquez-Montijo,J.A.2009.Distribución, estructurapoblacionalyutilización

del saguaro (Carnegia gigantea (Engelm.) Britt & Rose) en México. Universidad Nacional Autónoma de México. Instituto de Ecología. Informe

final SNIB-CONABIO proyecto No. ES008. México D.F.

Castellanos, A. E., Castillo, R. A., Quijada, A. 2010. Termite activity on

green tissues of saguaro (Carnegiea gigantea) in the Sonoran Desert.

In: Halvorson, W., C. Schwalbe and C. van Riper (eds.) Southwestern

Desert Resources. University of Arizona Press, Tucson, AZ. p. 171 - 180.

Castellanos-Villegas, A. E., L. C. Bravo, G. W. Koch, [and others] 2010.

Impactos Ecológicos por el Uso del Terreno en el Funcionamiento de

Ecosistemas Áridos y Semi-Áridos de Sonora. En: F. E. Molina-Freaner y

T. Van Devender, eds. Diversidad Biológica del Estado de Sonora. UNAM

- CONABIO, México, D. F. pp 157-186.

Drezner, T. D. and B. L. Lazarus. 2008. The population dynamics of columnar

and other cacti: a review. Geography Compass. 2:1-29.

Drezner, T. D. 2002. Saguaro (Carnegiea gigantea, Cactaceae) age-height

relationships and growth: the development of general growth curve.

American Journal of Botany. 90:911-914.

Godínez-Álvarez, H., Valverde, T. & P. Ortega-Baes. 2003. Demographic

trends in the Cactaceae. Botanical Review. 69: 173-203.

Hunt, D. R, Taylor N. P. and G. Charles. 2006. The new cactus lexicon. VolumesI. InternationalCactaceae Systematics Group, DH Books, Milbourne

Port. p. 373

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 479

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Vegetation Monitoring on Semi-Arid Grasslands Ungrazed 

by Domestic Livestock

Linda Kennedy

National Audubon Society, Appleton-Whittell Research Ranch, Elgin, Arizona

Dan Robinett

Robinett Rangeland Resources LLC, Elgin, Arizona

Abstract—The Research Ranch is an 8000-acre sanctuary and research facility in the semi-arid grasslands of 

southeastern Arizona, USA. Cattle were removed from the property in 1968 to provide a reference area by 

which various land uses, such as grazing and exurbanization, could be evaluated. Vegetation transects were 

established in 2000 and 2003 on several ecological sites in Major Land Resource Area 41.This monitoring 

program has tracked changes after wildfires and during drought. Trends associated with non-native, invasive 

species, particularly Eragrostis lehmanniana (Lehmann lovegrass) and E. curvula (Boer lovegrass) have been 

documented. For example, in an 8-year span, the frequency of Lehmann lovegrass on one loamy upland site 

grew from 1% to 60% at the expense of the native Eragrostis intermedia (Plains lovegrass) and Bouteloua 

gracilis (Blue grama). Results from monitoring provides feedback to Audubon personnel in the management 

of the property and are shared with other land managers to help tease apart the effects of grazing and other 

land use actions from the effects of climate.

Introduction

The Appleton-Whittell Research Ranch is an 8000-acre exclosure 

from which domestic livestock were removed in 1968. The facility 

is a cooperative effort among several landowners including the Bureau of Land Management, U.S. Forest Service, Resolution Copper 

Company, The Nature Conservancy, The Research Ranch Foundation, and National Audubon Society, and is managed by Audubon 

under contractual agreements with each. The Research Ranch is 

primarily Madrean mixed grass prairie. Its lands are tributary to the 

Babacomari River. Audubon staff work closely with the members of 

the cattle ranching community and ranchers are able to compare their 

management efforts with an ungrazed reference area. Many research 

projects have resulted from work conducted on the Research Ranch 

but vegetation monitoring has been a relatively recent addition to the 

scope of activities.

Methods

Eighteen vegetation transects were established in 2003 and 2004 

based on the work by Breckenfeld and Robinett (2001). Transects 

were located in representative areas within Ecological Sites. The 

Plant Frequency method (Despain and others, 1997) is used to gather 

vegetation data. Four parallel lines of 50 frames each yield 200 frames 

to determine frequency. Three points per frame are used to establish 

cover (data not shown). Simple rain gages are located at or near each 

transect and read twice per year.

Results

Data from a subset of four transects established in loamy uplands 

are summarized along with precipitation data (fig. 1) and some generalizations across all transects. 

Transect 650

This transect is on the western side of the Research Ranch and 

exhibits a 15% SW slope. The Ryan Fire burned this area in April 

Figure 1—The cumulative shortfall (long-term mean—actual precipitation) 

during the 9 years represented by this report equals 24 inches. 
480 USDA Forest Service Proceedings RMRS-P-67. 2013

Kennedy and Robinett Vegetation Monitoring on Semi-Arid Grasslands Ungrazed by Domestic Livestock

2002 during severe drought. Recent precipitation: Monsoon 2010:

14.7 inches; Winter 10/11: 0.8 inches; Monsoon 2011: 6.7 inches.

Frequencyofsideoatsgrama (Bouteloua curtipendula)hasbeenstable

or increased (fig. 2), but blue grama (B. gracilis) crashed in 2006 and

has not recovered. Wolftail (Lycurus setosus) is increasing and curly

mesquite (Hilaria belangeri) is maintaining despite the dry winter

of 2010/11.

Transect 651

	 This site is on the western side of the Research Ranch and has a

2% E slope. This transect also burned in the Ryan Fire of April 2002.

Recent precipitation: Monsoon 2010: 12.5 inches; Winter 10/11: 1.5

inches; Monsoon 2011: 7.1 inches. Lehmann lovegrass (Eragrostis 

lehmanniana) is exhibiting its invasive potential on this transect

(fig. 3), increasing from 1% frequency in 2003 to 55% in 2006,

and is retaining dominance. Both blue grama and plains lovegrass

(E. intermedia) show declines during this period. Wolftail is increasing on this transect.

Figure 2—Loamy upland transect 650.

Figure 3—Loamy upland transect 651.

Transect 706

	 This site is also on the eastern side of the research ranch. Slope is

12-15% E. This plot burned in the Ryan Fire of April 2002. Recent

precipitation was: Monsoon 2010: 11.7 inches; Winter 10/11: 0.9

inches; Monsoon 2011: 10.6 inches. NRCS had established and read

this transect in 1997 using the same protocol, so we were able to

compare changes and trendsfor a longer time period. Curly mesquite

was dominant in 1997 (96%), butfound in only 2 frames(1%)in 2011

(fig. 4). Frequency of cane beard grass (Bothriochloa barbinodis)

is trending upward, 18% to 52%. Lehmann lovegrass was found at

low frequencies (2-3%) from 1997 through 2005 before peaking at

41% in 2009. Complete plant mortality for this species brought the

frequency down to 10% in 2011. In 1997, plains lovegrass was noted

in 26% of the frames. By 2011 none was recorded.

Transect 665

	 Thistransecthas a slight(1-3%)northernexposure.Thisplotburned

in the Ryan Fire ofApril 2002 and the Canelo Fire of May 2009 (also
USDA Forest Service Proceedings RMRS-P-67. 2013 481

Vegetation Monitoring on Semi-Arid Grasslands Ungrazed by Domestic Livestock Kennedy and Robinett

Figure 4—Loamy upland transect 706.

a drought year). Recent precipitation: Monsoon 2010: 13.0 inches;

Winter 10/11: 1.0 inches; Monsoon 2011: 7.7 inches. Plainslovegrass

has exhibited a steady decline during this period,from39% frequency

in 2003 to 0% in 2011 (fig. 5). Lehmann lovegrass was found at low

levels until 2006 (62%). Curly mesquite declined from 16% to 2%.

Conclusions

	 We established/re-established sixteen transects in 2003 and added

two more in 2004. The transects experienced a range of fire activity

and all have been influenced by drought conditions. This is a monitoring project rather than a research project, so care must be taken to

assign causality, but two iconic native grasses, plains lovegrass and

blue grama have exhibited a steady decline in frequency and the trend

for the non-native, Lehmann lovegrass is upward. It was hoped that

Figure 5—Loamy upland transect 665.

grassland protected from grazing by livestock would be resistant to

invasion by non-native grasses, but this hope has proved groundless.

Other methods must be used ifthe objective isto protect and maintain

native plant species diversity.

References

Breckenfeld, D.; Robinett, D. 2001. Soil and range resource inventory of 

the National Audubon Society Appleton-Whittell Research Ranch Santa 

Cruz County, Arizona. Online: http://researchranch.audubon.org/PDFs/

SoilRangeResourceInventory2001.pdf.

Despain, D.W.; Ogden, P.R.; Smith, E.L. 1997. Plant frequency sampling for 

monitoring rangelands. In: Some methods for monitoring rangelands. G.B. 

Ruyle, ed. Arizona Cooperative Extension Publication 190043.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
482 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Mountain Pine Beetle in Southwestern White Pine in the 

Pinaleño Mountains

Ann M. Lynch

USDA Forest Service, Rocky Mountain Research Station, Tucson, Arizona

Christopher D. O’Connor

School of Renewable Natural Resources, University of Arizona, Tucson, Arizona

Abstract—Mountain pine beetle has rarely been found in the Madrean Sky Island Archipelago and has not 

been reported from the Pinaleño Mountains until recently. This insect began killing southwestern white pine 

in 1996 or earlier, with additional mortality each year since. Activity has increased in the last 2 years. The 

life cycle in the Pinaleños during this time has been univoltine, with some females surviving the winter to 

produce a second brood in the spring. Stain fungi are evident in most killed trees, but not all. In infested 

areas, total stocking and the abundance of southwestern white pine has been significantly reduced, creating 

small openings in the stand canopies. The future course of this outbreak is uncertain.

Introduction

	 Mountainpinebeetle (Dendroctonus ponderosaeHopkins(Coleoptera: Curculionidae, Scolytinae)) is a native bark beetle infrequently

found in the Madrean Sky Island Archipelago, and there is no record

of it in the Pinaleño Mountains (32.40 latitude). Scattered mountain

pine beetle-infested southwestern white pines (Pinus strobiformis)

were found in 2007 near Soldier Creek in the Pinaleño Mountains

(USDAForest Service 2008).Additional individual and small groups

of southwestern white pines have been infested each year since.

Mortality each year is minor, but the outbreak has persisted, damage

appearsto be increasing, and mortality has accumulated to significant

levels that affect resource values. Values at risk include recreation,

as some of the infestations are in the vicinity of developed recreation

sites, and wildlife, as the seeds, large trees, and snags are utilized by

the endangeredMountGrahamredsquirrel,Tamiasciurus hudsonicus 

grahamensis(Benkmanandothers1984;FrankandCox2009), among

others. Though mountain pine beetle is the most significant insect

disturbance agent in lodgepole pine (Pinus contorta) forests, and is

well studied elsewhere, little is known about its biology and effects

in southern Arizona or on southwestern white pine. We undertook

investigations to characterize the effects of infestation, determine

tree and site characteristics associated with mountain pine beetle

infestation, and establish a timeline of outbreak development

Background

	 Though well distributed throughout much of western North

America (Gibson and others 2009; Safranyik and others 2010; Wood

1982), records of mountain pine beetle and related tree mortality are

sparse in southern Arizona. Hopkins (1909) reported finding it in the

Chiricahua Mountains in 1907, but reported no further information

from that location. A tabulated Forest Service record of forest insect

activity during the 1960s inArizona and New Mexico (USDAForest

Service 1971) indicates that an infestation occurred on the Coronado

National Forest (N.F.) in the 1960s, at the same time that outbreaks

occurred on the Cibola N.F. and Carson N.F. in New Mexico. Though

survey recordsfrom individual yearsin the 1960s are no longer available, the information indicates that infested trees were on these three

National Forests and included ponderosa pine (Pinus ponderosa) and

southwesternwhitepine in1961-1965,1968, and1969.Theonlyother

insect activity recorded for the Coronado N.F. during that time period

was roundheaded pine beetle (Dendroctonus adjunctus Blandford)

in 1967-1970. No further reference to mountain pine beetle activity

in southern Arizona was found until 2007.

	 The historicalrange ofmountain pine beetle extendsfromnorthern

Baja, California, and the northern border of Mexico (310 latitude)

to central British Columbia (560) (Cibrián-Tavar and others 1995;

Safranyik and others 2010; Wood 1982). As a result of rising temperatures, the geographic range has recently expanded northward to

eastern Alberta (Safranyik and others 2010) and to higher elevations

(Bentz and others 2011; Logan and Powell 2009). Activity within

the historical range has also increased in recent years, and outbreaks

are occurring in areas where they either were not recorded or were

recorded infrequently (Bentz and others 2011; Bentz and SchenLangenheim 2007; Gibson and others 2009). The current outbreak

in British Columbia and Alberta is likely one of the largest in history

(Kurz and others 2008; Taylor and others 2006). All Pinus species

within the mountain pine beetle range except Jeffrey pine (Pinus jeffreyi) can be successfully attacked, including white pines and piñons.
USDA Forest Service Proceedings RMRS-P-67. 2013 483

Mountain Pine Beetle in Southwestern White Pine in the Pinaleño Mountains Lynch and O'Connor

	 Temperature regimes play a significant role in determining when

and where mountain pine beetle outbreaks occur (Gibson and others

2009; Logan and others 2010), and rising temperatures may have

contributed to recent changes (Bentz and others 2011; Safranyik and

others 2010) by influencing seasonal aspects of life history (Logan

and Bentz 1999). In order to overcome tree defenses, adult mountain

pine beetles must mass-attack trees with large numbers of beetles.

Synchronized adult emergence and aggregation pheromones make

it possible for sufficient numbers of beetles to be present at one

time to overcome tree defenses. Mountain pine beetle can adapt its

development ratesseasonally to optimize likelihood ofsynchronized

emergence (Logan and Bentz 1999). Throughout its wide geographic

range, mountain pine beetle has adapted to local temperature regimes

andhasthe capacitytoexhibitdifferentseasonallifehistories(Amman

andBartos 1991; Gibson and others 2009).Mountain pine beetle does

not have a winter diapause, and the life history is very responsive

to temperature. Mountain pine beetle is typically univoltine, but in

portions of its range with cooler summers (especially at high elevations), it is semivoltine (Safranyik and Carroll 2006). Outbreaks occur during warm periods, particularly when winter temperatures are

mild enough to allow survival of all life stages, and there issufficient

summer thermal energy to allow a univoltine life cycle (Logan and

others 2010) and when temperature regimes facilitate synchronous

adult emergence (Bentz andothers 1991;Safranyik andCarroll 2006);

suitable thermal windows may be quite narrow. The northern part of

the Madrean archipelago is the southern extreme for the mountain

pine beetle range, and the thermal habitat is probably marginal for

the species. Most of the time, activity is probably minimal because

synchronization of adult emergence is insufficient to allow massattack, a univoltine life cycle isinfrequent, or mortality occursto one

or more life stages because summer temperatures are too warm or

because an overwintering stage is susceptible to cold. One or more

of these conditions has probably changed recently.

Approach

	 Our objectives were to characterize the effects of infestation, determine tree and site characteristics associated with mountain pine

beetle infestation, and establish a timeline of outbreak development.

Our approach was to establish plots in infested areas, compare the

character ofthose plotsto randomly located southwestern white pineinhabiting plots, and date mortality of mountain pine beetle-attacked

trees using tree-ring analysis.

	 Infested areas were located by reconnaissance both on foot and by

vehicle. Plots were located completely within infested areas. Where

infested areas were large, plots were located 100 m apart, and along

an elevation gradient where possible. Trees larger than 20 cm DBH

were measured on a 0.05-ha plot fixed-radius plot while treessmaller

than 20 cm DBH but 3 cm DBH or larger were measured on a 0.02-ha

fixed-radius plot. At each plot, spatial coordinates, elevation, aspect,

slope, and species oftreesin the vicinitywere recorded.Data recorded

for each tree included species, diameter (DBH, or DIB if bark was

absent), dwarf mistletoe rating, canopy position, needle color and

abundance on live and dead trees, presence of bark and limbs of

various sizes on dead trees, evidence of bark beetle attack, species

of attacking bark beetle, and evidence ofstain fungi. When recording

canopy position we distinguished between small suppressed trees

and healthy understory trees. There are many bark beetle speciesthat

attack Pinus in southern Arizona, but mountain pine beetle galleries

are distinctive and easily distinguished from those of other species

(fig. 1a; Fairweather and others 2006). We had little difficulty finding evidence of mountain pine beetle activity in trees that had been

dead for 15 years (fig. 1b), as mountain pine beetle scores the wood

surface and these scars are evident after much weathering. Increment

cores were taken from mountain pine beetle-killed treesthat had been

dead long enough that the attack year could not be determined from

visual inspection of snag condition, as well as from some recentlykilled trees. The outer ring dates of these cores were determined in

the laboratory using standard tree-ring procedures(Stokes andSmiley

1968). We also opportunistically sampled individual mountain pine

beetle-killed treesfound during reconnaissance to assist in determining the outbreak timeline.

	 Characteristicsofmountainpinebeetle-infestedtrees andplotswere

compared to tree and plot characteristics of systematically located

plots in a Pinaleño Demography study (Falk and others 2009) on

whichsouthwesternwhitepinewasfound.ThePinaleñoDemography

plotsystem consists of two sampling grids acrossthe mountain range

above 2134 m elevation. The first grid is spaced at 1000 m, and the

second grid is offset east and south by 500 m. Plots consist of nested

0.017- and 0.05-ha plots for small and large trees, respectively. At

each plot, site and stand characteristics data were collected in 2009

and 2010 similar to what is described above for live and dead trees

of all species. Additionally, increment cores were collected from

each tree and cross dated and measured to establish demographics,

mortality events, growth release events, and recruitment events.

	 We did not set out to investigate the life cycle and seasonal history

of mountain pine beetle in the Pinaleño Mountains. However, during

the last several years the first author has repeatedly visited infested

trees and plots several times to monitor the situation, assist other

investigators in the collection of genetic material and infested bolts

for life history studies, train local managers in identifying mountain

pine beetle, and to develop this study. These observations are also

reported here.

	 Mountain pine beetle-infested areas were located below the Riggs

Lake campground, in and above the Treasure Park campground, on

the upperslopes ofWebb Peak, and in the SoldierCreek campground.

District personnel had removed many dead and infested trees at the

latterlocation forsafety reasons, and we sampled six plotsin the other

locations. Here we report the resultsfrom those plots, and are careful

Figure 1—(a) Fresh mountain pine beetle galleries, and (b) gallery scars 

from a tree killed in 1997 and photographed in 2011; both are southwestern white pine in the Pinaleño Mountains. Mountain pine beetle 

females construct long nearly-linear galleries with J- or L- shaped hooks 

at the bottom, along which eggs are laid; larvae feed in the phloem and 

construct galleries that initially are horizontal and do not score the wood 

(photos by A.M. Lynch).
484 USDA Forest Service Proceedings RMRS-P-67. 2013

Lynch and O'Connor Mountain Pine Beetle in Southwestern White Pine in the Pinaleño Mountains

to not over interpret the limited data, restricting our observations to

what we think can be supported by the data. In 2012, we will sample

additional plots and determine the causal agents of dead southwestern

white pine found in the Pinaleño Demography study.

Results

Insect Biology

	 The majority of the mountain pine beetle population was clearly

univoltine with mass-attacks in the autumn. A small number of trees

were successfullymass-attackedinspring.Thiswas evidencedbyvery

tight bark in trees with excessive sap flow on the outer bark in spring,

presence of pupae in early June 2011 (indicative of spring attacks

the previous year [B.J. Bentz, personal communication]), noticeable

fading offoliage in some treesin autumn (fading fromautumn attacks

usually occurs early inthe subsequent growing season), andformation

of a partial growth ring in the outer ring of a few dead trees. Our data

is not sufficient to accurately determine the proportion of trees that

were attacked in spring (it was low, perhaps 10-15%) or if a portion

of the population was semivoltine.

	 Evidence of stain fungi was found in most trees, but not all. We

did not record the presence of staining at the beginning of the study.

Twenty-five percent of sampled southwestern white pine and ponderosa pine (on the plots or not) and 12% of the cored trees did not

have stain fungi. Stain-free trees were killed in a range of years from

1998 to 2011.

Outbreak Development

	 We originally thought that the outbreak originated at the summit

of Webb Peak in lightning-struck trees, but this was not the case.

Individual dead trees with outer ring dates of 1996 and 1997, which

for autumn-attacked trees would be the year-of-attack, were found

scattered acrossthe landscape. Larger numbers oftrees were attacked

in 2001 and especially 2010.Thisistypical ofmountain pine beetle—

randomly distributed impaired trees are attacked when population

densities are low, but once population density increases beetlesselect

clustered populations of larger resource-rich trees (Boone and others

2011).

Character of Infested Trees

	 Fifty-fourmountainpinebeetle-attackedtreeswerefound,including

14 opportunistically sampled (off-plot) trees. All were southwestern

white pine except for two ponderosa pine and one Douglas-fir (Pseudotsugae menziesii).TheDouglas-firprobablyrepresents amistakeby

attacking beetles, as it was growing tightly intertwined with a similarly sized southwestern white pine (both were 13 cm DBH). Brood

developed in the southwestern white pine, but not in the Douglas-fir.

All attacked trees were larger than 17 cm DBH except for these two

trees, and the majority were larger than 20 cm.

	 On infested plots, a greater proportion of larger size classes were

attacked (fig. 2). Over 70% of trees 40 cm or larger in diameter were

killed by mountain pine beetle.

Character of Infested Plots

	 The Pinaleño Demography plot system (Falk and others 2009)

represents a random sample ofsitesin the Pinaleño Mountains above

2134 m. On plots with southwestern white pine, it is most commonly

associatedwithDouglas-fir,whitefir(Abies concolor), andponderosa

pine (89,51, and49%ofplots,respectively).Engelmannspruce (Picea 

engelmannii), corkbark fir (A. lasiocarpa var. arizonica), and aspen

were less frequently found (31, 31, and 26% of plots, respectively),

and Gambel oak (Quercus gambelii) and Arizona willow (Salix arizonica) were found occasionally. Southwestern white pine inhabited

sites with slopes less than 65% with a wide range of aspect, though

few faced west.

	 Species composition, slope, and aspect of mountain pine beetleinfested plots were very similar to those of the Pinaleño Demography

plots. Elevation ranged from 2670 to 3050 m. Southwestern white

pine stocking of infested plots varied considerably, between 5 and

36 sq m/ha, and stocking of all species varied between 34 and 73 sq

m/ha. Douglas-fir was found on all infested plots

Figure 2 –Proportion of southwestern white pine infested with or killed by mountain pine beetle on infested 

plots. Each size class, 15-50 cm, had 6 or more trees.
USDA Forest Service Proceedings RMRS-P-67. 2013 485

Mountain Pine Beetle in Southwestern White Pine in the Pinaleño Mountains Lynch and O'Connor

	 On average, infested plots had similar densities but half the basal

area compared to the random Pinaleño Demography plots. There

was a higher representation of southwestern white pine that were, on

average, smaller in diameter than those in the Pinaleño Demography

study (table 1).

Effects of Infestation

	 Mountain pine beetle infestation significantly reduced total stocking and density, and the abundance ofsouthwestern white pine (table

1). Mean basal area was reduced approximately 60%, creating small

openings in the canopy. Resulting species composition favored

Douglas-fir, with mean dominance by that species increasing from

22 to 56%.

Discussion

	 Some conclusions can be made from the study at this point. In the

Pinaleño Mountains, mountain pine beetle preferred areas dominated

by southwestern white pine indicates that insect attacks occur where

the host is more abundant. We found few ponderosa pine that were

attacked bymountain pine beetle, even in areas where ponderosa pine

was common.

	 The overall impacts ofthisinsect have been modest, creating small

openingsin mixed-coniferstands. Southwestern white pine is prolific

in the Pinaleño Mountains, with abundant reproduction. It is one of

the preferred species for several resource objectives, and has been a

stable component of the mixed-conifer forest. Though damage has

been modest to date, it increased significantly in 2010 and 2011.

	 Uncertaintyregardingthe future courseoftheoutbreakisthe largest

concern. Mountain pine beetle has aggressive population dynamics

once population densities are high enough that the defenses of healthy

trees can be overcome (Boone and others 2011; Gibson and others

2009), and once started, outbreaks can deplete the host resource over

large areas. The initial stages of outbreaks are marked by a change in

attack behavior from individual impaired trees to groups of healthy

trees, which appears to be the case in the Pinaleños. Attacks in the

last 2 years have clearly favored groups of healthy trees; however,

the scale of activity in the Pinaleños is rather small, so it is too soon

to conclude that a high-intensity outbreak has begun. Outbreaks

occur in warm periods when the population can sustain a univoltine

life cycle (Logan and others 2010), and that appears to be the situation in the Pinaleños. On the other hand, the temperature regime

needed to sustain that life cycle in southernArizona is unknown. The

thermal habitat for mountain pine beetle may be marginal, and it is

possible that small changes in the temperature regime could disrupt

the univoltine life cycle and synchronization of adult emergence,

which would subsequently dampen outbreak dynamics. Activity in

the1960s apparentlysubsidedwithout catastrophe.Toolittle isknown

about the course of that outbreak to assist in projecting mountain pine

beetle activity during the current outbreak. We cannot determine if

the mountain pine beetle population in the Pinaleñosis at an incipient

outbreak stage or in the initial stage of an intense outbreak.

Acknowledgments

	 We thank Matthew Littrell with the Coronado N.F. for assistance

with locating mountain pine beetle-infested areas, and José Negrón,

Brytten Steed, and Jaime Villa Castillo for their helpful comments

on earlier drafts.

References

Amman,G.D.;Bartos,D.L.1991.Mountainpinebeetleoffspringcharacteristics

associated with females producing first and second broods, male presence,

and egg gallery length. Environmental Entomology 20: 1562-1567.

Benkman, C.W.; Balda, R.P.; Smith, C.C. 1984. Adaptations for seed dispersal and the compromises due to seed predation in limber pine. Ecology

65(2): 632-642.

Bentz, B.J. 2006. Mountain pine beetle population sampling: inferencesfrom

Lindgren pheromone traps and tree emergence cages. Canadian Journal of

Forest Research 36: 351-360.

Bentz, B.; Campbell, E.; Gibson, K.; [and others]. 2011. Mountain pine beetle

in high-elevation five-needle white pine ecosystems. Pp. 78-84 in Keane,

R.E.;Tomback,DF.;Murray,M.P.;Smith,C.Y.(editors).The futureofhighelevation five-needle white pines in western North America: Proceedings

of the High five Symposium; 28-30 June 2010; Missoula, MT. Proceedings

RMRS-P-63. Fort Collins, CO: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station. 376 p.

Bentz,B.J.;Logan,J.A.;Amman,G.D.1991.Temperature-dependentdevelopment of the mountain pine beetle (Coleoptera: Scolytidae) and simulation

of its phenology. The Canadian Entomologist 133: 375-387.

Bentz, B.J.; Schen-Langenheim, G. 2007. The mountain pine beetle and

whitebark pine waltz: Has the music changed? Pp. 43-50 in Proceedings

of the conference Whitebark Pine: A Pacific coast perspective. R6-NRFHP-2007-01.PortlandOR:U.S.DepartmentofAgriculture,ForestService.

Boone, C.K.; Aukema, B.H.; Bohlman,n J.; [and others]. 2011. Efficacy of

tree defense physiology varies with bark beetle population density: A basis for positive feedback in eruptive species. Canadian Journal of Forest

Research 41: 1174-1188.

Cibrián-Tovar, D.; Méndez-Montiel, J.T.; Campos-Bolaños, R.; [and others].

1995. Insectos Forestales de Mexico. Universidad Autonoma Chapingo,

Secretaría de Agricultura y Recursos Hidraulicos, United States Department of Agriculture, Natural Resources Canada y Comision Forestal de

America del Norte. 455 p.

Table 1—Characteristics of study plots before and after mountain pine beetle infestation, and plots 

in the Pinaleño Demography study where southwestern white pine was present.

Before After Change (%) Demography

All species

stocking (sq m/ha) 49 ± 17 19 ± 5 -61 96 ± 39

density (tph) 660 ± 214 537 ± 193 -19 698 ± 335

Southwestern white pine

stocking (sq m/ha) 27 ± 9 14 ± 11 -48 20 ± 19

density (tph) 360 ± 135 240 ± 153 -33 152 ± 102

dominance (% stocking) 55 ± 9 37 ± 18 -32 22 ± 20

mean dbh (cm) 29.2 ± 7.5 28.0 ± 8.7 -4 39 ± 11
486 USDA Forest Service Proceedings RMRS-P-67. 2013

Lynch and O'Connor Mountain Pine Beetle in Southwestern White Pine in the Pinaleño Mountains

Fairweather, M.L.; McMillin,J.; Rogers, T.; [and others]. 2006. Field guide to

insects anddiseases ofArizona andNewMexico forests.Albuquerque,NM:

U.S.DepartmentofAgriculture,ForestService,SouthwesternRegion.271p.

Falk, D.A.; Lynch, A.M.; O’Connor, C.; [and others]. 2009. Growth and

demography of Pinaleño high elevation forests. Performance Report RJVA

07-JV-11221615317.U.S.DepartmentofAgriculture,ForestService,Rocky

Mountain Research Station. 25 p.

Frank, C.L.; Cox, S.R. 2009. The adaptive significance of seed hoarding by

the Mt. Graham red squirrel. Pp. 256-273 in Sanderson, H.R.; Koprowski,

J.L., (editors). The last refuge of the Mt. Graham Red Squirrel: Ecology of

endangerment. Tucson: The University of Arizona Press. 427 p.

Gibson, K.; Kegley, S.; Bentz, B. 2009. Mountain pine beetle. Forest Insect

& Disease Leaflet 2. FS-R6-RO-FIDL#2/002-2009. U.S. Department of

Agriculture, Forest Service, Pacific Northwest Region. 12 p.

Hopkins, A.D. 1909. Practical information on the Scolytid beetles of North

American forests. I. Barkbeetles of the genus Dendroctonus. Bulletin No.

83, Part I. Washington, DC: U.S. Department of Agriculture, Bureau of

Entomology.

Kurz, W.A.; Dymond, C.C.; Stinson, G.; [and others]. 2008. Mountain pine

beetle and forest carbon feedback to climate change. Nature 452: 987-990.

Logan, J.A.; Bentz, B.J. 1999. Model analysis of mountain pine beetle seasonality. Environmental Entomology 28: 924-934.

Logan, J.A.; Powell, J.A. 2009. Ecological consequences of forest insect

disturbance altered by climate change. Pp. 98-109 in Wagner F.H. (editor).

Climate warming in western North America: evidence and environmental

effects. Salt Lake City: University of Utah Press.

Logan, J.A.; MacFarlane, W.W.; Willcox, L. 2010. Whitebark pine vulnerability to climate-driven mountain pine beetle disturbance in the Greater

Yellowstone Ecosystem. Ecological Applications 20: 895-902.

Safranyik, L.; Carroll, A.L. 2006. The biology and epidemiology of the

mountain pine beetle in lodgepole pine forests. Pp. 33-66 in Safranyik

L., Wilson B. (editors). The mountain pine beetle: A synthesis of biology,

management and impactsin lodgepole pine. Victoria, BC: Canadian Forest

Service, Natural Resources Canada.

Safranyik,L.;Carroll,A.L.;Régnière,J.;[andothers].2010.Potentialforrange

expansion of mountain pine beetle in the boreal forest of North America.

The Canadian Entomologist 142: 415-442.

Six, D.L.; Adams, J. 2007. White pine blister rust severity and selection of

individual whitebark pine by the mountain pine beetle (Coleoptera: Curculionidae, Scolytinae). Journal of Entomological Science 47:345–353.

Stokes,M.A.;Smiley,T.L.1968.Anintroductiontotree-ringdating.University

of Chicago Press. 73 p.

Taylor, S.W.; Carroll, A.L.; Alfaro, R.I.; Safranyik, L. 2006. Forest, climate

and mountain pine beetle outbreak dynamicsin western Canada. Pp. 67-94

in Safranyik, L., Wilson, B. (editors). The mountain pine beetle: A synthesis of biology, management and impacts in lodgepole pine. Victoria, BC:

Canadian Forest Service, Natural Resources Canada.

USDA Forest Service. 1971. Major forest insect and disease losses 1961-70.

Prepared by I&D Staff, December 1971. Insect and disease surveys are

completed every year by the Forest Service. Reports from the 1960s are

lost or were never published, but a tabulated summary of activity inArizona

and New Mexico is available at the USDAForest Service, Region 3, Forest

Health Protection office, Albuquerque, NM.

USDAForest Service. 2008. Forest insect and disease conditionsin the Southwestern Region, 2007. R3-08-01. Albuquerque NM: U.S. Department of

Agriculture, Forest Service, Southwestern Region, 41 p.

Wood, S.L. 1982.The bark and ambrosia beetles ofNorth andCentralAmerica

(Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist

Memoirs No. 6. 1359 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 487

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Population Status of Prairie Dogs (Cynomys ludovicianus) 

in the San Pedro River Basin, Sonora 

Efrén Moreno-Arzate and Carlos A. López González

Universidad Autónoma de Querétaro, Querétaro, Mexico

Gerardo Carreón Arroyo

Naturalia A. C. Hermosillo, Sonora, Mexico

Abstract—The black tailed prairie dog (Cynomys ludovicianus) is a species of conservation concern for 

Mexico, the United States and Canada. Populations in Mexico (including those in Sonora), which are considered endangered by the Mexican authority, require additional conservation efforts to maintain them on 

the long term. Our objective was to determine population size and density of the colonies in Sonora, as a 

first step to translocate individuals into a new colony in the region (Los Fresnos Ranch). The size of prairie 

dog colonies is 93 ha for La Mesa and 58 ha for Las Palmitas. La Mesa was surveyed between November 

2010 and December 2011, and Las Palmitas from May to December 2011. Density was calculated with 

distance-based transect counts, using sampling software (Distance 4.0) for the analysis of the data. Our 

density estimates for La Mesa range from 1.03 to 5.29 individuals/ha, with an estimated peak population 

size of 488 individuals. In Las Palmitas, densities ranged from 1.50 to 6.45 individuals/ha, with an estimated 

peak population size of 362 individuals. Our results suggest that a controlled but limited translocation effort 

will allow the establishment of a new colony at Los Fresnos without compromising the long-term persistence 

of extant colonies.

Introduction

	 Prairie dogs(Cynomysspp) are considered keystone species asthey

contribute to increase regional biodiversity (Ceballos and Pacheco

2000; Cully and others 2010; Royo-Marquez and Baéz-Gonzalez

2001; Winter and others 2002). The genus Cynomys is restricted

to North America and includes five species of which two occur in

Mexico: the Mexican prairie dog (C. mexicanus) and the black-tailed

prairie dog (C. ludovicianus). Black-tailed prairie dogs in Mexico

are distributed in two populations (Ceballos and Pacheco 2000), one

in the grasslands of northwestern Chihuahua (Ceballos and Mellink

1993) and the other in northern Sonora (Castillo 2004).

	 Currently, this species is present in 2% of its historical range

(Ceballos and Mellink 1993; Ramirez and Keller 2010). Population

declines have been partially attributed to poor livestock husbandry

and agricultural practices that result in the loss, fragmentation, and

disturbanceofnaturalhabitats;poisoningcampaigns,plague (Yersinia 

pestis), andrecreationalshooting(Hoogland2006;Ramirez andKeller

2010)have alsocontributedtodeclines.Prairiedogpopulationdeclines

have been relatively well documented for northwestern Chihuahua

(Avila and others 2012) but little is known about population changes

in northern Sonora. The two colonies in Sonora are located on the

edge of their distribution (Hoogland 1996) so that their abundance

may be lower than those presented in the distribution center (Lawton

1993). The colonies have no genetic connectivity with the colonies

of Chihuahua and, in the case ofArizona, were considered extirpated

long ago (Knowles and others 2002).Although recentmapping efforts

indicate that the prairie dog colony in La Mesa Ranch lost 60 ha (34%

loss) between 2004 and 2011 (Moreno-Arzate and López-González

2012), no study has documented changes in population numbers in

Sonora. Our objective was to determine the population status of the

only two colonies left in Sonora, La Mesa and Las Palmitas, based

on visual counts and distance-based methods. With this information,

we expect to determine whether these populations are able to support

extractions of individuals for translocation efforts in the region.

Study Area

	 The study area is located near the international border between

Sonora andArizona in the upper San PedroRiverBasin.The colonies

are located between parallels 30°37’12’’ and 31°19’48’’ N, and between meridians 109°48’36’’ and 110°37’12’’W (fig. 1). This area is

dominatedbyChihuahuandesertgrasslands.Elevationrangesbetween

1400 and 1500 m with an average annual rainfall of 300 to 750 mm.

The area has been impacted by livestock pressure causing changes

in the structure and composition of plant communities, including an

evident spread of mesquite (Castillo 2004).
488 USDA Forest Service Proceedings RMRS-P-67. 2013

Moreno-Arzate, López González, and Arroyo Population Status of Prairie Dogs (Cynomys ludovicianus) in the San Pedro River Basin, Sonora

Methods

	 La Mesa was surveyed between November 2010 and December

2011, and Las Palmitas from May to December 2011. We used strip

transects(Burnhamandothers1980)delineatedapproximatelythrough

the center of each colony to conduct visual counts of prairie dogs.

The transect length in the colony of La Mesa was 2100 m, while the

transect in the colony at Las Palmitas was 1100 m long. Each transect was surveyed twice a day, once between 0700 and 0800 hr and

once between 1400 and 1500 hr for 2 consecutive days. We slowly

walked each transect and recorded both the position and distance of

each individual prairie dog during our daily surveys.

	 The distance from the observer to the observed prairie dog was

calculated using a rangefinder. We also recorded the distance from

the start of each transect to the observation point, documenting the

position of the animal to either left or right side of the transect. We

used the programDistance v4.0 (Thomas and others 2005) to estimate

the population density within each colony. We identified significant

changesin density throughout the year comparing monthly data with

one-way ANOVAs.

Results

Density

	 The highest density at La Mesa was documented in March (5.0

individuals/ha) and July (5.3 individuals/ha). However, we found

significant differences among sampled months throughout the study

period (F = 11.90, df = 11, p = 0.0001). In the case of Las Palmitas,

we documented the highest densities in September and July (6.3 and

4.5 individuals/ha, respectively) and the lowest in November (1.2

individuals/ha). Significant differences (F = 21.7, df = 6, p = 0.0001)

between the months are shown in figure 2.

Population Size

	 After the surveys at colony La Mesa and using the highest monthly

density(Julywith5.29individuals/ha),we estimatedapeakpopulation

size of 488 individuals. The highest density at Las Palmitas produced

a peak population size of 362 individuals during September.

Discussion and Conclusions

	 During the monthsfrom May to July, the densities calculated in La

Mesa and Las Palmitas colonies showed an increase that coincided

with the beginning of the rainy season and continued after heavy

rainfall from July to September. This finding could be attributed to

an increase in pasture growth, available biomass, reflected in the

prairie dogs feeding activity (Hoogland 1996, 2003) and an increase

in the activity of juveniles. A different pattern was observed for the

months closerto thewinter,where cold temperatures and the available

biomass decreases, which caused a decrease in activity.

	 Population size estimated for both colonies confirmed that it is

possible to carry out the translocation of individuals to create a new

colony in the region.In the case ofthe colonyLaMesa,which presents

a population estimate above 400 individuals, this number allows the

extraction of a group of prairie dogs that could range between 60 to

100 individuals.The latter number,which represents about 25%ofthe

estimated population, might be interpreted as the number of prairie

dogs that can be extracted without having a demographic impact on

Figure 1—Geographic location of black-tailed prairie dog colonies in Sonora.
USDA Forest Service Proceedings RMRS-P-67. 2013 489

Population Status of Prairie Dogs (Cynomys ludovicianus) in the San Pedro River Basin, Sonora Moreno-Arzate, López González, and Arroyo

the founder colony (Robinette and others 1995; Dullum and others

2005). Similarly, a translocation effort could be supported by the

Las Palmitas colony as it has a population of over 350 individuals

in which one could draw between 70 and 90 individuals (≤ 25%)

without compromising the long-term sustainability of that colony. It

isimportantto continuemonitoring in both coloniesto estimate trends

in the population and identify the most appropriate dates where all

the necessary elements for a successful translocation may occur.

References

Avila-Flores, R., G. Ceballos, A.Villa-Meza, [and others]. 2012. Factors associated with long-term changesin distribution of black-tailed prairie dogs

in northwestern Mexico. Biological Conservation. 145: 54–61.

Biggins D.E., J. G. Sidle, D. B. Seery and A. E. Ernst.2006. Estimating the

abundance of prairie dogs. 2006. In: Conservation of the Black-Tailed,

Prairie Dog. John L. Hooglan, ed. Island Press, E.U., pp.107-94.

Burnham, K. P., D. R. Anderson and J. L. Laake. 1980. Estimation of density

from line transect sampling of biological populations. Wildlife Monographs. 72: 1-202.

Castillo, A. R. 2004. Evaluación de las poblaciones del perrito llanero (Cynomys ludovicianus) y su hábitat en Sonora, México. Informe técnico

final presentado a World Wildlife Fund Programa Desierto Chihuahuense.

Carreón.A.G., N. E. Lara,A. Gonzalez, H. Coronel, C. Lòpez, R. E.Jiménez.

2010. Fortalecimiento e implementación de un programa de monitoreo de

especies clave para la Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe y la RPC Cuenca del Río San Pedro. Reporte final al

FMCN.NATURALIA,A.C.-UniversisdadAutónoma deQuerétaro. 97pp.

Ceballos G, and R. L. Mellink. 1993. Distribution and conservation status of

prairie dogs Cynomys mexicanus and Cynomys ludovicianus in Mexico.

Biological Conservation. 63:105-112.

Ceballos, G. and J. Pacheco.2000. Los perros llaneros de Chihuahua. Biodiversitas. CONABIO, México D.F. 31:3-5.

Cully, J.F. Jr., S.K. Collinge, R.E. Van Nimwegen, [and others]. 2010 Spatial

variation in keystone effects: Small mammal diversity associated with

black-tailed prairie dog colonies. Ecography. 33: 667-677, 2010

Dullum, J. A., K. Foresman, Marc R. Matchett. 2005. Efficacy of translocationsforrestoring populations of black-tailed prairie dogs.Wildlife Society

Bulletin. 33(3): 842-850.

Hoogland, I. J. 1996. Cynomys ludovicianus. Mammalia Species. 53:51-10.

Hoogland, I. J. 2006. Conservation of the black-tailed, prairie dog. Island

Press, E.U. 357 p.

Knowles,C.,J.Proctor andS.Forest.2002.Black-tailedprairiedogabundance

and distribution in the Great plains based on historic and contemporary

information. Great Plains Research. A Journal of Natural and Social Sciences. 12: 219-254.

Lawton, J. H. 1993. Range, population abundance and conservation. Trends

in Ecology and Evolution. 8:409–413.

Moreno-Arzate, E. and López-González,A.C. 2011. Protocolo para la Traslocación de Perritos de la Pradera cola negra (Cynomys ludovicianus) en la

Reserva Los Fresnos, Cuenca del Río San Pedro, Sonora. Reporte Técnico

para Naturalia A.C.: Mayo-Junio-Julio y Resumen de los monitoreos realizados en La Mesa en el periodo de noviembre del 2010 a Julio del 2011.

Ramírez, J. and G. Keller. 2010. Effects of landscape on behavior of blacktailed prairie dogs(Cynomys ludovicianus) in rural and urban habitats. The

Southwestern Naturalist. 55:167–171.

Robinette, K.V., W.F. Andelet and K.P. Burnham. 1995. Effect of group size

on survival of relocated prairie dogs. Journal of Wildlife Management.

59:867-874.

Royo-Marquez, M., and A. D. Baéz-González. 2001. Descripción del hábitat

de áreas colonizadas y sin colonizar por perrito llanero (Cynomys ludovicianus) en el noroeste de Chihuahua. Tecnica Pecuaria Mexicana. 39:89–104.

Severson, K. E., and G. E. Plumb. 1998. Comparison of methods to estimate

population densities of black-tailed prairie dogs. Wildlife Society Bulletin.

26:859–866.

Thomas,L.,J.L.Laake,S.Strindberg,[andothers].2005.Distance5.0.Release

5.0. Research Unit for Wildlife Population Assessment, University of St.

Andrews UK. Shttp: www.ruwpa.st-and.ac.uk/distance/.

Winter, S.L., J.F. Cully, Jr., and J.S. Pontius. 2002. Vegetation of prairie dog

colonies and non-colonized short-grass prairie. Journal of Range Management. 55:502-508.

Figure 2—Monthly density estimated for the two prairie dog colonies surveyed in Sonora, Mexico.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
490 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Babocomari River Riparian Protection Project

Dan Robinett

Robinett Rangeland Resources LLC, Elgin, Arizona

Linda Kennedy

Appleton-Whittell Research Ranch of the National Audubon Society, Elgin, Arizona 

Abstract—The Babocomari River is a major tributary of the San Pedro River in Santa Cruz and Cochise counties, Arizona. This 140,000 acre catchment includes rolling grasslands on the Sonoita plain, oak woodlands 

in the Canelo Hills and the pine-oak forests of the northwestern Huachuca Mountains. The Babocomari River 

runs for 22 miles from its headwaters near Sonoita at 5000 feet elevation, eastward to join the San Pedro at 

Fairbanks at an elevation of 3850 feet. The U.S. Geological Survey estimates that this important tributary 

contributes about 6000 acre- feet of water annually to the San Pedro River system. The Arizona Department 

of Water Resources funded this 5-year study with a grant (09-164WPF) in 2009. Monitoring transects were 

installed in 2009 and 2010 and will be re-read each year through 2013. Objectives: Construct 2 miles of 

riparian boundary fence to restrict access by livestock from the Babocomari River. Install six stream riparian 

vegetation and geomorphic monitoring transects and six vegetation and geomorphic transects on riparian 

grasslands (sacaton) on tributaries to the Babocomari River. Analyze and summarize data annually and present that information to the participating ranch properties for use in making management decisions. Riparian 

monitoring stations were established in May of 2010 at three locations along the Babocomari River below 

the Babocomari Ranch (BR) headquarters and at three locations in June of 2010 on the Appleton-Whittell 

Research Ranch (ARR) of the National Audubon Society. These riparian monitoring transects were re-read 

in June of 2011. Riparian grasslands include large bottomlands of giant sacaton (Sporobolus Wrightii Monro 

ex Scribn) on both the BR and the ARR. Riparian grassland monitoring stations (three on each property) 

were established in the fall of 2009, re-read in 2010 and 2011. In addition to this monitoring effort the 

Babocomari Ranch has begun to protect private lands along the Babocomari River by selling development 

rights and placing conservation easements on the land. The land on the Research Ranch is already protected 

from development.

Introduction

	 The Babocomari River is a major tributary of the San Pedro River

in Santa Cruz and Cochise counties, Arizona. This 310 square mile

catchmentincludesrollinggrasslandsontheSonoitaplain,Chihuahuan

desertscrub oftheWhetstone pediment, oak woodlandsin theCanelo

Hills and the pine-oak forests of the northwestern Huachuca Mountains. The Babocomari River runs for 25 miles from its headwaters

near Sonoita at 5000 feet elevation, eastward to join the San Pedro

at Fairbanks at an elevation of 3850 feet (Cook and others 2009).

This important tributary contributes about 6000 acre feet of water

annually to the San Pedro River system (ADWR, 2005). Protection

and monitoring of the Babocomari River and associated ecosystems

are an important part of the management of water resources in the

area. The Arizona Department of Water Resources funded a 5-year

study with a grant (09-164WPF) in 2009. Monitoring transects were

installed in 2009 and 2010 andwill be re-read each yearthrough 2013.

	 The objectives of the grant are (1) construct 2 miles of fence to

restrict access by livestock to the Babocomari River corridor; (2a)

install six vegetation and geomorphic transects on riparian grasslands (sacaton); (2b) install six riparian vegetation and geomorphic

monitoring transects; (3) present annual results to the participating

ranch propertiesfor use inmakingmanagement decisions. This paper

addresses objectives 2a and 2b.

Methods and Results

Riparian Grasslands (Sacaton)

	 Large bottomlands of giantsacaton (Sporobolus Wrightii Monro ex

Scribn) are found on both ranches. Six riparian grassland monitoring

sites were established in the autumn of 2009 and were re-surveyed

for canopy and basal cover by vegetative species in 2010 and 2011.

Transects are re-surveyed for changesin channel morphology annually.

	 We present data from one riparian grassland (sacaton) transect as

an example.

	 Transect # 1 is on the ARR just below the confluence of Post Canyon and O’Donnell Canyon and was established by Dr. Ron Tiller in

1997. A rain gauge is within 600 meters and a piesometer (7 m deep

well) nearby isread quarterly for depth to groundwater. Thistransect
USDA Forest Service Proceedings RMRS-P-67. 2013 491

Babocomari River Riparian Protection Project Robinett and Kennedy

is 448.5 meters long. The southern half of the cross section burned

in the Canelo fire (wildfire) on May 6, 2009. Everything north of

the Post/O’Donnell Canyon channel was unburned. Ten, 100-meter

lines were re-sampled for vegetative cover by species and soil cover

at fixed stations along this transect.

	 Precipitation in 2011 (13 inches) was much lower than in 2010

(21.5 inches) and was only 75% of the long-term average. Total

perennial grass foliar cover on this transect was 43% with 7% basal

cover, down slightly from 2010 with giant sacaton dominating. Vine

mesquite (Hopia obtusa), blue grama (Bouteloua gracilis), green

sprangletop (Leptochloa dubia) and sideoats grama (Bouteloua curtipendula) made up the remainder of the cover. Basal cover of litter

was higher and bare soil lower in 2011, although the differences are

within sampling error. Annual forb cover was less in 2011 due to a

“La Nina” winter and very poor winter-spring moisture. (figs. 1, 2).

Riparian Stream

	 Monitoring stations were established in May of 2010 at three locations along the Babocomari River below the ranch headquarters.

They were re-read in June 2011. Data from one monitoring station

is included as an example.

	 BR #1 is on the Babocomari River and begins about 100 meters

downstream of the USGS stream gauge (#09471380), about 4 miles

eastoftheRanchheadquarters. Itrunsfromthe southwesttonortheast.

This portion of the Babocomari River was fenced to make a separate

(River) pasture in 1996 and has been managed by grazing only during

the winter months and for short periods of time (2 to 3 weeks) since

then. The pasture was not grazed in either 2009 or 2010.

	 Themonitoring at thislocation consists of a cluster ofthree riparian

green-line transects for herbaceous vegetation paired with three belt

transects (3 m wide) on both banks to record tree species by canopy

Figure 1—Percent canopy cover for TRR #1.

Figure 2—Percent basal cover for TRR #1.
492 USDA Forest Service Proceedings RMRS-P-67. 2013

Robinett and Kennedy Babocomari River Riparian Protection Project

Figure 3—Percent canopy cover for BR #1.

cover. Green-line transects (Medina 2008) are 40 m in length along

both banks. This technique helps segregate the plant community into

banklovingandwaterlovingspecies.Data arepresentedseparatelyfor

bank plots and water plots. Herbaceous vegetative data are presented

as average canopy cover by species summed for all three transects

in a cluster. Tree vegetative data are presented as average canopy

cover by species and average total canopy cover for the transect area.

Total cover is often less than the sum of individual tree species cover

because individual tree canopies overlap.

	 Geomorphic monitoring includes three survey cross sections, one

perpendicular to the stream channel at the midpoint of each transect

in the cluster.

	 The vegetative data show a diverse plant community dominated by

native wetland species. Deergrass (Muhlenbergia rigens) dominates

thebankplotswithlesser amountsofhorsetail(Equisetum laevigatum)

and spike rush (Eleocharis palustrus) (fig. 3).

	 Deergrass and watercress (Rorippa nasturtium-aquaticum) dominate the water plots with lesser amounts of spike rush, horsetail, and

several important aquatic forbs. Banks are extremely well vegetated

with only 4 to 5 % bare (exposed) soil. There were no significant

changes in the understory plant community from 2010 to 2011. Total

tree canopy cover averaged over all three transects is 63%. Cottonwood (Populous fremontii) is dominant and Gooding willow (Salix 

goddingii)issub-dominant.There were no significant changesin tree

canopy from 2010 to 2011.

	 Geomorphic crosssectionswere surveyed at 2-to 3-meterintervals

andperpendiculartothestreamateachtransect.Crosssectionsspanned

the floodplain and low stream terraces on either side. Re-survey of

the cross sections show little change in the last 3 years.

Conclusions

	 Monitoringinformationdevelopedduringthisstudyisbeingusedto

helpmakemanagementdecisionsthatmaintainandimprovevegetative

conditions along theBabocomariRiver,severalmajortributaries, and

the Babocomari Cienega. Both the BR and ARR management intend

to continue monitoring at these sites after this WPF grant expires.

The water, plant, and animal resources of this system are diverse and

rarely found intact with such good environmental and hydrologic

conditions. In addition to this monitoring effort, the BRR has begun

to place conservation easements on the land. The land on the ARR is

already protected from development. The Babocomari River and its

upper watershed are worth preserving and protecting for generations

to come.

References

ADWR (Arizona Department of Water Resources). 2005. Upper San Pedro

Basin Active Management Area Review Report. Arizona Department of

Water Resources, Phoenix, AZ. 

Cook,JosephP.,AnnYouberg,PhilipA.Pearthree,[andothers].2009.Mapping

of Holocene River Alluvium along the San Pedro River, Aravaipa Creek,

and Babocomari River, Southeastern Arizona. Arizona Gelogical Survey.

Digital map RM 1. 75 p. www.azgs.az.gov.

Medina, Alvin. 2008. U.S. Rocky Mountain Forest and Range Experiment

Station, Flagstaff, AZ. (personal communication)

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 493

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Important Bird Areas of the Madrean Archipelago: A 

Conservation Strategy for Avian Communities

Vashti (Tice) Supplee

Arizona Important Bird Area Program, Audubon Arizona, Phoenix, Arizona

Jennie MacFarland

Arizona Important Bird Area Program, Tucson Audubon Society, Tucson, Arizona

Abstract—The Important Bird Areas (IBA) Program is a worldwide program through BirdLife International 

that identifies sites considered to provide important habitats for avian species. Criteria for designation are 

species abundance, diversity, and range restriction. As the United States Partner of BirdLife International, 

the National Audubon Society administers the IBA Program in the United States. In Mexico, the network of 

AICAs is administered by CONABIO. The Madrean Archipelago has a diverse range of Identified IBAs (AICAs,

Áreas de Importancia para la Conservación de las Aves, in Mexico) that feature riparian, wetland, grassland, 

and Madrean oak woodland habitats. Five of the 24 IBAs and three of the AICAs in the region are globally 

significant: the Chiricahua Mountains, San Pedro Riparian National Conservation Area, Whitewater Draw 

State Wildlife Area, and Willcox Playa. 

Introduction

	 The purpose of designation as an IBA is highlighting the value of

specificsites.ThelowerelevationriparianIBAsarevitalforneotropical

migrantsin the spring aswell as providing habitatforresidentspecies.

In these habitats,which are particularly productive in the spring, these

species can refuel and rest before continuing on their journey north

to breeding grounds. The high elevation “Sky Island” IBAs are very

productive in the fall and serve as vital stop-over points for migrants

headed south. Many birds complete their interrupted molt in these

IBAs. This network serves to identify those habitats most important

to conserve to benefit native birds.

Important Bird Areas

Bird Conservation Region 34: Sierra Madre 

Occidental-Sonoran Joint Venture

	 The Sierra Madre Occidental “Sky Islands” and associated grasslandsandripariancorridorsareasharedecologicalsystemwithArizona

and New Mexico, USAand Sonora, Mexico. The biological diversity

ofthisregioniswelldescribedandincludessignificantsitesinArizona

such as the Chiricahua and Huachucha Mountains, the San Pedro

River, and the San Rafael Grasslands.The NationalAudubon Society

Appleton-Whittell Research Ranch is located within the grasslands

and Madrean oak woodland communities. The headwaters of the San

Pedro River and the southern portion of the San Rafael grasslands

are in Sonora, Mexico, where the continuation of the mountain Sky

Islands provides a connection south into the sub-tropical regions of

the Sierra Madre Occidental.

	 Among the species whose range extends into the United States in

this region, highest priorities include Mexican Spotted Owl, Eared

Quetzal, Lucy’s Warbler, Red-faced Warbler, Strickland’s (Arizona)

Woodpecker, and Montezuma Quail. Riparian areas in lowlands

support many in-transit migrants as well as breeding Thick-billed

Kingbirds, Bell’s Vireo, and Western Yellow-billed Cuckoos.

	 Identification of Mexico AICAs in northern Sonora and United

States IBAs within the Apache Highlands ecoregion as a globally

significant International IBA/AICA management zone will serve as

a unifying statement of shared birds and habitats. An International

and global AICA/IBA is supported by the Sonoran Joint Venture and

Audubon as a contributing strategy to promote conservation ofshared

border birds.

Sistema de Islas Sierra Madre Occidental/

Sierra Madrean Sky Islands 

Globally Important Species

Strix occidentalis Spotted Owl (USA and Mexico)

Euptilotis neoxenus Eared Quetzal (Mexico)

Vireo bellii Bells’ Vireo (USA and Mexico) Riparian

Calcarius ornatus Chestnut-collared Longspur (winter) (USA and

Mexico) grasslands

Global Biodiversity—Assemblage of Biome-restricted species

Continental Important Species

Cyrtonyx montezumae Montezuma Quail

Picoides stricklandi Arizona Woodpecker

Coccyzus americanus Western Yellow-billed Cuckoo

Asio flammeus Short-eared Owl (Winter)

Otus trichopsis Whiskered Screech Owl
494 USDA Forest Service Proceedings RMRS-P-67. 2013

Supplee and MacFarland Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities

Micrathene whitneyi Elf Owl

Caprimulgus ridgwayi Buff-collared Nightjar

Lampornis clemenciae Blue-throated Hummingbird

Calypte costae Costa’s Hummingbird

Melanerpes lewis Lewis’s Woodpecker (winter)

Colaptes chrysoides Gilded Flicker

Tyrannus crassirostris Thick-billed Kingbird

Vermivora luciae Lucy’s Warbler

Dendroica graciae Grace’s Warbler

Cardellina rubrifrons Red-faced Warbler

Pipilo aberti Abert’s Towhee

Aimophila carpalis Rufous-winged Sparrow

Aimophila quinquestriata Five-stripped Sparrow

Calcarius mccownii McCown’s Longspur (Winter)

Spizella breweri Brewer’s Sparrow (Winter)

Spizella atrogularis Black-chinned Sparrow

Ammodramus bairdii Baird’s Sparrow

Species of Conservation Concern

Callipepla squamata Scaled Quail

Arizona Important Bird Areas

Appleton-Whittell Audubon Research Ranch—National Audubon,

BLM, Coronado National Forest-Grasslands

Globally important bird species: Montezuma Quail

Chestnut-collared Longspur and McCown’s Longspur (winter)

Buenos Aires National Wildlife Refuge—U.S. Fish and Wildlife

Service

Globally important bird species: Masked Bobwhite Quail

California Gulch—Coronado National Forest

Globallyimportantbirdspecies:Five-strippedSparrow,SpottedOwl

Chiricahua Mountains—Coronado National Forest

Globally important bird species: Spotted Owl,Whiskered ScreechOwl,Violet-crownedHummingbird,ArizonaWoodpecker,Sulphurbellied Flycatcher, Buff-breasted Flycatcher, Grace’s Warbler and

Mexican Chickadee

Huachuca Mountains—Coronado National Forest

Globally important bird species: Spotted Owl

Santa Cruz River—Arizona State Park, U.S. Park Service

Globally important bird species: Bell’s Vireo

San Pedro River Riparian National Conservation Area—Bureau of

Land Management

Globally important bird species: Bell’s Vireo

San Rafael Valley—Coronado National Forest, Arizona State Parks

Globally important bird species: Chestnut-collared Longspur

(winter), McCown’s Longspur (winter)

Santa Rita Mountains—Coronado National Forest

Globally important bird species: Spotted Owl

Sonoita Creek—The Nature Conservancy and Arizona State Park

Globally important bird species: Bell’s Vireo

Sycamore Canyon—Coronado National Forest

Globally important bird species: Five-stripped Sparrow, Spotted

Owl, Buff-collared Nightjar

Whitewater Draw State Wildlife Area—Arizona Game and Fish

Globally important bird species: Sandhill Crane

Potential Arizona Important Bird Areas:

Patagonia Mountains and Pinaleno Mountains–Coronado National

Forest

Globally important bird species: Spotted Owl

LasCienegasNationalConservationArea—BureauofLandManagement

Rufous-winged Sparrow

New Mexico Important Bird Areas:

Clanton Canyon—Coronado National Forest

MontezumaQuail,WhiskeredScreech-Owl,ArizonaWoodpecker,

Whip-poor-will,ElfOwl,Dusky-cappedFlycatcher,Grace’sWarbler

Gray Ranch Grasslands—Animas Foundation

Botteri’s Sparrow

Guadalupe Canyon – Bureau of Land Management

Costa’s, Broad-billed, Blue-throated, Magnificent, and Violetcrowned Hummingbird, Northern Beardless-Tyrannulet, Thickbilled Kingbird, and Varied Bunting.

References

American bird conservancy guide to 500 most important bird areas in the

United States. 2003. Random House. 518 p. ISBN 0-8129-7036-5.

Arizona Important Bird Areas webpages: www.aziba.org.

Birds of North America Online. [n.d.] Website. Ithaca, NY: Cornell Laboratory of Ornithology. Online: http://bna.birds.cornell.edu/bna/. [accessed

January 3, 2013].

Devenish,C.;Diaz-Fernandez,D.F.;Clay,R.P.; [andothers], eds.2009.Important bird areas:Americas-priority sitesfor biodiversity conservation.Quito,

Ecuador: BirdLife International (BirdLife Conservation Series No. 16).

Latta,M.J.;Beardmore,C.J.;Corman,T.E.1999.ArizonaPartnersinFlightBird

Conservation Plan. Ver. 1.0. Nongame and Endangered Wildlife Technical

Report 142, Arizona Game and Fish Department, Phoenix.

NationalAudubonSociety,ImportantBirdAreas,Arizonawebpageshttp://iba.

audubon.org/iba/viewState.do?state=US-AZ and New Mexico webpages

http://iba.audubon.org/iba/viewState.do?state=US-NM
USDA Forest Service Proceedings RMRS-P-67. 2013 495

Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities Supplee and MacFarland

Appendix: Migratory Birds of Conservation Focus 

Map sources: Birds of North America; Cornell Laboratory of Ornithology: green = winter; dark tan = 

breeding; light tan = migration; blue = yearlong. 

Gray Hawk Asturina nitida Common Black-Hawk Buteogallus anthracinus. Mexico has resident populations (in 

the South) and migratory (in the north).

Elf owl Micrathene whitneyi

Mexican whip-or-will Caprimulgus 

vociferous

Thick-billed kingbird Tyrannus crassirostris Buff-breasted flycatcher Empidonax 

fulvirons
496 USDA Forest Service Proceedings RMRS-P-67. 2013

Supplee and MacFarland Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities
USDA Forest Service Proceedings RMRS-P-67. 2013 497

Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities Supplee and MacFarland

Rose-throated becard Pachyramphus 

aglaiae

Northern Beardless tyrannulet Camptostoma imberbeOlive warbler Peucedramus taeniatus Lucy’s warbler Vermivora luciae
498 USDA Forest Service Proceedings RMRS-P-67. 2013

Supplee and MacFarland Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities

Grace’s Warbler Dendroica graciae Red-faced warbler Cardellina rubrifrons Five-stripped sparrow Aimophila quinquestriata

Varied bunting Passerina versicolor Botteri’s sparrow Aimophila botteri Baird’s sparrow Ammodramus bairdi
USDA Forest Service Proceedings RMRS-P-67. 2013 499

Important Bird Areas of the Madrean Archipelago: A Conservation Strategy for Avian Communities Supplee and MacFarland

McCowan’s longspur Calcarius mccownii

(winter) (winte

Chestnut-collared longspur Calcarius ornatus Lark bunting Calamospiza melanocorys (winter) (winter)

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
500 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; 498comps. 

2013. Merging science and management in a rapidly changing world: 

Biodiversity and management of the Madrean Archipelago III; 2012 

May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research 

Station. 

Preliminary Flora of Ojo de Agua Tonibabi, Sierra La 

Madera, Sonora, Mexico 

Melissa Valenzuela-Yánez, Gertrudis Yanes-Arvayo, Maria de la Paz Montañez-Armenta, and 

Hugo Silva-Kurumiya

Universidad de la Sierra, Moctezuma, Sonora, Mexico

Thomas R. Van Devender

Sky Island Alliance, Tucson, Arizona 

Abstract—Ejido Tonibabi is located 12 kilometers east-northeast of Moctezuma in east-central Sonora, 

México (29°50’16’’N 109°33’45’’W, 780 m elevation). The vegetation is riparian in the wetlands and foothills 

thornscrub on slopes. The flora was inventoried on transects in different areas in Ejido Tonibabi. Specimens 

were collected to identify by comparing with specimens in the University of Arizona and Universidad de 

Sonora Herbaria and for voucher specimens. A total of 162 species in 125 genera and 47 families were 

documented in the area. The families with the most species were Fabaceae, Asteraceae, Euphorbiaceae, 

Convolvulaceae, Poaceae and Cactaceae, which contain 53.1% of the flora. The most important life form was 

annual herbs (47 taxa), followed by perennial herbs (40 taxa), trees (21 taxa), shrubs (15 taxa), succulents 

(10 taxa), grasses (9 taxa), subshrubs (9 taxa), vines (8 taxa) and aquatic herbs (3 taxa). The Tonibabi flora 

contains 4.4% of the state flora. 

Introduction

Sierra La Madera is a Sky Island mountain range located in the 

transition between the New World tropics and the northern temperate zone at about 29°N in east-central Sonora, which enhances the

diversityof floraand fauna.Complextopographyandanelevational

range of ca. 1685 m (from 615 m along Río Bavispe at Huásabas to

over 2300mon the highest peak)resultin diverse habitatsin theSierra

la Madera. The Ojo deAgua Tonibabi isin the southwestern foothills

of the Sierra la Madera in east-central Sonora, México. There are no

published florasin foothillsthornscrub in Sonora. Yanes-Arvayo and

others (2011) present a photographic guide to the common plants of

the Sierra La Madera. We report a preliminary flora of the Tonibabi

area, and compare it with the flora of the Río Bavispe region in

northeastern Sonora (White 1948) in this paper.

Study Area

	 The Ojo de Agua Tonibabi (29°50’16’’N, 109°33’45’’W, 780 m

elevation) is in the Ejido Tonibabi 12 kilometers east-northeast of

Moctezuma intheMunicipio(=County)deMoctezuma ineast-central

Sonora, México. Soils are very thin soil and near the wet areassaline,

which indicates that groundwater is near the surface. The vegetation

in the Tonibabi study area is willow (Salix gooddingii) and mesquite

(Prosopis velutina) in riparian areas along streams and foothills

thornscrub on slopes. Foothills thornscrub is a tropical shrub-tree

formation that occurs in a broad transition between the Sonoran

Desert to the west and tropical deciduous forest or oak woodland in

the Sierra Madre Occidental to the east (Búrquez and others 1999;

Martínez-Yrízar and others 2010).

Methods

	 Selectedsiteswiththehighest concentrationofplantsintheTonibabi

area were visited on eight field trips between April and December

2011. Plants were collected and observed in lowland riparian habitats

near streams, and on rocky slopes and disturbed roadsides. Plant collectionswere identified usingfloristic references andcomparisonwith

specimens in the University of Arizona and Universidad de Sonora

(USON) herbaria. Online resources consulted were the University of

Arizona herbarium (http://ag.arizona.edu/herbarium/), the Madrean

ArchipelagoBiodiversityAssessment(MABA)(http://www.madrean.

org), and the United States Department of Agriculture Plants (http://

plants.usda.gov) databases. Voucher specimens were deposited into

USON. Plant collections and imagesin this study are available in the

MABA database.

Results

	 A total of 162 plant species in 125 genera and 47 families were

identified in the Tonibabi area. The families with the most species

were Fabaceae (27 species, 16.7%), Asteraceae (20 species, 12.3%),

Euphorbiaceae (12 species, 7.4%), Cactaceae (10 species, 6.2%),

Poaceae (9 species, 5.6%), and Convolvulaceae (8 species, 4.9%; fig. 1).

These families contained 86 species in 60 genera and 86 species
USDA Forest Service Proceedings RMRS-P-67. 2013 501

Preliminary Flora of Ojo de Agua Tonibabi, Sierra La Madera, Sonora, Mexico Valenzuela-Yánez and others

(53.1% of the flora). The genera with the most species were Acacia,

Boerhavia, Bouteloua, Bursera, Desmodium, Euphorbia, Ipomoea,

Jatropha, Opuntia, and Portulaca with 19.5% of the total species.

	 The most important life forms in Tonibabi are herbs and trees

(66.7% of the flora), followed by cacti and grasses (11.7% each),

shrubs (9.5%), subshrubs (5.5%), vines (4.9%) and aquatic herbs

(1.8%). Riparian species are Goodding willow (Salix gooddingii),

mesquite (Prosopis velutina), tésota (Acacia occidentalis), batamote

(Baccharis salicifolia), cadillo (Xanthium strumarium), and Mimulus 

guttatus. Matelea pringlei and Tragia jonesii are regional endemic

species only found in Sonora. Pseudabutilon thurberi also occurs in

adjacent Arizona. Amoreuxia palmatifida and Olneya tesota have a

PR (Protected) status under NOM-059-SEMARNAT-2010. Only

four species in the Tonibabi flora (2.5%) were non-native plants:

Nasturtium officinale, Nicotiana glauca, Melinis repens and Pennisetum ciliare (Van Devender and others 2009).

Discussion

	 The State of Sonora has a diverse flora with 3483 species (3237

natives) including 104 varieties, 43 subspecies, and 30 interspecific

hybrids. Only 78 taxa (68 species, 8 subspecies, and 2 varieties) are

endemic to mainland Sonora (Van Devender and others 2010). The

Tonibabi flora has widespread riparian/wetlands species and typical

tropical foothills thornscrub species. As in many floras, herbs are

numerically dominant in the flora, with relatively few trees, shrubs

and large succulentssuch asBursera fagaroides, B. laxiflora, Mimosa 

distachya, Lysiloma divaricatum, Jatropha cordata, and Stenocereus 

thurberi, structural dominantsin the vegetation.Treeswith 21 species

are only 13% of the flora.

	 Stephen S. White from the University of Michigan led expeditions to study the flora of the Río Bavispe Region in northeastern

Sonora from 1938 to 1941 (White 1948). He collected in a wide

range of habitats in the region, resulting in 1200 species (currently

995 taxa after revision) in 549 genera in 114 families. In July1938,

he collected in the Moctezuma area including eleven specimens at

Tonibabi.Eustoma exaltatum, Martynia althaeifolia,Phaulothamnus 

spinescens, and Samolus valerandissp. parviflorus were not found in

the present study. Catchfly prairie gentian (E. exaltatum) is an herb

with showy lavender flowersfound in seeps and springs and israre in

Sonora. Only 76 species (47%) in the Tonibabi flora are shared with

the Río Bavispe flora, reflecting White’s brief visit to the area, and

that foothills thornscrub is only found in the Río Bavispe Region in

the southern end of the Sierra El Tigre west across the to the Sierra

la Madera.

	 Tonibabi with its extensive wetlands is a unique area in foothills

thornscrub. Large areas of whitish soil visible from satellite images

suggest that the wetlands were much more extensive in the past.

Study of pollen in the spring sedimentsshow that the ciénega formed

8300 years ago in the early Holocene and was wetter than today until

after 2300 years ago (Espinoza-Encinas 2012). Plants in the Ojo de

Agua Tonibabi are disjunct from other wetlands and possibly relict

populations from wetter past climates.

Figure 1—Species and genera in the most important families in the Tonibabi flora.
502 USDA Forest Service Proceedings RMRS-P-67. 2013

Valenzuela-Yánez and others Preliminary Flora of Ojo de Agua Tonibabi, Sierra La Madera, Sonora, Mexico

Acknowledgments

We thank George M. Ferguson for his careful review.

References

Búrquez,Alberto;Martínez-Yrízar,Angelina; Felger,Richard S.; andYetman,

David. 1999. Vegetation and habitat diversity at the southern edge of the

Sonoran Desert. Pg. 36-67 in Robichaux, Robert H., ed. Ecology of Sonoran Desert plants and plant communities.Tucson: University ofArizona.

Espinoza-Encinas, Iván Rosario. 2012. Paleoambientes Cuaternarios de la

Ciénega de Tonibabi, Moctezuma, Sonora. Masters thesis. Hermosillo:

Universidad de Sonora. 144 p.

MABA database (Madrean Archipelago Biodiversity Assessment). Online:

http://www.madrean.org/maba/symbflora/

Martínez-Yrízar,Angelina;Felger,RichardS.; andBúrquez,Alberto.2010.Los

ecosistemas terrestres: un diverso capital natural. Pg. 129-156 in MolinaFreaner, Francisco E.; and Van Devender, Thomas R. eds. Diversidad BiológicadeSonora. Hermosillo:UniversidadAutónomaNacionaldeMéxico.

Norma Oficial Mexicana. 2010. NOM-059-SEMARNAT-2010. Diario Oficial de la Federación. Segunda sección. Jueves 30 de diciembre de 2010.

Mexico. 78 p.

Van Devender, Thomas R.; Felger, Richard S.; Fishbein, Mark; [and others].

2010. Biodiversidad de las plantas vasculares. Pg. 229-261 in MolinaFreaner, Francisco E.; and Van Devender, Thomas R. eds. Diversidad

Biológica de Sonora. Hermosillo: Universidad Autónoma Nacional de

México. Online: http://www.skyislandalliance.org/Sonoran%20vascular%20plants-VanDevender%20et%20al.2010.pdf.

Van Devender, Thomas R.; Felger, Richard S.; Reina-Guerrero, Ana L.; and

Sánchez–Escalante,J.Jesús. 2009.Sonora:Non-native and invasive plants.

Pp. 85-124 in Van Devender, Thomas R.; Espinosa-García, Francisco J.;

Harper-Lore, Bonnie L.; and Hubbard, Tani. eds., Invasive plants on the

move. Controlling them in North America. Proceedings of Weeds Across

Borders 2006 Conference; Hermosillo, Sonora; May 25-28, 2006. Tucson,

AZ:SkyIslandAlliance.Online:http://www.skyislandalliance.org/WAB06-

c19-Van%20Devender%20et%20al-Sonoran%20non-natives-Jan08.pdf.

White, Stephen S. 1948. The vegetation and flora of the region of the Río de

Bavispe in northeastern Sonora, Mexico. Lloydia.11: 229‐302.

Yanes-Arvayo, Gertrudis; Montañez-Armenta, María de la Paz; SilvaKurumiya, Hugo; [and others]. 2011. Catálogo de Plantas de Sierra La

Madera. Arbóreas, Arbustivas y Herbáceas. Universidad de la Sierra,

Hermosillo, Sonora.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 503

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Examining Wildlife Responses to Phenology and Wildfire 

Using a Landscape-Scale Camera Trap Network

Miguel L. Villarreal, Leila Gass, Laura Norman, Joel B. Sankey, Cynthia S.A Wallace, and Dennis 

McMacken

USGS Western Geographic Science Center, Tucson, Arizona

Jack L. Childs

Borderlands Jaguar Detection Project, Amado, Arizona 

Roy Petrakis

University of Arizona, Tucson, Arizona

Abstract—Between 2001 and 2009, the Borderlands Jaguar Detection Project deployed 174 camera traps 

in the mountains of southern Arizona to record jaguar activity. In addition to jaguars, the motion-activated 

cameras, placed along known wildlife travel routes, recorded occurrences of ~ 20 other animal species. We 

examined temporal relationships of white-tailed deer (Odocoileus virginianus) and javelina (Pecari tajacu) 

to landscape phenology (as measured by monthly Normalized Difference Vegetation Index data) and the 

timing of wildfire (Alambre Fire of 2007). Mixed model analyses suggest that temporal dynamics of these two 

species were related to vegetation phenology and natural disturbance in the Sky Island region, information 

important for wildlife managers faced with uncertainty regarding changing climate and disturbance regimes.

Introduction

	 The Madrean Archipelago is known to have exceptional wildlife

diversity, related in part to the wide variety of available habitat in the

area as well as its location at the confluence of ranges of subtropical

and northern species (Felger and Wilson 1994). Monitoring wildlife

populationsin this area has become increasingly important as humancaused and natural disturbances (e.g. United States/Mexico Border

fence,drought andclimate change) affectwildlifemovement corridors

and the quality and quantity of available habitat. Populations of terrestrial vertebrates, both herbivores and predators, are dependent on

primary productivity of vegetation for their survival, and fluctuate

according to resource availability (White 2008). This dynamic is

often amplified in arid and semi-arid ecosystems where populations

respond to resource pulses, in particular growth of vegetation, driven

by both local and global climatic patterns(Marshal and others 2002).

Vegetationdynamics,whichincludeplantphenology, are capturedand

quantified by satellite imagery and have been used to model wildlife

habitat,vegetationcover,andspeciesabundance (Wallace2008;Sesnie

and others 2011). In addition to climate-induced variability in plant

productivity, resource availability for mammals can also be limited

or enhanced by natural disturbances like wildfire (Smith 2000). For

example, ungulates are responsive to changesin vegetation structure,

feeding in areas with increased plant growth after high-intensity fires,

and carnivores and omnivores are believed to be opportunistic and

will often feed near areas of recent burns where prey abundance has

increased (Smith 2000).

	 Monitoringwildlifepopulationsistraditionallyaccomplishedusing

labor-intensive human observation (e.g. avian surveys) or expensive

trapping methods (e.g. capture-recapture). An alternative to human

observationandtrappingare remote,motion-triggeredcamera “traps.”

Camera traps are typically set up to capture information on rare and

secretive species of conservation concern (McCain andChilds 2008),

but are increasingly being used tomonitor generalwildlife population

dynamics (Rowcliffe and Carbone 2008). Motion-triggered cameras

offerseveral benefits overtraditionalmethods of wildlife observation

(see Cutler and Swann 1999); they are noninvasive, can be set up

in rugged and remote locations, and provide continuous monitoring

while the camera remains operational. This last point is an important

distinction in that repeat, long-term camera monitoring provides a

photographic time series that can be analyzed with continually collected satellite and climate data allowing us to examine connections

between wildlife abundance and vegetation phenology.

	 In this project, we used an existing, landscape-scale camera trap

network to examine the relationship between wildlife dynamics,

vegetation phenology, and fire.We chose two example species,whitetailed deer (Odocoileus virginianus) and javelina (Pecari tajacu), 

which are the primary prey of the jaguar and other felids as well as

economically important game species in Arizona. It was anticipated

that the multi-species information gathered from the photographic

database might provide insight into decadal and annual population

dynamics, by capturing ameasure ofwildlife abundance atthe camera

locations as well as activity in the movement corridors on which the

cameras were located. In addition, this study was conceived to help
504 USDA Forest Service Proceedings RMRS-P-67. 2013

Villarreal, Gass, Norman, and others Examining Wildlife Responses to Phenology and Wildfire . . .

guide future trap efforts and study design by demonstrating the value

of incidental, non-targeted species data for understanding wildlife

dynamics related to climate and vegetation, and illustrating some of

the challenges of data analysis at a landscape scale.

Study Area

	 The study area is located at the western portion of the Madrean

Archipelago north of the United States/Mexico border stretching

from the Altar Valley to the San Rafael Valley (McCain and Childs

2008). Wildlife monitoring sites were located at the Baboquivari,

Sierrita,Tumacácori,Atascosa, Patagonia, andSantaRitaMountains.

Vegetation ofthe study sites consists ofMadrean evergreen woodland

and semidesert scrub grassland (Brown 1994) and elevations range

from 934-1944 m. The average maximum temperature recorded at

Tumacácori National Historical Park (1948-2005) was 36 °C, (July),

the average minimum was 0 °C (January), and average annual precipitation was 40 cm.

Data and Methods

	 Primary data for this project were (1) animal detections collected

by the Borderlands Jaguar Detection Project (McCain and Childs

2008) using a series of remote cameras and (2) 250-meter resolution

NormalizedDifferenceVegetation Index (NDVI) data.The following

sections provide details of the camera and vegetation data sets.

Camera Trap Data

	 A total of 8 (174 total cameras) different camera sites were monitored between 2001 and 2009 using passive infrared camera systems

that detect heat-in-motion. Cameras were intentionally located in

areas with high probability of detecting jaguar movement (i.e. game

trails, washes, and canyon bottoms). Cameras were typically left

in the original location for the entire length of the study in order to

detect variations in movement over time; however, some cameras

were moved to more strategic nearby sites if higher animal activity

was found in the area.

Moderate Resolution NDVI

	 WeusedNormalizedDifferenceVegetationIndex(NDVI)calculated

fromModerateResolutionImagingSpectroradiometer(MODIS)data.

Composites ofMODISNDVI datawere acquired fromtheUniversity

ofArizonaRemoteSensingCenterfor2000-2009.NDVIis calculated

using a ratio of the spectral properties of vegetation reflectance in the

red and near-infrared wavelengths of the electromagnetic spectrum;

green vegetation typically haslow reflectance in red wavelengths and

high reflectance in the near-infrared wavelengths. NDVI values are

positively correlated with the relative abundance of green, growing

vegetation.

Analysis

	 Detection photographs were compiled into a digital database with

each camera location attributed with the number of animal sightings

per month by species, monthly NDVI, and time-since-fire values

(months) for camerasthat were in or near theAlambre burn perimeter

(July 7, 2007 in the Baboquivari Mountains).

	 Counts of animal sightings per month were analyzed and histograms of whitetail sightings were constructed by mountain range

(8 total). The relationship of whitetail sightings with NDVI among

mountain ranges was evaluated with quantile regression. Additionally, a mixed model analysis was performed with whitetail sightings

as the response, mountain range, month, and NDVI as fixed effects,

and year and camera as random effects. The relationship of javelina

sightings with NDVI near theAlambre burn perimeter was evaluated

using a mixed model analysis with NDVI as a fixed effect and year

and camera as random effects. The relationship of javelina sightings

with months since fire was similarly evaluated with the time variable

as a fixed effect and year and camera as random effects.

Results and Discussion

	 Monthly sightings of whitetail ranged from very low to as large

as 50 in the different mountain ranges. Two-tailed Student’s t-tests

show that camera locations with whitetailsightings have significantly

lower NDVI values than camera locations with no sightings (fig. 1).

Figure 1—Quantile regression results for monthly white-tailed deer sightings as a function of NDVI. Tau (T) refers to the quantile of whitetail sightings 

conditional to NDVI, and B1 values are the slopes of the regression lines, which were significant with p < 0.05 (left). Mean (standard error) monthly 

NDVI for camera locations with Whitetail deer sightings during the study period (right).
USDA Forest Service Proceedings RMRS-P-67. 2013 505

Examining Wildlife Responses to Phenology and Wildfire . . . Villarreal, Gass, Norman, and others

Mixed model analysis indicated that mountain range (p = 0.1) and

NDVI(p = 0.06) had a weaklymoderately significant correlation with

whitetailsightings, whereas month did not appear to be significant (p

= 0.27). Quantile regression analysis showed a significant negative

relationship forthe largest values of whitetailsightings conditional to

NDVI(fig. 1).Thissuggeststhat lowabundance ofwhitetailsightings

occurred for a wide range of NDVI; however, the largest monthly

sightings occurred during periods of relatively low NDVI. On average, large values of NDVI occurred during the summer monsoon and

post-monsoon months ofAugust, September, and October(fig. 2) and

NDVI was comparably lower during the other months of the year

when sightings were greater. This observed pattern may be related to

greater whitetail activity during the breeding season (winter months).

	 Mixed model analyses indicated that NDVI and time since fire

were significant for javelina sightings (p = 0.02, respectively). Javelina sightings increased with time (1-24 months) since burning. The

relationship of javelina sightings to NDVI was also positive. NDVI

often decreases immediately after fire, then increases with time as

vegetation recovers, and therefore might be expected to covary with

time since burning; however, histograms depicting observationsfrom

a camera in the fire perimeter compared to those outside suggest that

the increase of javelina was related to post-fire effects (fig. 2).

Figure 2—Distribution of javelina sightings at the Coyote Mountain study site before and after the Alambre Fire of July 7, 2007. One camera located 

inside the burn perimeter showed a marked increase in javelina sightings in 2008 when compared to cameras outside the burn area.

References

Cutler, Tricia L., and Don E. Swann. 1999. Using remote photography in

wildlife ecology: a review. Wildlife Society Bulletin. 27(3): 571–581.

Felger, Richard S., and Michael F. Wilson. 1994. Northern Sierra Madre 

Occidental and its Apachian outliers: a neglected center of biodiversity.

Pages 36–59 in DeBano, F.L., Ffolliott, P.F., Ortega–Rubio, A., Gottfried, 

G.J., and Hamre, R.H., eds. Biodiversity and management of the Madrean 

Archipelago: the sky islands of southwestern United States and northwestern 

Mexico. U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, CO.

Marshal, Jason P., Paul R. Krausman, Vernon C. Bleich, [and others]. 2002.

Rainfall, El Niño, and dynamics of mule deer in the Sonoran Desert, Cal

ifornia. Journal of Wildlife Management. 66(4): 1283–1289.

McCain, Emil B., and Jack L. Childs. 2008. Evidence of resident jaguars

(Panthera onca) in the Southwestern United States and the implications

for conservation. Journal of Mammalogy. 89(1): 1–10.

Rowcliffe, J. Marcus, and Chris Carbone. 2008. Surveys using camera traps:

are we looking to a brighter future?Animal Conservation. 11(3): 185–186.

Wallace,Cynthia S.A. 2002.Extracting temporal and spatial information from

remotely sensed data for mapping wildlife habitat: Tucson, The University

of Arizona. Ph.D. Dissertation. 198 p.

White, T. C. R. 2008. The role of food, weather and climate in limiting the abundance of animals. Biological Reviews. 83(3): 227–248.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
506 USDA Forest Service Proceedings RMRS-P-67. 2013

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane 

G.; Collins, Loa C., comps. 2013. Merging science and management in 

a rapidly changing world: Biodiversity and management of the Madrean 

Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. 

Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 

Mapping Landscape Phenology Preference of Yellow-Billed 

Cuckoo With AVHRR Data

Cynthia S. A. Wallace and Miguel Villarreal

U.S. Geological Survey, Western Geographic Science Center, Tucson, Arizona

Charles van Riper, III

U.S. Geological Survey, Southwest Biological Science Center, Sonoran Desert Research Station, Tucson, Arizona 

Abstract—We mapped habitat for threatened Yellow-billed Cuckoo (Coccycus americanus occidentalis) in 

the State of Arizona using the temporal greenness dynamics of the landscape, or the landscape phenology. 

Landscape phenometrics were derived from Advanced Very High Resolution Radiometer (AVHRR) Normalized Difference Vegetation Index (NDVI) data for 1998 and 1999 by using Fourier harmonic analysis 

to analyze the waveform of the annual NDVI profile at each pixel. We modeled the spatial distribution of 

Yellow-billed Cuckoo habitat by coupling the field data of Cuckoo presence or absence and point-based 

samples of riparian and cottonwood-willow vegetation types with satellite phenometrics for 1998. Models 

were validated using field and satellite data collected in 1999. The results indicate that Yellow-billed Cuckoo 

occupy locations within their preferred habitat that exhibit peak greenness after the start of the summer 

monsoon and are greener and more dynamic than “average” habitat. Identification of preferred phenotypes 

within recognized habitat areas can be used to refine habitat models, inform predictions of habitat response 

to climate change, and suggest adaptation strategies. 

Introduction

	 The Yellow-billed Cuckoo (Coccycus americanus occidentalis) is

a neo-tropical migrant bird that travels north from Central and South

America into the southwestern United States to nest during the summer. Riparian forest and woodland nesting habitat of this threatened

species hasdeclined in thewesternUnitedStates overthe past century.

As a result, Western Yellow-billed Cuckoos have been petitioned for

possible listing under the 1973 Endangered Species Act.

	 Riparian habitats contain dense, water-loving deciduous species

including cottonwood (Populus fremontii) and willow (Salix gooddingii)—vegetation preferred by Yellow-billed Cuckoo. In central

and southernArizona, there is a marked contrast between the riparian

vegetation and the surrounding arid and semi-arid upland vegetation

that dominates most of the landscape. This contrast produces strong

vegetation reflectance signalsthat can be detected by satellite remote

sensing both in terms of the strength of the signal due to the difference in vegetation type and density and also in terms of the temporal

pattern of the greenness dynamics, or landscape phenology.

	 The objectives of this study were to identify characteristic phenology patterns preferred by Yellow-billed Cuckoo as captured by

satellite data and use these characteristics to map preferred habitat

of the Yellow-billed Cuckoo in the State of Arizona.

Data

	 TheYellow-billed Cuckoo field data used in the models came from

a state-wide census (Johnson and others 2010). The satellite data

used are the Advanced Very High Resolution Radiometer (AVHRR)

1-kilometer resolution Normalized Difference Vegetation Index

(NDVI) bi-weekly composites. These data are sensitive to biophysical vegetation characteristics, including biomass and percent cover

(Huete and others 2002). The riparian vegetation map was created by

the Arizona Game & Fish Department (AZGF) in 1994 (www.land.

state.az.us). For thisstudy, we created two binary images: all riparian

(RIP) vegetation and riparian Cottonwood-Willow (CW) vegetation.

We excluded any riparian vegetation in the far north and northeast of

the state because no bird sightings were reported in those areas.

Methods

	 Five sets of point data were compiled including 78 survey locations

withYellow-billedCuckoo presentfor 1998; 43 survey locations with

birds absent for 1998; point-sampling polygons of CW vegetation;

point- sampling polygons of RIP vegetation; and 360 bird sightings

for 1999.

	 Phenometrics were derived by applying a Fourier harmonic

analysis to the temporal AVHRR NDVI profiles at each image pixel

in the State of Arizona (Jakubauscas 2001; Wallace 2002). Fourier

analysis expresses a complex curve as the sum of an “additive term”

and simple cosine waves of different frequencies. Three new images each representing a phenometric were created: (1) the amount

of greenness—the “additive term” component that is the best fit of

a flat line to the profile and is related to the vegetation productivity;

(2) dynamics or variability of the greenness—the magnitude of the

first frequency component that is the amplitude of the single cosine
USDA Forest Service Proceedings RMRS-P-67. 2013 507

Mapping Landscape Phenology Preference of Yellow-Billed Cuckoo With AVHRR Data Wallace, Villarreal, and van Riper

wave that fitsthe annual profile best; and (3)the timing of peak greenup—the phase of the first frequency component that is the position

of the peak of the single cosine wave that fits the annual profile best,

rescaled to represent a day of the year (DOY) between 0 and 365.

	 The phenometrics were sampled and summarized for each of

the 5 point data sets. Two-tailed Student’s t-tests were performed

to identify the phenometrics that were significantly different in the

survey locations with Cuckoos present compared to the other point

locations; these phenometrics were used to create Cuckoo phenology preference models. We rescaled the phenometrics based on the

statistical distribution ofthe values atthe set of 1998Cuckoo presence

data to create uniform units of the standard deviation distance (SDD)

by calculating the difference between the phenometric and the mean

value observed at the 1998 Cuckoo sightings divided by the observed

1998 standard deviation.

	 Three models were created, one continuous and two categorical.

For each model, each phenometric wasstratified into three categories

: “high” targeted to contain 50% ofthe sightings, “medium” with 30%

(cumulative 80%), and “low” with 19-20% (cumulative >99%), as

follows:

	 1. Standard Deviation Distance-based Model—The SDDs of

selected Phenometrics were averaged to produce a phenology preference map based on the average SDD.

	 2. Standard Deviation Stratified Model—The SDD of each

phenometric was stratified into three preference classes using the

SDD values of +/- 0.67, +/- 1.28 and +/- 3.0; given a normal distribution, these cutoffs would capture cumulative percentages of 50, 80

and 99+, respectively. The three stratified phenometrics were then

summed, producing a single map with nine classes ranging from 1 =

low preference to 9 = high preference.

	 3.Rank-based Model—TheRankandPercentilewascalculatedfor

each 1998 phenometric at the CW points. The 1998 CW ranks were

identified for the phenometrics at the 50-80-99 SDD values observed

at the 1998 bird sightings; these same ranks were then identified for

the ranked 1999 CW phenometrics, and the corresponding 1999

phenometric values extracted. Each phenometric was stratified into

three preference classes using these extracted values and the three

maps were summed, producing nine classes from 1 = low preference

to 9 = high preference.

	 The models were validated by determining how well they predicted Cuckoo presence in 1999. Refinements of the model based on

observed phenometric relationships within and between years were

also explored and applied.

Results

	 Allthreemodels createdfromphenometricvaluesobservedin1998

were effective at predicting the locations of Yellow-billed Cuckoo

sighted in 1999.We consideredthe standard deviationstratifiedmodel

(fig. 1A) superior given it identified a less generalized mapping and

predicts 64-98-100% cumulatively of the 1999 bird sightings in the

High(greens)-Medium(orange)-Low(darkgray)classes,respectively.

The model was then clipped to probable landscapes, from most to

least limited with each a subset of the next (figs. 1B, C, D).

	 Examination of the statistics and histograms for the locations with

Birds Present and the CW vegetation types shows the phenometrics

Figure 1—Standard Deviation Stratified Model, showing High-Medium-Low phenology preference classes. View B is clipped to CW 

vegetation, view C is clipped to RIP vegetation, and view D is clipped to the Arizona Streams Data, digitized from 1988 1:100,000 USGS 

topographic maps. The insets show the 1999 bird locations overlaid onto the clipped models.
508 USDA Forest Service Proceedings RMRS-P-67. 2013

Wallace, Villarreal, and van Riper Mapping Landscape Phenology Preference of Yellow-Billed Cuckoo With AVHRR Data

of these landscapes share parallel relationships in 1998 and 1999.

In both years, birds occupy landscapes that have a more variable,

dynamic greenness, achieve a maximum greenness later in the year,

and are significantly greener than the “average” CW vegetation. We,

therefore, refined the model by averaging the observed intra-annual

phenometricsoffsets andcreatedgeneralizedmodelsforboth1998and

1999 that defineYellow-billedCuckoo habitatrelative to the observed

phenology of the CW vegetation. These models correctly predict in

High-Medium-Low classes cumulative bird presence percentages of

57-99-100 and 81-99-100, for 1998 and 1999, respectively.

Discussion

	 We found thatYellow-billed Cuckoo occupy locations within their

preferred habitat that green up later in the year, have a more dynamic

greenness and are greener than the average cottonwood-willow habitat. These could be areas of healthier and denser trees or could be a

vegetation signature in the understory that reflect conditionssupporting larger insects—a food that the Yellow-billed Cuckoos favor and

exploit. Phenology preference models were effective at predicting

Cuckoo sightings in both years, and could be refined as additional

data become available. These models are successful because there is

a consistent relationship between the intra-annual phenometrics for

Cuckoosightingsrelativetothoseintheirpreferredcottonwood-willow

habitat. It is likely that other species will exhibit similar phenotype

preferences within their habitat types. Such relationships can be used

to refine habitat models, predictspecies presence in other areas of the

Southwest and better inform habitat response to climate change.

References

Huete, A.; Didan, K.; Miura, T. 2002. Overview of the radiometric and biophysical performance of the MODIS Vegetation Indices. Remote Sensing

of Environment 83:195–213.

Jakubauskas, M.E.; Legates, D.R.; Kastens, J. 2001. Harmonic analysis of

time-seriesAVHRRNDVIdata.PhotogrammetricEngineeringandRemote

Sensing 67(4): 461-470.

Johnson, M.J.; Magill, R.T.; van Riper, C., III. 2010. Yellow-billed Cuckoo

distribution and habitat associations in Arizona, 1998-1999. Pp. 197-212,

In: The Colorado Plateau IV: Shaping conservation through science and

management. van Riper, C., III; Wakeling, B.F.; Sisk, T.D., eds. Tucson,

AZ: University of Arizona Press. 335 pp.

Wallace, C.S.A. 2002. Extracting temporal and spatial information from

remotely sensed data for mapping wildlife habitat. Tucson: The University

of Arizona. Ph.D. Dissertation. 198 p.

The content of this paper reflects the views of the authors, who are responsible for the facts and accuracy of the information presented herein.
USDA Forest Service Proceedings RMRS-P-67. 2013 509

Abstracts
510 USDA Forest Service Proceedings RMRS-P-67. 2013
USDA Forest Service Proceedings RMRS-P-67. 2013 511

In: Gottfried, Gerald J.; Ffolliott, Peter F.; Gebow, Brooke S.; Eskew, Lane G.; Collins, Loa C., comps. 2013. Merging science and management in a rapidly 

changing world: Biodiversity and management of the Madrean Archipelago III; 2012 May 1-5; Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station. 

Abstracts of Oral and Poster Presentations at the Conference: English and Spanish 

(Some titles and authors are different from the finalized versions)

Resúmenes en Inglés y Español de las Presentaciones Orales y en Cartel del 

Congreso (Alugunos títulos y autores son diferentes al de la versión final)

AUSTIN, VALER, David Hodges, and Cindy Tolle, Cuenca Los Ojos A.C. Agua Prieta, Sonora México and Cuenca Los Ojos Foundation,

12626 E. Turkey Creek Road, Pearce AZ 85625; vaustin@elcoronadoranch.net.

Restoration of Ciénegas and Headwater Tributaries of the Río San Bernardino. The Cuenca Los Ojos Foundation restores lands, wetlands,

and wildlife in Arizona and Sonora. This semi-arid area receives limited rainfall in a given year with extreme fluctuations, sometimes receiving

up to 45% of the annual rainfall in one event. Many streams are seasonal and depending upon the watershed, can go from dry to flooding in an

hour’s time. Harvesting water on both slopes and drainages is therefore crucial to achieving maximum conservation outcomes. Historically,

a large ciénega wetland was located where Black Draw, Hay Hollow, and Silver Creek came together to form the Río San Bernardino, just

south of the International Border. The wetlands were a major stopping place for migrating birds and animals. In the early 1900’s these wetlands

dried due to incision of the adjacent streams. Cuenca Los Ojos has been working for more than a decade to restore these streams and adjacent

upland areas. Despite historic drought conditions during the entirety of this restoration period, our work has resulted in a dramatic increase in

the ciénega acreage and riparian and grassland vegetation. This in turn, has led to an increase in wildlife such as native fish, frogs, riparian and

grassland bird species, and a host of mammals.

Restauración de ciénegas y afluentes de la cuenca alta del río San Bernardino. La fundación de la Cuenca los Ojos trabaja para restaurar

terrenos, humedales, fauna y flora en Arizona y Sonora. Esta región semiárida tiene muy baja precipitación con una variabilidad extrema, en

ocasiones recibiendo hasta el 45% de la precipitación anual en un chubasco. Muchos arroyos son temporales y dependiendo de la cuenca, pueden cambiar en una hora de secos a inundados. Así, el recolectar el agua de lluvia tanto de las laderas como de las vertientes es decisivo para

lograr el máximo resultado en conservación. En el pasado un gran humedal de ciénega se ubicaba donde se juntan los afluentes Black Draw,

Hay Hollow y Silver Creek para formar el río San Bernardino, justo al sur de la frontera internacional. Estos humedales fueron un importante

refugio para aves y otros animales migratorios. A principios de 1900 estas ciénegas se secaron por la fragmentación causada por los arroyos

adyacentes. La Cuenca los Ojos ha estado trabajando por más de una década restaurando estos arroyos y las tierras altas adyacentes. A pesar

de las condiciones de sequía histórica durante estos años de restauración, hemos logrado el aumento dramático de la ciénega y la vegetación

ribereña y el pastizal. Ésto a su vez a creado un aumento de fauna nativa como son: peces nativos, ranas, aves ribereñas y de pastizal y una

gran variedad de mamíferos.

AVILA-VILLEGAS, SERGIO1; Jessica Lamberton2; Jeannette P. Hanby3; J. David Bygott4. 1M.S. Sky Island Alliance, 300 E. University Blvd.,

Tucson,AZ 85705,sergio@skyislandalliance.org; 2Sky IslandAlliance, 300 E. University Blvd., Tucson,AZ 85705, jessica@skyislandalliance.

org; 3 Ph D. Visiting scholar U. of Arizona Anthropology Department; Sky Island Alliance volunteer, kibuyu@yahoo.com; 4Ph D. Sky Island

Alliance volunteer, davidbygott@yahoo.com.

Wildlife survey and monitoring in the Sky Island Region with an emphasis on Neotropical felids.The Sky Island region of southwestern

U.S. and northwestern Mexico consists of isolated mountain ranges separated by deserts and grasslands. It mixes elements from four major

ecosystems - the Rocky Mountains, Sierra Madre Occidental and the Sonoran and Chihuahuan deserts, and Neotropical species reaching their

northern ranges, such as jaguars (Panthera onca) and ocelots (Leopardus pardalis). Sky Island Alliance seeks to build cooperative relationships

with landowners in Sonora and public land managers in Arizona to facilitate scientific research, encourage large predator conservation and

establish corridors connecting patches of continuous habitat throughout the region. We have conducted wildlife surveys with remote cameras in

northern Sonora since 2007 and in Arizona since 2009. Our photographic records cover 23 species of wild mammals, including jaguar, ocelot,

puma (Puma concolor), black bear (Ursus americanus), coati (Nasua narica) and four species of skunks (Mephitidae). In 2009 we documented

Arizona’s first live ocelot, and recently we documented two individual jaguars 30 miles south of the border in Sonora. Our records reveal spatial

and temporal activity patterns of wildlife species, their distribution, habitat preference, and corridor use. The project’s results have increased

our knowledge of borderland wildlife, especially the Sonoran ocelot, and supported the certification of a 10,000-acre private ranch as a wildlife

preserve in northern Sonora.
512 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Evaluación y monitoreo de fauna en la Región de las Islas Serranas con énfasis en felinos neotropicales. La región de las Islas Serranas del

noroeste de México y suroeste de Estados Unidos consiste de montañas aisladasseparadas por desiertos y pastizales. La región une elementos de

cuatro ecosistemas –la Sierra Madre Occidental, las Montañas Rocallosas, los desiertos Sonorense y Chihuahuense y especies Neotropicales que

alcanzan aquí su distribución más norteña, como jaguares (Panthera onca) y ocelotes (Leopardus pardalis). Sky Island Alliance busca construir

relaciones de cooperación con propietarios privados en Sonora y encargados de áreas naturales enArizona para facilitar investigación científica,

fomentar la conservación de grandes carnívoros y establecer corredores que conecten hábitats continuos en la región. En este proyecto hemos

realizado evaluaciones de fauna con cámaras remotas en el norte de Sonora desde 2007 y enArizona desde 2009. Nuestros registros fotográficos

incluyen 23 especies de mamíferos silvestres, como el jaguar, ocelote, puma (Puma concolor), oso negro (Ursus americanus), coati (Nasua 

narica) y cuatro especies de zorrillos (Mephitidae). En 2009 documentamos el primer ocelote vivo en el estado de Arizona y recientemente

documentamos dos jaguares distintos a 45 kilómetros al sur de la frontera en Sonora. Nuestros registros revelan patrones de actividad de fauna

espaciales y temporales, su distribución, preferencia de hábitat y uso de corredores. Los resultados del proyecto han incrementado nuestro

conocimiento sobre la fauna en la región fronteriza, especialmente de la subespecie de ocelote Sonorense y respaldan la certificación de una

reserva privada de 4,000 hectáreas para fauna en el norte de Sonora.

AVILA-VILLEGAS, SERGIO1; Caroline Patrick-Birdwell2; Jenny Neeley3; Louise Misztal4; Nicholas Deyo5. 1M.S. Sky Island Alliance, 300

E. University Blvd., Tucson, AZ 85705, sergio@skyislandalliance.org; 2M.S. The University of Arizona, School Geography and Development,

453E Harvill Building, Box #2, Tucson,AZ 85721, cpatrick@email.arizona.edu; 3J.D. Sky IslandAlliance, 300 E. University Blvd., Tucson,AZ

85705, jenny@skyislandalliance.org; 4Sky Island Alliance, 300 E. University Blvd., Tucson, AZ 85705, louise@skyislandalliance.org; 5M.L.A.

Sky Island Alliance, 300 E. University Blvd., Tucson, AZ 85705, nick@skyislandalliance.org.

Mapping and assessing the environmental impacts of border tactical infrastructure in the Sky Island Region. In this project we mapped

the different types of border wall, identified impacts of border infrastructure on public and private lands and conducted spatial analyses within

the approximately 200 miles of international border in the Sky Island region. The Sky Island region, bisected by the U.S.-Mexico border, is

critically important for its biodiversity and continental connectivity; only on the Arizona side the border is lined with protected areas such as

San Bernardino and Buenos Aires National Wildlife Refuges, San Pedro Riparian National Conservation Area, Coronado National Memorial,

and several Wilderness Areas. In recent years the construction of border barriers, walls and other tactical infrastructure built to deter human

and vehicle traffic across the border has had substantial impacts on ecological processes and created an impermeable boundary with effects

that reach far beyond the footprint of the wall. Due to the lack of environmental impact studies, construction of border infrastructure proceeded

without input from public land managers or private landowners. With the use of Geographic Information Systems we analyzed and graphically

compiled results from over-flights along the border generating photographic data of the current state and effects of infrastructure, field visits to

obtain complementary information and ground-truth aerial photographs, and observations of the direct effects of barriers to wildlife movements

and the flow of ecological and hydrologic processes.

Trazado de mapas y evaluación de los impactos ambientales de la infraestructura táctica fronteriza en la Región de las Islas Serranas.

En este proyecto se trazaron mapas de los diferentes tipos de barreras fronterizas, se identificaron impactos de la infraestructura en áreas del

gobierno y privadas, y se realizaron análisis espaciales en cerca de 320 kilómetros de frontera internacional en la Región de las Islas Serranas.

La Región de lasIslas Serranas, dividida por la frontera internacional México-Estados Unidos, esimportante porsu biodiversidad y conectividad

paisajística a nivel continental; solo en Arizona la frontera esta delineada por áreas naturales protegidas como los Refugios de Fauna Silvestre

San Bernardino y Buenos Aires, el Área Nacional de Conservación del Río San Pedro, el Coronado National Memorial, y otras zonas naturales. En años recientes la construcción de barreras, muros y otra infraestructura táctica fronteriza construida para detener el paso de personas y

vehículos a través de la frontera ha tenido impactos substanciales en los procesos ecológicos y ha creado una barrera impermeable con efectos

que llegan más allá de la huella del muro. Debido a la falta de estudios de impacto ambiental, la construcción de infraestructura fronteriza ha

procedido sin la participación de encargados de áreas naturales o propietarios privados. Con el uso de sistemas de información geográfica se

analizan y representan gráficamente los resultados de vuelos en la frontera generando información fotográfica del estado actual de la infraestructura, visitas de campo para recabar información complementaria, verificar fotografías aéreas, y observar los efectos directos de las barreras

sobre los movimientos de fauna y el flujo de procesos ecológicos e hidrológicos.

BACKER, DANA, and Perry Grissom. Saguaro National Park, 3693 S. Old Spanish Trail, Tucson, Arizona, dana_backer@nps.gov, perry_grissom@nps.gov.

Perennial invasive grass fires threaten to convert the Sonoran Desert ecosystem. Pyrophilic buffelgrass (Pennisetum ciliare) is invading

many ecosystems in the United States and abroad, including the Sonoran Desert. Buffelgrass, an African bunch grass, poses a significant threat

to Saguaro National Park’s natural and cultural resources.Abuffelgrass-fueled fire islikely to cause unprecedented and irreparable damage to the

desert ecosystem. Iconic Sonoran Desert species such as the saguaro cactus and desert tortoise are not fire-adapted and will suffer fire-induced

mortality from buffelgrass fueled fires. Buffelgrass will contribute to a positive grass-fire cycle with the potential to convert the Sonoran Desert

ecosystem into exotic grassland. Land managers in the Sonoran Desert face novel challenges beyond large scale control treatments, including

developing restoration techniques, fire management strategies, and post-fire treatments. The Park has been managing buffelgrass for more than

10 years; we will present fire behavior results, economical and logistical issues of controlling buffelgrass in a wilderness, and lessons learned.
USDA Forest Service Proceedings RMRS-P-67. 2013 513

Abstracts/ Resúmenes

Incendios de zacates perennes invasores amenazan cambiar el ecosistema del Desierto Sonorense. El zacate buffel amante del fuego (Pennisetum ciliare) está invadiendo muchos ecosistemas en Estados Unidos y el extranjero, incluso el Desierto Sonorense. El buffel, un zacate

africano que crece en macollos, representa una amenaza seria para los recursos naturales y culturales del Parque Nacional Saguaro. Un incendio

de buffel puede causar un daño sin precedentes e irreparable al ecosistema del desierto. Las especies emblemáticas del Desierto Sonorense,

como el sahuaro y la tortuga de los cerros no están adaptados al fuego y sufrirán una mortandad inducida por el fuego de incendios del buffel. El

buffel inducirá un ciclo de zacate-fuego con el potencial de convertir el ecosistema del Desierto Sonorense a un pastizal exótico. Los encargados

del manejo de recursos naturales del Desierto Sonorense enfrentan nuevos retos además del control en gran escala como son: el desarrollo de

técnicas de restauración, estrategias de manejo del fuego, y tratamientos posteriores al incendio. El Parque ha controlado el zacate buffel por

más de 10 años; se presentarán resultados de comportamiento del fuego, problemas económicos y logísticos para controlar buffel en un área

naturals, y las lecciones aprendidas.

BALLARD, GRANT1, Carol Beardmore2, Elva Ivonne Bustamante-Moreno3, Jennifer N. Duberstein2, Nathan Elliot1, Geoff Geupel1, Dennis Jongsomjit1

, Robert Mesta2, Leo Salas3, and Sam Veloz3. 1PRBO Conservation Science, 3820 Cypress Drive #11, Petaluma, CA 94954.

prbo@prbo.org; 2Sonoran Joint Venture, 738 North 5th Ave., Ste. 102, Tucson, AZ 85705, jennie_duberstein@fws.gov; 3CONANP (Mexican

National Commission for Protected Areas), Camino al Ajusco No. 200, Col. Jardines en la Montaña, Deleg. Tlalpan, C.P. 14210, México D.F.,

ebustama@conanp.gob.mx.

Detecting climate change impacts on birds and their habitats in the southwest United States and northwest México. Failing to incorporate the projected impacts of climate change into habitat management strategies will almost certainly lead to a failure to achieve conservation

objectives. Changes in bird populations have long been used to indicate the direction and magnitude of change in ecosystems, and they can help

us assess current and future impacts of climate change. This information can then be used by managers to take appropriate conservation action

for birds and bird habitat and can help focus investment in overlap between current and future conservation priorities. The goal of this project

is to develop a foundation for monitoring environmental change in the Sonoran Joint Venture region. We provide land managers and biologists

with methodological tools, data resources, analyses, and training to understand and detect climate change effects, define adaptation options, and

increase capacity for making conservation decisions for bird populations and habitats in the areas under their jurisdiction. We use climate and

bird observation data to identify locations where we predict the greatest changes in climate, habitats, and bird communities. During a workshop

with experts, we will use the locations identified by our analyses to define priority bird species, sites, and regions and develop a coordinated

bird monitoring strategy for the region with respect to climate change. We will also develop a bilingual web portal to view predicted changes

and store and query data, and provide training opportunities for managers and biologists on the use of these resources, monitoring strategies,

and data collection and management.

Cómo detectar los impactos del cambio climático en aves y sus hábitats en el suroeste de Estados Unidos y noroeste de México. Si no se

incluyen los impactos previstos del cambio climático en las estrategias de manejo del hábitat muy probablemente no se lograrán los objetivos

de conservación. Los cambios en las poblaciones de aves se han usado desde hace tiempo para indicar la dirección y magnitud del cambio en

los ecosistemas y pueden ayudar a evaluar los impactos actuales y futuros del cambio climático. Esta información puede ser útil al personal de

manejo de recursos para tomar decisiones de conservación para las aves y sus hábitats, y a dar prioridad a la inversión donde coincidan preferencias actuales y futuras de conservación. Este proyecto tiene como meta sentar las bases para el monitoreo del cambio ambiental en la región

Sonoran Joint Venture. Se brinda a los encargados de manejo y biólogos metodología, fuentes de datos, análisis y capacitación para entender

y detectar los efectos del cambio climático, identificar adaptaciones y aumentar su capacidad para tomar decisiones de conservación para las

poblaciones de aves y sus hábitats en las áreas bajo su jurisdicción. Utilizamos datos climáticos y observaciones de aves para identificar las

localidades donde pronosticamos los cambios mayores en clima, hábitat y comunidades de pájaros. En un taller con especialistas, usaremos las

localidades identificadas en nuestros análisis para determinar las especies de aves, sitios y regiones prioritarias y planear una estrategia coordinada para el monitoreo de aves en la región con relación al cambio climático. También diseñaremos un portal de internet bilingüe para ver los

cambios previstos, subir y guardar información y consultas. Así mismo tendremos oportunidades de capacitación para el personal de áreas de

conservación y biólogos sobre el uso de estos recursos, estrategias de monitoreo, toma de datos y manejo.

BENTON, MARY BETH1, Siria A. Cerda-Navarro2, Katie R. Keck4 and Brittany N. McKnight3. 1Saguaro National Park, Tucson, AZ 85730.

benton.marybeth@gmail.com; 2School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721, siriac@email.

arizona.edu, 3bmcknigh@email.arizona.edu; 4Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721,

khooker@email.arizona.edu.

Spatial and temporal analysis of predator-prey interactions within Saguaro National Park. FromApril 2011 toApril 2012, we investigated

the spatial and temporal patterns of mammals within Saguaro National Park’s Rincon and Tucson Mountain Districts. Data was analyzed using

Cuddeback Capture camera images provided by Saguaro National Park. The 91,440 acre park was divided into 1-km2 plots ranging in elevation

from 664 m to 2,641 m using a grid feature of ArcMap. The location of the plots as well as the camera placement in each plot was randomly

chosen using a number generator in ArcMap. Over a twelve-month period, images from 126 cameras were analyzed. The location and time of

mammal images were recorded and evaluated and correlations of predator-prey interactions were examined based on these factors. Our findings suggest that the mammal community is successfully coexisting both temporally and spatially within Saguaro National Park. Knowledge

of how mammals utilize the landscape of Saguaro National Park will enable wildlife managers to implement more effective conservation and

management strategies.
514 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Análisis espacial y temporal de las interacciones entre depredador y presa del Parque Nacional Saguaro. Se hizo un estudio de los patrones espaciales y temporales de mamíferos en los distritos Tucson Mountain y Rincon del Parque Nacional Saguaro, desde abril de 2011 a

abril de 2012. La información se analizó usando imágenes de cámaras ‘Cuddeback Capture’ del Parque. El área del parque (370 has) se dividió

en parcelas de 1-km2 variando en elevación de 664 m a 2,641 m usando una cuadrícula de ArcMap. La ubicación de las parcelas, así como la

colocación de la cámara en las mismas, fue seleccionada al azar usando un generador de números de ‘ArcMap’. Se analizaron las imágenes de

126 cámaras generadas en un período de 12 meses. La ubicación y tiempo de las imágenes de mamíferos se capturaron y analizaron, y basado

en estos factores se examinó las interacciones entre el depredador y la presa. Nuestros resultados sugieren que la comunidad de mamíferos tiene

una coexistencia exitosa tanto espacialmente como temporalmente en el Parque Nacional Saguaro. El saber cómo los mamíferos utilizan el área

del Parque permitirá que los encargados de manejo de fauna implementen estrategias de conservación y manejo más efectivas.

BERMÚDEZ-ENRÍQUEZ, ERICK OSWALDO1, Rosa Elena Jiménez-Maldonado2, Gertrudis Yanes-Arvayo1 and Maria De La Paz MontañezArmenta1

. 1Universidad de la Sierra. División de Ciencias Biológicas, Moctezuma, Sonora; 2Reserva Forestal Nacional y Refugio de Fauna

Silvestre Ajos Bavispe, Cananea, Sonora.

Monitoring of medium and large mammals in the Sierra La Madera, Sonora, Mexico. Sierra La Madera is part of the group of mountains

called “Sky Islands” Madrean Archipelago and comprises the Reserve Fraction V “ Ajos-Bavispe”. The objective was to determine the composition of medium and large mammals of wildlife in the area, through the use of camera traps. Were placed 18 camera traps WildView 2®

brand in three monitoring campaigns: August-September, September to November and November-December. The first and second campaign

was placed cameras on the farms La Bellota, La Palmita y San Fernando, the third campaign was placed on ranches La Mesa, El Mezquite y

Brecha CONAFOR, located in the municipalities of Cumpas, Montezuma, and Huásabas y Villa Hidalgo, where there is thorny scrubs, oak

and pine forest. The results show 17 species in five orders Carnivora, Artiodactyla, Didelphimorphia, Lagomorpha and Rodentia. Additionally,

there was four species of birds and animals.

Monitoreo de mamíferos medianos y grandes en Sierra La Madera, Sonora, México. Sierra La Madera forma parte del conjunto de sierras

denominadas “Islas Serranas” del Archipiélago Madrense y comprende la Fracción V de Reserva “Ajos-Bavispe”. El objetivo fue determinar

la composición de fauna silvestre de mamíferos medianos y grandes en el área, a través del uso de cámaras-trampa. Se colocaron 18 cámarastrampa marca WildView 2® en tres campañas de monitoreo: agosto-septiembre, septiembre-noviembre y noviembre- diciembre. La primera

y segunda campaña se colocó cámaras en los ranchos La Bellota, La Palmita y San Fernando; la tercera campaña se colocó en los ranchos La

Mesa, El Mezquite y Brecha CONAFOR, estos ubicados en los municipios de Cumpas, Moctezuma, Huásabas y Villa Hidalgo, donde existe

matorral espinoso, bosque de encino y pino. Los resultados muestran 17 especies en cinco órdenes Carnívora, Artiodactyla, Didelphimorphia,

Lagomorpha y Rodentia. Adicionalmente se registró cuatro especies de aves, así como animales domésticos.

BERTELSEN, C. DAVID. Herbarium, University of Arizona, david.bertelsen8@gmail.com.

Conceptual model of plant responses to climate and implications for monitoring for ecosystem change. Although land management agencies have expressed increasing concern for monitoring for ecosystem health and resiliency in a changing climate, current practices, designed

primarily to monitor for erosion and impacts of livestock and focusing on a small number of the most resilient (and economically desired)

species, are inadequate to assess early change in vegetative communities or ecosystems. Optimal and adverse climatic conditions result in different vegetative assemblages. Ecosystem change will be more readily seen if the focus is on those species that respond positively to optimal

conditions but are absent or non-reproductive in adverse conditions: what isn’t present may be more important than what is. The conceptual

model is a starting point for determining how land managers can identify ecosystem change before it reachesthe tipping point ofsystem collapse.

Modelo conceptual de las respuestas de plantas al clima y sus implicaciones en el monitoreo de cambios en el ecosistema. Aunque las

agencias de manejo de recursos naturales han expresado una mayor preocupación por el monitoreo del equilibrio ecológico y la resistencia del

ecosistema en un clima cambiante, las prácticas actuales diseñadas principalmente para el monitoreo de la erosión y los impactos del ganado y

centrándose en unas cuantas de las especies más resistentes (y económicamente deseables) son inadecuadas para evaluar los primeros cambios

en las comunidades vegetales o ecosistemas. Las condiciones climáticas adversas y óptimas producen diferentes agrupaciones de plantas. El

cambio en el ecosistema se podrá ver más fácilmente si el trabajo se enfoca en las especies que responden de forma positiva a las condiciones

óptimas, pero están ausentes o sin reproducir en condiciones adversas: las ausentes pueden ser más importantes que las presentes. El modelo

conceptual es el punto de partida para ayudar a los encargados de manejo a identificar el cambio en el ecosistema antes de alcanzar el momento

crítico del desplome del sistema.

BODNER, GITA1, Dave Gori2, Peter Warren1, and Karla Sartor2. 1The Nature Conservancy in Arizona, 1510 E Ft Lowell Road, Tucson, AZ

85719, gbodner@tnc.org, pwarren@tnc.org; 2The Nature Conservancy in New Mexico, 212 E. Marcy St., Suite 200, Santa Fe, NM 87501,

dgori@tnc.org, ksartor@tnc.org.

Sustaining the grassland sea: Regional perspectives on restoring and protecting the Sky Island region’s most intact grassland valley 

landscapes. Grasslands of the Sky Islands Region once covered over 11 million acres in southeastern Arizona and adjacent portions of New

Mexico, Sonora, and Chihuahua. Attempts to evaluate current ecological conditions suggest that approximately two third of this remains as

intact or restorable grassland habitat. These grasslands provide watershed services such as flood control and aquifer recharge across the region,

and continue to support dozens of species of concern. Prioritizing conservation interventions for these remaining grassland blocks has been

challenging. Reliable data on condition and conservation value of grasslands in the region have not been systematically summarized. State and
USDA Forest Service Proceedings RMRS-P-67. 2013 515

Abstracts/ Resúmenes

national boundaries further complicate efforts to identify where the best remaining habitats and populations of grassland obligate wildlife still

exist. We present results of an effort to merge grassland condition assessmentsfrom adjacent geographies, compile information on targetspecies

locations, and identify “priority grassland valley landscapes” across the region. We evaluate these priority landscapes in terms of the number of

targetspecies, critical threats, and enabling conditionsfor long-term conservation successsuch as activity by local cooperative groups dedicated

to sustaining their landscapes. Lastly, we discuss the opportunities and challenges of designing and implementing effective restoration activities

in these large multi-jurisdictional landscapes.

Conservación del mar de pastizales: Perspectivas regionales sobre la restauración y protección del paisaje de pastizal más intacto de 

la región de las Islas Serranas. Los pastizales de la región de las Islas Serranas, años atrás, cubrieron más de 4.5 millones de hectáreas en el

sureste de Arizona y partes adyacentes de Nuevo México, Sonora y Chihuahua. Las evaluaciones de las condiciones ecológicas actuales indican

que aproximadamente dos terceras partes están intactas o son hábitat de pastizales recuperables. Estos pastizales proporcionan servicios de

cuencas, tales como control de inundaciones y recarga de los acuíferos en la región, y mantienen a decenas de especies de interés. Priorizar las

intervenciones para la conservación de estos bloques restantes de pastizales ha sido un desafío. Datos confiables sobre la condición y valor para

la conservación de los pastizales de la región no han sido sistemáticamente condensados. Fronteras estatales y nacionales, complican aún más

los esfuerzos para identificar dónde quedan los mejores hábitats y las poblaciones de vida silvestre dependiente de los pastizales que aún existen.

Presentamoslosresultados de un estudio para incorporarlas evaluaciones de la condición de los pastizales geográficamente adyacentes,recopilar

información sobre la ubicación de especies de interés, e identificar “paisajes de llanuras de pastizal prioritarios “ de la región. Evaluamos estos

paisajes prioritarios en términos del número de especies importantes, amenazas críticas y condiciones propicias para el éxito de la conservación a largo plazo como la participación de grupos locales colaborativos dedicados a mantener sus áreas naturales. Por último, analizamos las

oportunidades y retos de diseño e implementación de medidas de restauración efectivas en estas grandes áreas naturales de variasjurisdicciones.

BODNER, GITA1, Linda Kennedy2, Karen Simms3, Jeff Williamson4. 1The Nature Conservancy, Tucson, AZ, gbodner@TNC.ORG ; 2National

Audubon Society, Elgin, AZ, lkennedy@audubon.org; 3Bureau of Land Management, Tucson, AZ. ksimms@blm.gov; 4Cienega Watershed

Partnership, Phoenix, AZ, JWilliamson@thephxzoo.com.

Science on the Sonoita Plain. The Sonoita Valley Planning Partnership (SVPP) is a voluntary ad hoc association of agencies, user groups,

conservation organizations, and individuals who come together to achieve community-oriented solutions to local and national issues affecting

public lands within the Sonoita Valley. The SVPP was created in 1995 in response to BLM's initiation of a collaborative planning process for

Las Cienegas National Conservation Area. The SVPP meets quarterly and provides a forum for participants to share information and work

together to perpetuate naturally functioning ecosystems while preserving the rural, grassland character of the Sonoita Valley for future generations. Science on the Sonoita Plain symposiums are held annually to bring together and share results of scientific investigations occurring within

and informing us about the unique and diverse resources of the Sonoita Plain in the upper watersheds of Cienega Creek, Sonoita Creek, and

the Babocomari River. The third annual symposium was held on June 4, 2011; the next is scheduled for June 9, 2012. While several groups in

the region organize lectures, just a few host annual events designed to concentrate scientists, community members, and agency staff to learn

together about their local area. We welcome attendees and recommend this approach for other local areas.

Ciencia en la llanura de Sonoita. The Sonoita Valley Planning Partnership (SVPP) es una asociación voluntaria de agencias, grupos de

usuarios, organizaciones de conservación e individuos que se formó para buscar soluciones dirigidas a la comunidad sobre temas locales y

nacionales que afectan las áreas naturales del valle de Sonoita. La SVPP fue creada en 1995 como respuesta al proceso colaborativo del BLM

para planear el Área Nacional de Conservación de Las Ciénegas. La SVPP se reúne trimestralmente y es un foro donde los participantes comparten información y trabajan juntos para conservar los ecosistemas naturales y preservar el carácter de pradera rural del valle de Sonoita para

las generaciones futuras. Los simposios de ciencia de la llanura de Sonoita se realizan cada año para compartir resultados de investigaciones

científicas e información sobre la diversidad de los recursos naturales de la llanura de Sonoita en la cuenca alta de Cienega Creek, Sonoita

Creek, and the Babocomari River. El tercer simposio anual se llevó a cabo el 4 de junio 2011; el próximo está programado para el 9 de junio

2012. Aunque varios grupos de la región organizan conferencias, sólo unos ofrecen eventos anuales diseñados para unir a científicos, miembros

de la comunidad y personal de las agencias para juntos informarse sobre su zona local. Invitamos a que nos acompañen y recomendamos este

enfoque para otras regiones locales.

BOERSMA, KATE S.1, Michael T. Bogan2, and David A. Lytle3. 1Department of Zoology, Oregon State University, Corvallis, OR 97331, kate.

boersma@science.oregonstate.edu; 2Department of Zoology, Oregon State University, Corvallis, OR 97331, boganmi@science.oregonstate.

edu; 3Department of Zoology, Oregon State University, Corvallis, OR 97331, lytleda@science.oregonstate.edu.

Top predator extinctions in drying streams modify community structure and ecosystem functioning. Climate change and water withdrawals in the Madrean Sky Island region are causing many once-perennial streams to fragment or dry completely. One of the most notable changes

associated with this transition is the disappearance of large aquatic predators, such as fish and the invertebrate top predator Abedus herberti

(Hemiptera: Belostomatidae). These extinctions presumably cause changes in stream community structure and functioning, however manipulative experiments are necessary to disentangle the effects of local predator extinctions from other co-occurring environmental changes. We

seeded 16 mesocosms with stream invertebrates, removed Abedus from 8 mesocosms as a treatment, and assessed community and ecosystem

changes after 6 weeks. We found that the removal of Abedus initiated a trophic cascade resulting in decreased algal biomass, likely caused by the

release of the algae-grazing mayfly Callibaetis from Abedus predation pressure. Abedus removal also affected overall invertebrate community

structure. The abundance and diversity of secondary predators were higher in mesocosms without Abedus, suggesting a competitive release
516 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

of these smaller predators. Our findings demonstrate that the indirect biotic effects of stream drying can be as important as abiotic effects and

must be considered when planning aquatic conservation in the region.

La extinción de los depredadores principales en riachuelos que se están secando modifica la estructura de la comunidad y funcionamiento del ecosistema. El cambio climático y la extracción de agua en la Región de las Islas Serranas Madrense están causando que muchos

arroyos perennes se fragmenten o se sequen por completo. Uno de los cambios más notables asociados con esta transición es la desaparición

de grandes depredadores acuáticos, como peces y el depredador principal de los invertebrados Abedus herberti (Hemiptera: Belostomatidae).

Estas extinciones probablemente causan cambios en el funcionamiento y la estructura de la comunidad del riachuelo, sin embargo, hacen falta

experimentos de manipulación para separar los efectos de la extinción de los depredadores locales de otros cambios ambientales concurrentes. Una muestra de invertebrados de arroyos se introdujo en 16 mesocosmo, en ocho de los cuales se extrajo Abedus como tratamiento, y se

evaluaron cambios en la comunidad y el ecosistema después de 6 semanas. La eliminación de Abedus inició una cascada trófica que resultó en

la disminución de la biomasa de algas, probablemente causado por la falta del depredador Abedus sobre la mosca del agua Callibaetis que se

alimenta de algas. La eliminación de Abedus también afectó la estructura general de la comunidad de invertebrados. La abundancia y diversidad

de depredadores secundarios fueron mayores en los mesocosmos sin Abedus, lo que sugiere menos competencia entre estos depredadores más

pequeños. Nuestros hallazgos demuestran que los efectos bióticos indirectos de riachuelos que se están secando pueden ser tan importantes

como los efectos abióticos y deben ser considerados en la planificación de la conservación acuática en la región.

BOGAN, MICHAELT.1, Oscar G. Gutierrez2,J.AndrésAlvarado3, and DavidA. Lytle4. 1Department of Zoology, Oregon State University, Corvallis, OR, boganmi@science.oregonstate.edu; 2

Centro de Estudios Superiores Del Estado De Sonora, Hermosillo, Sonora, México, ruachogr@

hotmail.com; 3 Centro de Estudios Superiores del Estado de Sonora, Hermosillo, Sonora, México, zulu_1237@hotmail.com; 4 Department of

Zoology, Oregon State University, Corvallis, OR, david.lytle@science.oregonstate.edu.

Habitat type and permanence determine local aquatic invertebrate community structure in the Madrean Sky Islands. Aquatic habitats

in the Madrean Sky Islands (MSI) consist of a matrix of perennial and intermittent stream segments, seasonal rain ponds, and artificial cattle

tank habitats that all contribute to the regional biodiversity of aquatic macroinvertebrates. Although environmental conditions and aquatic

communities are generally quite distinct in lotic and lentic habitats, MSI streams are characterized by isolated perennial pools for much of the

year, and thus seasonally approximate lentic environments. In this study, we compared habitat characteristics and Coleoptera and Hemiptera

assemblages of stream pools with those of true lentic habitats (seasonal ponds and cattle tanks) across the MSI. We identified 150 species across

the 38 study sites, with distinct faunas found in stream pools versus seasonal ponds. Stream-exclusive species included many long-lived and

poor dispersing species, while pond-exclusive species tended to have rapid development times and good dispersal abilities. At a smaller set of

streams, we also compared aquatic invertebrate community structure between perennial and intermittent streams. We found lower diversity in

intermittent streams, but most intermittent stream species were specialists not found at perennial sites. We suggest that, in addition to perennial

streams, seasonal aquatic habitats should also be a focus of conservation planning in the MSI.

El tipo de hábitat y la permanencia determinan la estructura de la comunidad local de invertebrados acuáticos en las Islas Serranas 

Madrenses. El hábitat acuático de las Islas Serranas Madrenses (ISM) está compuesto de una red de riachuelos intermitentes y perennes,

charcos temporales y represos artificiales de abrevadero, donde todos contribuyen a la biodiversidad regional de especies acuáticas. Aunque las

comunidades acuáticas y las condiciones ambientales prevalecientes en riachuelos y charcas son por lo general muy diferentes, los riachuelos

de las ISM se caracterizan por tener tinajas perennes aisladas la mayor parte del año y son hábitats casi lénticos. En este estudio se compararon

las comunidades acuáticas de coleópteros y hemípteros de tinajas de arroyos con hábitat léntico verdadero (charcos temporales y abrevaderos)

a lo largo de las ISM. Se identificaron 150 especies a lo largo de 38 sitios de estudio. Entre los represos de abrevaderos y las tinajas se presentó

fauna diferente. Las especies exclusivas de las tinajas incluyeron muchas especies longevas y de pobre dispersión, mientras que las especies

exclusivas de losrepresostuvieron en general un periodo de rápido desarrollo y una buena dispersión. En otrasseis corrientes,se compararon las

comunidades de invertebrados acuáticos entre corrientes perennes e intermitentes. Se encontró menos diversidad de invertebrados en corrientes

intermitentes, pero la mayoría de estas especies fueron especialistas y no se encontraron en corrientes perennes. Se sugiere, que en adición a los

arroyos perennes, también se les preste atención a los charcos temporales para la conservación de organismos acuáticos en las ISM.

BONILLAS-MONGE, MARIO ERANDI1 and Carlos Manuel Valdez-Coronel1. 1Departamento de Investigaciones Científicas y Tecnológicas

de la Universidad de Sonora, Luis Donaldo Colosio s/n, entre Sahuaripa y Reforma, Col.Centro, Hermosillo, Sonora, Mexico, C.P. 83000,

marioer.bonillasmo@correoa.uson.mx, jim.jim.furia@hotmail.com.

Mammal richness of San Bernardino Ranch, Agua Prieta, Sonora, Mexico. Historically, economic activities such as grazing and intensive

agriculture have been the support of San Bernardino Ranch, however this has caused environmental damage in the region. The ranch has mayor

ecological value because of its semi desert grasslands, riparian areas and a ciénega, in which restoring efforts take place; we harvest water with

retaining walls. Mammals are important in the ecosystem because they occupy several niches, therefore by knowing the spices present in each

ecosystems, we will also know the status and healthiness of the ecosystem. We used direct and indirect methods; we identified 20 species of

terrestrial mammals. We can compare richness and diversity of the ranch to the San Bernardino National Wildlife Refuge, in the other side of

the border; in which restoration efforts started earlier.
USDA Forest Service Proceedings RMRS-P-67. 2013 517

Abstracts/ Resúmenes

Riqueza de mamíferos del rancho San Bernardino, municipio de Agua Prieta, Sonora, México. Históricamente, en elrancho SanBernardino

se realizaron actividades económicas como son la ganadería y agricultura intensivas, las cuales provocaron el deterioro de los ecosistemas de

la región. El rancho es de gran importancia ecológica debido a que cuenta con varios tipos de hábitats de interés para su conservación como el

pastizal semidesértico, el ribereño y una gran ciénega, la que se está restaurando con la recolección de agua de lluvia por medio de gaviones.

Los mamíferos son importantes en el ecosistema ya que ocupan una gran variedad de nichos, y el conocer las especies actuales que habitan los

diferentes sitios del rancho, nos indicará el balance ecológico del hábitat. Para conocer los mamíferos presentes se usaron métodos de muestreo

directo e indirecto con los cuales se identificaron 20 especies de mamíferos terrestres. Con éstos datos se puede empezar a comparar la riqueza

y diversidad de especies en este rancho y las especies presentes en el San Bernardino National Wildlife Refuge, ubicado exactamente del otro

lado de la frontera, el cual tiene más tiempo con obras de restauración.

BOYLE,BRAD1. 1The University ofArizona, Department ofEcology andEvolutionaryBiology,Tucson,AZ85721, bboyle@email.arizona.edu.

Flora of the Rancho Los Fresnos grasslands: A high-desert crossroads between the Sonoran, Chihuahuan and Great Plains floras. 

Rancho Los Fresnos, a private nature preserve at approximately 1500 m along the U.S. border in northern Sonora, protects over 4,000 hectares

of desert grasslands, oak woodland and riparian vegetation along the headwaters of the San Pedro River. The Los Fresnos flora consists of 245

species in 65 families, with over a third of these species in the three families Poaceae, Asteraceae, and Fabaceae. Approximately 70% of the

species are perennial. Blooming is distributed roughly bimodally between spring and fall, with a slight preponderance of fall-blooming species.

Due the presence of springs, permanent streams, and a large marsh or ciénega, Los Fresnos has a relatively rich riparian flora. Two species,

Lilaeopsis schaffneriana subsp. recurva and Echeandia flavescens, are listed as threatened or endangered in the US. Seven species, among them

Sporobolus giganteus, are apparently new records for the state of Sonora. Multivariate comparison of Los Fresnos with 246 North American

floras showed almost equal floristic affinity with adjacent Sonoran deserts and Chihuahuan desert floras of NM, TX and north-central Mexico,

as well as to short grass prairies of the SW Great Plains and semi-arid sites in central Mexico.

Flora de los pastizales del rancho Los Fresnos: un cruce de floras de desiertos altos entre el Desierto Sonorense, Desierto Chihuahuense 

y las Grandes Llanuras. El rancho Los Fresnos, una reserva natural privada con alrededor de 1500 m de la frontera de México con E.U. en

el norte de Sonora, protege más de 4,000 hectáreas de pastizal del desierto, encinal y vegetación ribereña a lo largo del nacimiento del río San

Pedro. La flora de Los Fresnos consta de 245 especies en 65 familias, con más de la tercera parte de estas especies en tres familias: Poaceae,

Asteraceae y Fabaceae. Casi el 70% de las especies son perennes. En general la floración presenta un patrón bimodal entre primavera y otoño,

con un ligero predominio de las especies con floración de otoño. Debido a la presencia de aguajes, riachuelos permanentes y una gran ciénega,

Los Fresnos cuenta con una flora ribereña relativamente diversa. Dos especies Lilaeopsis schaffneriana subsp. recurva y Echeandia flavescens

están clasificadas como amenazadas o en peligro de extinción en E.U. Siete especies, entre ellas Sporobolus giganteus, parecen ser registros

nuevos para el estado de Sonora. En una comparación multivariada de Los Fresnos con 246 floras deAmérica del Norte se encontró una afinidad

florística casi igual con las floras adyacentes del Desierto Sonorense y el Desierto Chihuahuense en Nuevo México, Texas y la región norte

centro de México, así como las praderas de zacates cortos del suroeste de Las Grandes Llanuras y regiones semiáridas del centro de México.

BRIGGS, M. K.1, Sharon Joy Lite, G. Garfin3, D. Cecil4, and B. Wickel5. 1Chihuahuan Desert Program, World Wildlife Fund, mark.briggs@

wwfus.org; 2Tucson, Arizona, sjoylite@earthlink.net; 3CLIMAS, University of Arizona, gmgarfin@email.arizona.edu; 4National Oceanic Atmospheric Administration, dewayne.cecil@noaa.gov; 5

Conservation Science Program, World Wildlife Fund, bart.wickel@wwfus.org.

Planning river restoration in a changing climate. On December 7-11, 2010, a conference was convened in Tucson, Arizona that brought

together river practitioners, scientists, private citizens, and conservationists from federal and state agencies, academic institutions, and nongovernmental organizations (NGOs) in the southwestern United States (U.S.), northern Mexico, and southeastern Australia to discuss lessons

learned from their river restoration experiences. Key conference topics included the development of viable restoration objectives, planning

and implementing restoration, monitoring results, climate change, environmental flow, native fish conservation, and restoration along transboundary rivers. Results are being incorporated into an applied river restoration guidebook, which summarizes the main steps of developing a

stream restoration project, from planning and implementation to monitoring and evaluation. Some of the main lessons from the conference and

overall strategy of the guidebook will be presented, with emphasis on the guidebook chapter that reviews considerations for developing river

restoration in a changing climate.

Restauración de ríos bajo los efectos del cambio climático. Del 7 al 11 de diciembre del 2010 se llevó a cabo una reunión en Tucson, Arizona, en la cual se congregaron expertos en ríos, científicos, ciudadanos y personal dedicado a la conservación de agencias estatales, federales,

instituciones académicas y organizaciones no gubernamentales (ONG) del sureste de Estados Unidos, norte de México y sureste de Australia,

con la finalidad de compartir experiencias con relación a la restauración de ríos. Los tópicos claves de las conferencias incluyeron desarrollo

de metas de restauración viables, planeación e implementación de la restauración, monitoreo de resultados, cambio climático, flujo ambiental,

conservación de peces nativos y la restauración de ríos transfronterizos. Los resultados se están compilando en una guía de restauración de

ríos, donde se resume como desarrollar proyectos de restauración, desde la planeación y la implementación, hasta el monitoreo y evaluación.

Se presentarán enseñanzas de la reunión y el contenido general del libro, con especial énfasis en el capítulo de restauración de ríos bajo efectos

del cambio climático.
518 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

BUECHER, DEBBIEC. 1 andRonnie Sidner2. 1BuecherBiologicalConsulting, 7050 E. KatchinaCourt,Tucson,AZ 85715, dbuecher@comcast.

net; 2Ecological Consulting, 1671 N. Clifton Street, Tucson, AZ 85745, sidner@email.arizona.edu.

Long distance commute by Lesser Long-nosed Bats (Leptonycteris yerbabuenae) to visit residential hummingbird feeders. Each spring,

thousands of lesser long-nosed bats (Leptonycteris yerbabuenae) migrate from México to northern Sonora and southern Arizona to have their

young and to forage on nectar, pollen and fruit of columnar cacti. Once the pups are volant, the population begins to disperse across the grasslands of southeastern Arizona, foraging on flowering agaves (Agave palmeri) and day-roosting in Madrean Sky Islands during their seasonal

migration back to México. Although the grasslands surrounding Arivaca, Arizona have no naturally occurring fields of agaves, nectar bats are

documented visiting area hummingbird feeders to obtain sugar water. To better understand foraging patterns by these bats in an area with no

natural food plants, we radio-tracked 28 lesser long-nosed bats in 2010 - 2011. We captured bats at area hummingbird feeders and tracked them

to a day-roost in the Santa Rita Mountains, approximately 25 miles away. We discovered that bats nightly performed a long-distance commute

from the Santa Rita Mountains to Arivaca, bypassing available hummingbird feeders closer to their day-roost. It may be that the large colony

size in the day-roost necessitates long-distance dispersal as a mechanism to reduce competition for limited food resources.

Desplazamientos largos de murciélagos nectarívoros (Leptonycteris yerbabuenae) para visitar bebederos residenciales de chuparrosas. 

Cada primavera miles de murciélagos nectarívoros (Leptonycteris yerbabuenae) migran desde México hasta el norte de Sonora y el sur de

Arizona para parir y alimentarse de néctar, polen y frutos de cactus columnares. Una vez que las crías pueden volar, la población empieza a

dispersarse en los pastizales del sureste de Arizona, alimentándose de las flores de maguey (Agave palmeri), encontrando refugio diurno en

las Islas Serranas Madrenses durante su migración anual de regreso a México. Los pastizales que rodean Arivaca, Arizona no cuentan con

poblaciones de magueyes, aún así los murciélagos nectarívoros se han documentado visitando los bebederos de chuparrosas para tomar el agua

azucarada. Con el fin de tener un mejor conocimiento de los patrones de alimentación de estos murciélagos en un área sin plantas alimenticias

nativas, de 2010 a 2011, usamos la técnica de rastreo con radio telemetría en 28 murciélagos nectarívoros. Capturamos los murciélagos en los

bebederos de chuparrosas y los rastreamos a sus refugios diurnos en la Sierra Santa Rita, ca. 40 km de distancia. Descubrimos que cada noche

los murciélagos se desplazaron una larga distancia desde la Sierra Santa Rita a Arivaca, eludiendo los bebederos más cercanos a sus refugios

diurnos. Una explicación podría ser que las colonias grandes de los refugios diurnos necesitan una dispersión larga como mecanismo para

reducir la competencia por recursos alimenticios limitados.

CASTELLANOS-VILLEGAS,ALEJANDROE.1,LuisC.Bravo-Peña2,HernánCelaya-Michel1,ArturoIbarra-Acosta1, andJoséR.Romo-León3. 1

DICTUS, Universidad de Sonora, Hermosillo, Sonora, 83000; 2UMC, Universidad Autónoma de Cd. Juárez, Cd. Cuauhtémoc, Chihuahua; 3

SNRE, University of Arizona, Tucson, AZ 85721.

Biodiversity effects on ecosystem function due to land use: The case of buffelgrass savannas in the Sky Islands seas in the Central Region 

of Sonora. Buffelgrass savannas have been an important landscape in cattle grazing ranches in Sonora over the past 50 years or so. Changes

in land use results in biodiversity changes that may result in ecosystem functional changes, however these are less documented. Although fire

cycle driven processes have been proposed for Buffelgrass savannas, this is not generally the case, and other processes seem to be driving ecosystem function. Several years of studying above- and below ground processes allow us to propose how water and nutrient dynamics change

in established buffelgrass savannas, as well as how biodiversity changes affect functional processes in arid and semiarid ecosystems. Water

and nutrient biogeochemical cycles changed in buffelgrass dominated savannas with regard to those in natural ecosystems, following effects of

land use changes in soil biological and physical changes. Our findings may be extrapolated to what could be expected to happen to other highly

invaded areas were buffelgrass is becoming a dominant exotic species.

Cambios en el uso del suelo afectan la biodiversidad y la función del ecosistema: El caso de pastizales de zacate buffel en las Islas 

Serranas del centro de Sonora. Los pastizales de buffel han sido comunes en los ranchos ganaderos de Sonora en los últimos 50 años. Los

cambios del uso del suelo producen cambios en la biodiversidad y pueden resultar en cambios funcionales del ecosistema, los cuales han sido

poco documentados.Aunque los procesos causados por incendios han sido propuestos para pastizales de buffel, esto no es generalmente el caso,

y otros procesos parecen afectar la función del ecosistema. Varios años de estudio de los procesos sobre y debajo del suelo, nos permitieron

registrar como la dinámica del agua y los nutrientes cambian en praderas de buffel y como los cambios en la biodiversidad afectan los procesos

funcionales de ecosistemas áridos y semiáridos. Debido al efecto del cambio del uso del suelo en los procesos físicos y biológicos, los ciclos

biogeoquímicos de agua y nutrientes cambian en pastizales dominadas por buffel comparados con ecosistemas naturales. Nuestros resultados

pueden extrapolarse a otras áreas altamente invadidas donde el buffel es la especie exótica dominante.

CHEN, HSIANG LING and John L. Koprowski. Wildlife Conservation and Management, School of Natural Resources and the Environment,

The University of Arizona, Biological Sciences East, Tucson, AZ 85721, cherlene@email.arizona.edu

Effects of roads on wildlife: A review of impacts, mitigation and implications within the Sky Islands. Roads are conspicuous and pervasive

features of landscapes and represent one of the most significant anthropogenic impacts on natural areas and wildlife. The goal of this review

is to summarize recent studies on road impacts on wildlife and identify gaps in our understanding of the topic in the Madrean Archipelago.

The sky islands region is defined by natural levels of fragmentation due to geography; however, human population growth and transportation

needs threaten to exacerbate levels of isolation in the region. We review road ecology related peer-reviewed publications and governmental

research reports and synthesize geography, biodiversity, and studies’ focus. Study areas are concentrated along highways and state routes in

Arizona. A taxonomic bias toward large mammals is evident. Despite a prevalence of studies on wildlife road crossing, most research focuses
USDA Forest Service Proceedings RMRS-P-67. 2013 519

Abstracts/ Resúmenes

on distribution and movements; whereas population impacts are rarely described. Our review will help to guide the future direction of research

and management in the Madrean Archipelago to minimize road impacts.

Efecto de carreteras en fauna silvestre: análisis de impactos, mitigación y consecuencias en las Islas Serranas. Las carreteras, una característica visible y generalizada del paisaje, representan uno de los impactos antropogénicos más significativos de las áreas naturales y fauna

silvestre. El objetivo de este análisis es condensar los estudios recientes sobre impactos de carreteras en fauna silvestre e identificar lagunas

de información sobre el Archipiélago Madrense. Las islas serranas se definen naturalmente por diferentes niveles de fragmentación debido a

la geografía de la región, sin embargo el crecimiento de la población humana y por ende la necesidad de trasportación amenaza con aumentar

los niveles de aislamiento de la región. El análisis incluye publicaciones científicas y reportes gubernamentales sobre ecología de carreteras, y

se sintetiza la geografía, biodiversidad y estudios relevantes. Los sitios de estudio se distribuyen a lo largo de autopistas federales y carreteras

estatales enArizona. Elsesgo taxonómico hacia mamíferos mayores es evidente.Apesar de la frecuencia de estudios de fauna silvestre cruzando

carreteras, la mayoría se enfoca en distribución y movimiento; y por lo general se ignora el impacto sobre la población. Este análisis servirá

para guiar la investigación y manejo futuros para minimizar impactos de carreteras en el Archipiélago Madrense.

CHOATE, BRITTANY1, Joanna B. Nadeau2, Candice Rupprecht3, Aaron M. Lien4, and Sharon B. Megdal5. 1Water Resources Research Center,

The University of Arizona, Tucson, AZ 85721, bchoate@email.arizona.edu; 2Water Resources Research Center, The University of Arizona,

Tucson, AZ 85721, jnadeau@cals.arizona.edu; 3Water Resources Research Center, The University of Arizona, Tucson, AZ 85721, candicer@

cals.arizona.edu; 4Water Resources Research Center,The University ofArizona,Tucson,AZ 85721, amlien@cals.arizona.edu; 5Water Resources

Research Center, The University of Arizona, Tucson, AZ 85721, smegdal@cals.arizona.edu.

Conserve to Enhance: An innovative mechanism for environmental benefits. Arizona’s riparian ecosystems have been susceptible to degradation because state water laws do not consider environmental water needs. This lack of legal authority has led to surface water diversion and

groundwater consumption with little or no consideration of impacts on water dependent ecosystems (Megdal et al. 2011). To help address this

challenge, The University of Arizona Water Resources Research Center (WRRC) has developed and is implementing the innovative Conserve

to Enhance (C2E) mechanism to bring the environment to the table as a paying water customer. Conserve to Enhance addresses the perception

that municipal water conservation does not directly benefit the environment (Schwarz and Megdal 2008). C2E offers water customersthe option

to donate money saved through voluntary water conservation actions to projects that support environmental water needs. The Sonoran Institute,

Tucson Water, Watershed Management Group, and WRRC have partnered to implement C2E in Tucson since January 2011. This poster details

outcomes from the first year of the Tucson pilot, including the program development process, the amount of funds raised, and the quantity of

water saved. Lessons learned during program development in Tucson will help inform implementation of C2E in other communities.

Conservar para Mejorar: un mecanismo innovador con beneficios ambientales. Los ecosistemasribereños deArizona han sido susceptibles

a la degradación debido a las leyes estatales que no toman en cuenta las necesidades de agua del medio ambiente. Esta falta de autoridad legal

ha dado lugar a la desviación del agua en la superficie y el consumo de aguas subterráneas con poca o ninguna consideración a los impactos

en los ecosistemas que dependen del agua (Megdal et al. 2011). Para enfrentar este desafío, la Universidad de Arizona Centro de Investigación

sobre los recursos del agua “Water Resources Research Center” (WRRC) ha desarrollado y está implementando el mecanismo innovador de

Conservar para Mejorar “Conserve to Enhance” (C2E) para poner al medio ambiente en la mesa de discusiones como un cliente más que paga

el agua. Conservar para Mejorar aborda la percepción de que la conservación del agua municipal no beneficia directamente al medio ambiente

(Schwarz y Megdal 2008).C2E ofrece a los consumidoresla opción de donar el dinero ahorrado en la conservación voluntaria de agua a proyectos

que apoyan las necesidades de agua del medio ambiente. El Instituto Sonorense, Organismo de Agua de Tucson, Grupo de Manejo de Cuencas

Hidrográficas y WRRC se unieron para poner en práctica C2E en Tucson desde enero de 2011. Este cartel detalla los resultados del primer año

de este proyecto piloto, el proceso de desarrollo del programa, la cantidad de fondos recaudados, y la cantidad de agua ahorrada. Las lecciones

aprendidas durante el desarrollo del programa en Tucson servirán para aplicar el C2E en otras comunidades.

CLARK, JESS1, Carolyn Napper2, Kevin Cooper3, and Marc Stamer4. 1U.S. Forest Service, Geospatial Management Office, Remote Sensing

Applications Center, Salt Lake City, UT, 84119; 2U.S. Forest Service, Mt. Shasta McCloud Management Unit, Mt. Shasta CA, 96067; 3U.S.

Forest Service, Santa Lucia Ranger District, Santa Maria, CA, 93454; 4U.S. Forest Service, Mountaintop Ranger District, Fawnskin, CA, 92333.

Utilization of Satellite Imagery to Evaluate and Predict Out-Year Post-Fire Watershed Response and Potential Application in the Southwest. Burned Area Emergency Response (BAER) teams, along with other agencies that have jurisdictional responsibility (Natural Resource

Conservation Service, state, counties, cities, etc.), prescribe and implement treatments based on expected post-fire response and associated

threats to life, property, and resources. Vegetative cover is often used to evaluate watershed recovery and the start of a return to pre-fire response

behavior. The objective of this project is to use remote sensing technologies to assess the rate of natural vegetative regeneration post-fire and

evaluate the level of risk for continued increased post-fire watershed response. If remote sensing indices are effective in determining the rate of

natural vegetative recovery in burned areas, then we can determine how peak flows, debris flows, and sedimentation and erosion factors have

changed, and this information will enable us to determine the level of risk for an area? This discussion summarizes steps taken, initial findings

from the Horseshoe 2, Monument, and other fires, and identifies future needs to implement a decision support tool with national application that

will give land managers and emergency response personnel an indication of the level of risk relative to pre-fire or immediate post-fire levels.
520 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

El uso de imagen satelital para evaluar y predecir la recuperación posterior a incendios en las cuencas y su aplicación potencial en el 

Suroeste Norteamericano. Las brigadas de BAER (Respuesta de Emergencia para Áreas Incendiadas, por sus siglas en inglés), junto con otras

agencias que tienen responsabilidad jurídica (Servicio de Conservación de Recursos Naturales, estado, municipios, ciudades, etc.), prescriben

e implementan tratamientos basados en respuestas esperadas posteriores al incendio y amenazas a la vida, propiedad y recursos. La cobertura

vegetal se emplea para evaluar la recuperación de la cuenca y el regreso al estado previo al incendio. El objetivo de este proyecto es emplear

tecnologías como sensores remotos para analizar el tiempo de regeneración de cobertura vegetal y evaluar el nivel de riesgo de recuperación

de la cuenca, después del incendio. Sí los índices desarrollados a partir de sensores remotos son efectivos en la determinación de recuperación

de cobertura vegetal en zonas quemadas, entonces se pudiera determinar la corriente máxima, flujo de detritos, sedimentación, erosión, y usar

esta información para determinar niveles de riesgo. Este trabajo analiza resultados preliminares de los incendios “Horseshoe 2”, “Monument”,

y otros, e identifica necesidades futuras para implementar una estrategia de decisión, con aplicaciones a nivel nacional que permita a los encargados de manejo de recursos y brigadistas identificar el nivel de riesgo previo y posterior al incendio.

COGAN, ROGER C. National Audubon Society, Appleton-Whittell Research Ranch, Elgin, AZ.

Rattlesnake Wintering Sites at the Appleton-Whittell Research Ranch. The Research Ranch is an 8000-acre sanctuary and research facility in the semi-arid grasslands of southeastern Arizona managed by the National Audubon Society. At the Research Ranch we work to protect

native species and the natural systems that occur both on the preserve and the surrounding area. Snakes are an integral element within all of

the habitats where they are found. Snakes not only help maintain the fitness of their prey but they themselves are food for a wide variety of

predators. They are a vital part of the landscape at all times of the year, yet little is know about the habitat requirements necessary for successful overwintering. Four rattlesnake species are known to occur at the Research Ranch: Western Diamondback (Crotalus atrox), Black-tailed

(Crotalus molossus), Rock (Crotalus lepidus) and Mohave (Crotalus scutulatus) rattlesnakes. During the months of February, March and April

2011, a survey for potential rattlesnake wintering sites was conducted. This search was conducted whenever time permitted or when weather

conditions seemed most likely to expose snakes at the surface. Surveys during February, March and April are conducted when climate conditions are optimal for snakes to exit their den sites for brief periods. Confirmed den sites are photographed and georeferenced and, if possible,

the snakes are photographed for vouchering.

Guaridas de invierno de víboras de cascabel en la estación biológica Appleton-Whittell Research Ranch. El centro de investigación es un

santuario de 3,237 has en los pastizales semiáridos del sureste de Arizona a cargo de la Asociación Nacional Audubon. En la estación se trabaja

para proteger las especies nativas y los sistemas naturales que ocurren tanto en la reserva como en el área adyacente. Las serpientes son un

elemento esencial de todos los hábitats donde se encuentran y no sólo mantienen el balance de sus presas sino que también son el alimento de

una gran variedad de depredadores. Aunque son parte vital del ecosistema todo el año, se conoce muy poco sobre los requisitos del hábitat para

que sobrevivan el invierno. En esta reserva ocurren cuatro especies de víboras de cascabel: Crotalus atrox (cascabel de diamantes), Crotalus 

molossus (cascabel de cola negra), Crotalus lepidus (chichámora) y Crotalus scutulatus (cascabel mohavense). En febrero, marzo y abril de

2011 se realizó un censo de los sitios potenciales de invierno para víboras de cascabel Las condiciones climáticas de esos tres meses son óptimas

para que las serpientes salgan de sus guaridas por tiempos cortos, lo que se aprovecha para censar. Las guaridas donde se confirma su presencia

se fotografían, se ubican con coordenadas y cuando es posible también se toman fotos de las serpientes para documentarse en una colección.

COGAN, ROGER C. National Audubon Society, Appleton-Whittell Research Ranch, Elgin, AZ.

Herpetofauna at the Appleton-Whittell Research Ranch. A rich diversity of amphibian and reptile species occurs at the Appleton-Whittell

Research Ranch. The Research Ranch is an 8000-acre sanctuary for native biota and research facility in the semi-arid grasslands of southeastern

Arizona, managed by the National Audubon Society. Since cattle were removed from the sanctuary in the late 1960s, nine species of amphibians and forty-two species of reptiles have been identified by staff and researchers within the preserve. Several species have not been seen in

the recent years and may no longer occur on the Research Ranch. Ongoing efforts are underway to locate and document the presence of the

herpetofauna. Our challenge into the future is to safeguard the continued existence of those that are still found and return native species that

may have been extirpated.

Herpetofauna de la estación biológica Appleton-Whittell Research Ranch. La estación biológica Appleton-Whittell Research Ranch cuenta

con una gran diversidad de especies de anfibios y reptiles. El centro de investigación es un santuario de 3,237 has de biota nativa en los pastizales

semiáridos del sureste de Arizona a cargo de la Asociación Nacional Audubon. El personal y los investigadores visitantes han identificado 9

especies de anfibios y 42 especies de reptiles en la reserva desde que se desalojó el ganado a finales de la década de 1960. Varias especies no se

han visto los últimos años y quizás ya no ocurren en la estación, por lo que se están llevando a cabo investigaciones para localizar y documentar

la herpetofauna. El reto es proteger la existencia continua de las especies presentes y reintroducir especies nativas extirpadas.

COLE, JULIA. Geosciences and Atmospheric Sciences, University of Arizona, 1040 E 4th St., Tucson, AZ 85721, jecole@email.arizona.edu.

Climate change, megadroughts, and the North American monsoon. Climate change in theSouthwesttranslatesto changesinwater availability,

and records of past hydroclimate help us delineate the natural range of those changes. Past droughts are apparent in tree-ring based records and

emerge from longer paleoclimate records as well. But do these droughts reflect changes in winter or summer moisture? This question is more

than academic: as we move into a warmer world, we can expect generally drier winters and springs. But projections of future climate disagree
USDA Forest Service Proceedings RMRS-P-67. 2013 521

Abstracts/ Resúmenes

with respect to changes in the North American monsoon, and global models do not yet simulate this system well. Here we use paleoclimate

records to explore how summer rainfall has responded during past climate regimes. In this talk I will describe new results from cave-based

climate records that demonstrate how, over the past 7000 years, the monsoon has clearly weakened. This result is consistent with other monsoon

systems globally and with climate model simulations. I will explore implications for future hydroclimate regimes in our Southwest region.

El cambio climático, megasequías y el monzón norteamericano. El cambio climático en el Suroeste Norteamericano induce cambios en la

disponibilidad de agua, y los registros ambientales hidrológicos nos ayudan a delinear el rango natural de esos cambios. Las sequías anteriores

son evidentes en los registros de anillos de árboles (dendroclimatología) y también emergen de registros de paleo-climas más largos. Pero estas

sequías, ¿reflejan cambios en la humedad de invierno o verano? Esta pregunta es más que académica: conforme el mundo se vuelve más cálido podemos esperar inviernos y primaveras más secas. Pero las proyecciones del clima del futuro no coinciden en los cambios en el monzón

norteamericano y los modelos mundiales todavía no simulan bien este sistema. Aquí usamos registros paleo climáticos para explorar como

las lluvias de verano han respondido durante regímenes climáticos anteriores. En esta charla describiré nuevos resultados basados en registros

climáticos de cuevas para mostrar como los últimos 7000 años, el monzón claramente se ha debilitado. Este resultado es consistente con otros

sistemas monzónicos mundiales y con modelos climáticos. Exploraré implicaciones para regímenes hidroclimáticos futuros en nuestra región

del Suroeste Norteamericano..

CORONEL-ARELLANO, HELÍ2, Nalleli E. Lara-Díaz1, Alejandro González-Bernal2, Gerardo Carreón-Arroyo2, Mario Cirett-Galán y Carlos

A. López-González1. 1Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Av. de las Ciencias S/N, Col. Juriquilla, Santiago

de Querétaro, Querétaro, C.P. 76230; 2Naturalia A. C., Av. 32 entre Calle 3 y 4 Villa Satélite 3250, Agua Prieta, Sonora, C.P. 84200; 3CONANP.

Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Calle 2a Este final y Sinaloa, Col. Centro, Cananea, Sonora. C.P. 84620.

Abundance and biomass of potential prey species for Mexican wolves (Canis lupus baileyi) in Sonora, México. Quantification of prey

availability is a necessary step toward the successful implementation of a carnivore reintroduction program, under the assumption of a higher

probability of success when a community is more diverse reflected on a lower biomass turnover, consequently creating a buffer for the newly

introduced predator. The objective of this study was to assess the diversity, abundance and prey biomass potentially available to reintroduce

Mexican wolves. We used camera traps between 2009 to 2011 in a 29,000 sq. km area, used a site-specific protocol to assess minimum number

of prey and this was transformed to biomass by multiplying the weight times the density of a given species. We documented a total of 11 species, where white-tailed deer, Gould ́s Turkey and rock squirrels each contribute 50 kg/km2

. The remaining species although can be considered

occasional or rare, contribute about 11.22 kg/km2. Our results show that the area can support 25 to 30 Mexican wolves, with no negative effects

toward prey species or livestock losses.

Diagnóstico de la biomasa disponible de presas útiles para el lobo mexicano (Canis lupus baileyi) en Sonora, México. Es necesario conocer la cantidad de presas disponibles para el éxito de un programa de reintroducción de carnívoros. Se asume que la probabilidad de éxito

es más alta en una comunidad con mayor diversidad, dándole cierta protección al depredador recién introducido. El objetivo de este estudio

fue evaluar la diversidad, abundancia y biomasa potencial de presas disponibles para la reintroducción del lobo mexicano. Se usaron cámaras

trampa de 2009 a 2011 en un área de 29,000 km2 y una metodología adaptada al sitio para evaluar la cantidad mínima de presas. La biomasa

de cada especie se obtuvo al multiplicar el peso por la densidad de la especie. Se documentaron un total de 11 especies, donde el venado cola

blanca, el guajolote silvestre y el ardillón contribuyen 50 kg/km2 cada uno. Aunque el resto de las especies se consideran ocasionales cada una

contribuye cerca de 11.22 kg/km2. Estos resultados muestran que el área puede mantener de 25 a 30 lobos mexicanos sin afectar negativamente

a las presas o al ganado.

CORTÉS-MONTAÑO, CITLALI1, P.Z. Fulé1, T.J. Ayers2. 1School of Forestry, Northern Arizona University, P.O. Box 15018, Flagstaff, AZ,

86011, USA; 2Department of Biology and Deaver Herbarium, Northern Arizona University, P.O. Box 5640, Flagstaff, AZ, 86011.

Preliminary standard for old-growth forests in the Sierra Madre Occidental. The Sierra Madre Occidental of México has temperate oldgrowth forests that have been seldom studied. They have been in decline since the advent of industrial logging, and currently lack protection

unlessincluded in protected areas. We collected structure and composition data in two temperate forestsin Chihuahua, and based on this propose

a regional preliminary old-growth standard. Our results showed that the basal area of old-growth forests was concentrated in diameter classes

>40 cm, and that trees in diameter classes >60 cm were present in all the sampled sites. The minimum basal area was 25 m2, dominated by

species in the Pinaceae (Pinus and Pseudotsuga), but Quercus and broadleaved species contributed to overstory diversity. Snags and large logs

were present. The understory was dominated by native species in the Asteraceae, Fabaceae, Poaceae and Lamiaceae. We found that indicator

species that favor old-growth, such as the Thick-billed Parrot, can be useful in identifying sites and to promote their conservation. Since the

information presented here was based on a dataset generated in two locations, it needs to be updated as more sites are explored, but meanwhile

it can be used to negotiate protection of old-growth sites of northern México.

Estándar preliminar de bosques antiguos en la Sierra Madre Occidental. Los bosques antiguos de la Sierra Madre Occidental de México

han sido poco estudiados. Su cobertura ha declinado desde el inicio de la tala industrializada, y solo se encuentran protegidoslegalmente si están

incluidos en áreas protegidas. Proponemos un estándar regional preliminar de bosques antiguos con base en datos de estructura y composición

colectados en dos bosquestemplados de Chihuahua. Nuestrosresultadosindican que el área basal de los bosques antiguosse concentró en clases

de diámetro >40 cm, y en todos los sitios de muestreo registramos árboles con diámetros >60 cm. El área basal mínima fue 25 m2, dominada

por especies de pinacéas (Pinus y Pseudotsuga), mientras que Quercus y otras especies de latifoliadas contribuyeron a la diversidad del dosel.
522 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

En todos los sitios registramos la presencia de arbolado muerto en pie y troncos sobre el piso forestal. El estrato herbáceo y arbustivo estuvo

dominado por especies nativas de Asteraceae, Fabaceae, Poaceae y Lamiaceae. El uso de especies indicadoras que favorecen las condiciones

de hábitat de los bosques antiguos, como las guacamayas o cotorras serranas, es útil para la identificación de sitios y para promover su conservación. La información presentada se basó en datos generados en dos localidades, por lo que es importante actualizar este estándar a medida

que se exploran otros sitios. Mientras tanto, se puede utilizar como medida temporal para proteger los bosques antiguos de las montañas del

noroeste de México.

CORTÉS-MONTAÑO, CITLALI1, P.Z. Fulé1, D.A. Falk2, T.J. Ayers3. 1School of Forestry, Northern Arizona University, P.O. Box 15018,

Flagstaff, AZ, 86011, USA, 2School of Natural Resources and the Environment and Laboratory of Tree-Ring Research, University of Arizona,

Tucson, AZ, 85721 USA; 3Department of Biology and Deaver Herbarium, Northern Arizona University, P.O. Box 5640, Flagstaff, AZ, 86011.

Two old-growth forests from Chihuahua, México: composition, structure, age, and fire disturbance. We studied two high elevation

(>2000 meters) old-growth forests in the Sierra Madre Occidental in Chihuahua, México: Mesa de las Guacamayas (MDG), located ~100 km

from the U.S.-México border, and Parque Nacional Cascada de Basaseachi (PNCB), located in central Chihuahua. We measured the structure

and composition of their overstory and understory, and collected increment cores and fire scars to create age reconstructions and fire histories.

Both sites had high understory ß-diversity, while overstory ß-diversity was higher at PNCB. Understory species were all native at MDG, and

we only collected one exotic at PNCB. Forests in the two sites had snags >40 cm and logs on the forest floor, as well as trees with diameters

>60 cm. The oldest trees at the two sites established in the mid 1700s, and their fire histories indicate that frequent fires that burn at least once

every decade are part of their disturbance regimes. Both sites have high conservation value and are included in protected areas, while MDG is

the northernmost known nesting site for Thick-billed Parrots. Conservation of these forests should incorporate integrated fire management, and

structural and compositional complexity should be maintained in order to ensure habitat quality.

Dos bosques antiguos de Chihuahua, México: composición, estructura, edad e historia de incendios. Estudiamos dos bosques antiguos

en elevaciones >2000 metros en la Sierra Madre Occidental de Chihuahua, México: Mesa de las Guacamayas (MDG), ubicada a ~100 km de

la frontera México-E.U., y el Parque Nacional Cascada de Basaseachi (PNCB), ubicado en el centro del estado. Medimos la estructura y composición del dosel y el estrato herbáceo y arbustivo, y colectamos cilindros de incremento y cicatrices de incendios para crear reconstrucciones

de edad e historias de incendios. La diversidad ß del estrato herbáceo y arbustivo fue alta en los dos sitios, mientras que la diversidad ß del dosel

fue más alta en PNCB. Todas las especies del estrato herbáceo y arbustivo colectadas en MDG fueron identificadas como nativas, y solo una de

nuestras colectas en PNCB fue identificada como exótica. Los bosques en los dos sitios presentaron árboles muertos en pie con diámetros >40

cm y troncos en el piso forestal, así como árboles con diámetros >60 cm. Los árboles más antiguos en los dos sitios se establecieron a mediados del siglo XVIII, y sus historias de incendios indican que los regímenes de perturbación de ambos sitios presentan incendios frecuentes, al

menos una vez cada década. Ambos sitios son de alto valor para la conservación y están incluidos en áreas protegidas, y MDG es el sitio más

norteño de anidación de la guacamaya o cotorra serrana. La conservación de estos bosques debe de incorporar manejo integrado del fuego, y

la complejidad estructural y de la composición debe de mantenerse con el fin de asegurar la calidad del hábitat.

CRIMMINS, THERESA M.1,2, Michael A. Crimmins3, and C. David Bertelsen4. 1Office of Arid Lands Studies, University of Arizona, 1955 E.

Sixth St., Tucson, AZ 85721, theresam@u.arizona.edu; 2National Coordinating Office, USA National Phenology Network, Tucson, AZ, 3Department of Soil, Water and Environmental Science, University of Arizona, Tucson, AZ, crimmins@u.arizona.edu; 4

Herbarium, University of

Arizona, Tucson, AZ, david.bertelsen8@gmail.com.

Temporal patterns in species flowering can inform long-term plant monitoring. The flowering patterns of plants of the Finger Rock trail,

located in the Santa Catalina Mountains of southern Arizona, have been very carefully documented on an approximately weekly basis for

nearly three decades. These observations, made along a gradient encompassing more than 4,000 ft in elevation, have yielded key insights into

climatic flowering cues as well as range shifts among particular species. These long-term observations of flowering plant phenology have also

revealed that the regularity with which plant species flower from year to year is highly variable. We implemented a clustering algorithm to

elucidate recurrent temporal flowering patterns in species. Several patterns emerged, and these patterns are consistent with plant life forms and

the water use strategies they employ as well as native/non-native status. The findings of this analysis are valuable for guiding species selection

for long-term monitoring efforts.

Uso de los patrones temporales de floración en el monitoreo a largo plazo de plantas. La floración de plantas en la vereda Finger Rock, de

la Sierra Santa Catalina del sur de Arizona, se documentó en detalle casi semanalmente durante tres décadas. Las observaciones en un gradiente

altitudinal de más de 1220 m, arrojaron información clave sobre estímulos climáticos de floración y cambios de distribución de ciertas especies.

Estas observaciones de la fenología de floración, hechas por muchos años, también mostraron que la regularidad con que las especies florecen

año tras año es muy variable. Se implementó un algoritmo de ordenamiento para esclarecer los patrones de floración temporales recurrentes. Se

obtuvieron varios patrones consistentes con las formas de vida de las plantas y sus estrategias de uso de agua, y también su estatus de nativa o

introducida. Los resultados de estos análisis son muy útiles en la selección de especies para el monitoreo a largo plazo.
USDA Forest Service Proceedings RMRS-P-67. 2013 523

Abstracts/ Resúmenes

CURÉ,JENNIE.Arizona State University School ofApplied Math and Science,International EnvironmentalTechnology Management Graduate

Program, PO Box 242,Cave Creek, AZ 85327, JCure@asu.edu.

Agricultural field restoration utilizing native grasses; Lower San Pedro River watershed. Agricultural Field Restoration Utilizing Native

Grasses; Lower San Pedro River Watershed is a case study documenting results from three representative projects. The projects are partnerships

between The Nature Conservancy and individual property owners. The study documents the challenges with the process of agricultural field

restoration, the methods utilized for change over to native grasses at the specific sites and the results. The suite of potential benefits of grass

cropping projects is numerous and reviewed within the study analysis. Potential benefits include; reduction in water and power usage, allow

businesses to be more sustainable and resilient to drought in the future, restore the integrity of the riparian habitat and address the affects of

climate change. The longer-term outcomes we hope to see from sharing this research at the conference are; an interest in change over to native

grasses, an increased market in native grass hay and the development of seed banking for rehabilitation projects. One of the goals of the research

is to inspire others to try to answer some of the questions within the framework of the process of the restoration projects. The research provides

documentation to support further study and has utility for different audiences.

Restauración de campos agrícolas mediante el uso de zacates nativos en la cuenca baja del río San Pedro. La restauración de campos

agrícolas utilizando zacates nativos en la cuenca baja del río San Pedro es un estudio de caso que documenta resultados de tres proyectos representativos. El proyecto es una colaboración entre la organización The Nature Conservancy y propietarios. El estudio documenta los retos en

el proceso de restauración de campos agrícolas, los métodos usados para el cambio a zacates nativos y los resultados. La variedad de beneficios

potenciales del cultivo de zacates nativos son numerosos y se examinan en el análisis del estudio. Los beneficios incluyen la disminución en el

uso de agua y electricidad, durabilidad y resistencia a futuras sequías, restauración de los hábitats ribereños y adaptación al cambio climático.

Al presentar este trabajo se espera: un interés en el cultivo de zacates nativos, aumento del mercado de forraje de zacates nativos y la creación

de un banco de semillas para proyectos de restauración. Una de las metas de este proyecto es inspirar para buscar respuestas a los problemas

que se presentan en los proyectos de restauración. Esta investigación tiene la documentación necesaria para apoyar estudios adicionales útiles

a diferentes audiencias.

CZAPLEWSKI, N.J.1 and SteveWillsey2. 1Oklahoma Museum of Natural History, 2401 ChautauquaAvenue, University of Oklahoma, Norman,

OK 73072, nczaplewski@ou.edu; 2P.O. Box 963, Hereford, AZ 85615, stee25bee@msn.com.

A Late Quaternary Brown Bear (Ursinae: Ursus cf. arctos) from a cave in the Huachuca Mountains, Arizona. In 2008 one of the authors

(SW) discovered the fragmentary cranium of a bear, loose on the floor of a cave at about 2270 m elevation near the crest of the Huachuca Mountains. In 2009 we revisited the cave to examine the specimen in order to identify the species. We photographed and measured the main pieces

and left them in the cave. The skull is from an adult, possibly male with prominent sagittal crest. Bears are highly variable morphologically

and their remains are difficult to identify. Many North American black bears were larger in the late Pleistocene and underwent a size reduction

in the Holocene. The morphological features and measurements of the present cranium are somewhat equivocal, but most available features

indicate a brown bear, Ursus cf. arctos. Some parts are encrusted with carbonate and could be better examined after collection and preparation

as well as comparison with late Pleistocene brown and black bears. Based on its state of preservation, the cranium possibly represents a late

Pleistocene occurrence, which could be determined by radiometric dating. There is no previous fossil record of U. arctos in the Sky Islands,

nor in the rest of Arizona, New Mexico, Chihuahua, or Sonora.

 Registro de oso pardo (Ursinae: Ursus cf. arctos) del Cuaternario tardío de una cueva en la Sierra Huachuca, Arizona. En 2008 uno de

los autores (SW) encontró el fragmento de un cráneo de oso en el suelo de una cueva a 2270 m de elevación, cerca de la cumbre de la Sierra

Huachuca. En 2009 se visitó de nuevo la cueva para examinar el ejemplar e identificar la especie. Se tomaron fotografías y medidas de las partes principales sin removerse de la cueva. El cráneo es de un adulto, posiblemente macho con una marcada cresta sagital. Los osos tienen una

morfología muy variable y es muy difícil identificar sus restos óseos. Muchos osos negros de América del Norte fueron más grandes a finales

del Pleistoceno pero su tamaño se redujo en el Holoceno. Las medidas y características morfológicas de este cráneo son un poco ambiguas, sin

embargo indican que es un oso pardo Ursus cf. arctos. Algunas partes están incrustadas con carbonato y podrían examinarse con más detalle

después de recolectarse, prepararse y compararse con osos pardos y negros de finales del Pleistoceno. Según su estado de preservación, es

posible que el cráneo represente un registro del Pleistoceno lo que podría determinarse por el método del carbono 14. No hay registros previos

de fósiles de U. arctos de las Islas Serranas de Arizona, Nuevo México, Chihuahua o Sonora.

DEAN, VIRGINIA. Appleton-Whittell Research Ranch of the National Audubon Society, Elgin, AZ.

Biodiversity database at the Appleton-Whittell Research Ranch. The Appleton-Whittell Research Ranch provides the location, resources,

and opportunity for natural-history research projects in ungrazed, unimproved grasslands of southeastern Arizona, USA. At ranch headquarters,

research since 1968 is archived as hard-copy pages of published and unpublished documents, conference posters, electronic files of data and

photographs, specimen collections, and species checklists for visitors. A recent addition to the archiving methods is an Excelâ database of all

plants or animals that have been documented as occurring on the Research Ranch. Each record includes several fields that are filled in whenever

the information is available. These include order, family, genus, and species names; date of the sighting; location by GPS, landmark, or local

place-name; type of habitat; life stage of the specimen; names and titles of the observer and species identifier; observed associated species

and relationships; and photographs. A comment column or additional separate columns contain other interesting observed data, e.g., weather
524 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

conditions, time of day, activity, health, size, color, unique attributes. The database consolidates local historical data that can be updated and

used by future researchers in their reports.

Base de datos de la Estación Biológica Appleton-Whittell. La Estación Biológica Appleton-Whittell proporciona la ubicación, recursos y

oportunidades para proyectos de investigación en historia natural en las praderas sin pastoreo, ni mejoradas del sureste de Arizona, E.U..En sus

instalaciones, desde 1968 se archivan copias de los documentos publicados y sin publicar, carteles en congresos, archivos electrónicos de datos

y fotografías, colecciones de especimenes y listados de especies, disponibles para visitantes. Recientemente se creo una base de datos de Excelâ 

de todos los animales y plantas registrados en la Estación Biológica. Cada registro cuenta con varios campos que se capturan con la información

disponible. Éstos incluyen orden, familia, género y especie; fecha de observación; ubicación con GPS, puntos de referencias o nombre local del

lugar; tipo de hábitat; forma de vida; nombre y título del observador y el identificador de la especie; especies asociadas observadas y relaciones;

y fotografías. Columnas adicionales son para comentarios u otros datos interesantes observados, ej.. condiciones del tiempo, hora, actividad,

salud, tamaño, color, y características únicas. La base de datos consolida los datos locales históricos que pueden ser actualizados y usados por

futuros investigadores en sus estudios.

DELGADO-ZAMORA, DAVID A.1, J. Jesús Sánchez-Escalante1 y Denise Zulema Ávila-Jiménez1. 1Department of Scientific Research and

Technology, University of Sonora, Hermosillo, Sonora, Mexico, C.P. 83000, Botanica.son@gmail.com.

Vascular plants of the riparian habitats of El Aribabi Ranch, Municipio de Ímuris, Sonora, Mexico. The state of Sonora, with an area of

184,934 km2,relies on an inventory of vascular plants including as many as 3,659 taxa, but the region defined as ‘The Borderlands’ alone has

an inventory of 1,706 taxa. The aim of the current work is to increase knowledge of the plants of the Borderlands as well as to get to know the

floral diversity of its riparian habitats; the latter through a study in a representative zone (El Aribabi Ranch), chosen for its proximity to the

border and the convergence of different biotic communities. The inventory of documented plants for the riparian habitats of El Aribabi Ranch

adds to a total of 354 taxa, distributed in 78 families and 245 genera; with respect to the number of species, the most well represented families

were Asteraceae (57), Poaceae (31) and Fabaceae (31). The plants present in the riparian habitats of El Aribabi Ranch represent 77% of the

total number of plants of the ranch, and the plant composition seems to be similar to the flora of other rivers in the region such as the San Pedro

(Cananea) and the Santa Cruz River (Santa Cruz).

Plantas vasculares de hábitats ribereños del rancho El Aribabi, municipio de Ímuris, Sonora, México. El estado de Sonora, con una superficie de 184 934 km2

, cuenta con un inventario de plantas vasculares que llega a 3,659 taxones, pero solo la región definida como La Frontera

presenta un inventario de 1,706 taxones. La finalidad del presente trabajo es incrementar el conocimiento sobre las plantas de La Frontera así

como conocer la diversidad florística de sus hábitatsribereños; esto último a través de un estudio en una zona representativa (rancho ElAribabi),

elegida por su cercanía a la frontera y la convergencia de diferentes comunidades bióticas. El inventario de plantas registrado para los hábitats

ribereños del rancho El Aribabi suma un total de 354 taxones, distribuidos en 78 familias y 245 géneros, en cuanto al número de especies, las

familias mayormente representadas fueron Asteraceae (57), Poaceae (31) y Fabaceae (31). Las plantas presentes en los hábitats ribereños del

Aribabi representan un 77% del numero total de plantas para el rancho, y la composición florística parece presentar similitud con las floras de

otros ríos de la región como son el río San Pedro (Cananea) y el río Santa Cruz (Santa Cruz).

DEYO, NICHOLAS S.1 and Edward Gilbert2. 1The Sky Island Alliance, 300 E University Blvd., Suite 270, Tucson, AZ 85705, nick@skyislandalliance.org; 2

Global Institute of Sustainability, Arizona State University, Tempe, AZ 85287, egbot@asu.edu.

Database Workshop: Documenting the biodiversity of the Madrean Archipelago. The Madrean Archipelago Biodiversity Assessment

(MABA) of Sky Island Alliance in Tucson, Arizona, is an ambitious project to document the distributions of all species of animals and plants in

the Madrean Archipelago, focusing particularly on northeastern Sonora and northwestern Chihuahua. The information gathered is made available through MABA’s online database (madrean.org). The sources of these records are museum collections, herbaria, scientific literature, other

databases, observations, and field notes. Many observations come from MABA expeditions to isolated mountain ranges in northeaster Sonora

involving taxonomic specialists, professors, students, and others from universities in Arizona and Sonora; personnel from Mexican agencies

like CEDES and CONANP; and other participants. The databases for flora and fauna are separate; fauna has over 100,000 records and flora has

more than 500,000 records. This workshop is an opportunity to better understand the status of digital plant and animal records in the region and

learn to use the database. The MABA database has a complex structure, allowing searches for scientific records by name, location, collector,

geographical area, etc. Records in the database can be displayed as detailed specimen records, taxonomic lists, or interactive Google maps.

Species lists and identification keys for areas of interest can be created. The database is a powerful resource for researchers, conservationists,

and natural history enthusiasts working in the Madrean Archipelago.

Taller: Documentación de la Biodiversidad del Archipiélago Madrense. El Programa de Evaluación de la Biodiversidad del Archipiélago

Madrense (MABA en inglés) de Sky Island Alliance en Tucson, Arizona, es un proyecto ambicioso para documentar las distribuciones de todas

las especies de animales y plantas en la región de las Islas Serranas, enfocado particularmente en el noreste de Sonora y noroeste de Chihuahua.

Toda la información se captura en la base de datos MABA, accesible en línea (madrean.org). Las fuentes de registros son colecciones en museos

y herbarios, literatura científica, otras bases de datos, observaciones y apuntes de campo. El proyecto MABA realiza expediciones a sierras

aisladas en Sonora con especialistas de distintos taxones, profesores, estudiantes de universidades en Sonora y Arizona; personal de oficinas

de gobierno como CEDES y CONANP; y otros participantes. Las bases de datos de fauna y flora están separadas; fauna tiene más de 100,000

registros y flora tiene más de 500,000 registros. Este taller es una oportunidad para entender mejor el estado de registros digitales de plantas
USDA Forest Service Proceedings RMRS-P-67. 2013 525

Abstracts/ Resúmenes

y animales en la región y el uso de la base de datos. La base de datos MABA tiene una estructura compleja. Se pueden buscar registros por

nombres científicos, localidades, colectores, o áreas geográficas. Los datos se pueden obtener como registros detallados, listas taxonómicas o

mapas interactivos de Google. Se pueden crear listas y claves de identificación de especies por áreas de interés. La base de datos de MABA es

un recurso poderoso para investigadores, conservacionistas, y entusiastas de historia natural trabajando en el Archipiélago Madrense.

DILLON, KRISTEN G.1 and Courtney J. Conway2. 1Arizona Cooperative Fish and Wildlife Research Unit, School of Natural Resources and the

Environment, 325 BioSciences East, University of Arizona, Tucson, AZ 85721, kgdillon@email.arizona.edu; 2U.S. Geological Survey, Arizona

Cooperative Fish and Wildlife Research Unit, School of Natural Resources and the Environment, 325 BioSciences East, University of Arizona,

Tucson, AZ 85721, cconway@usgs.gov.

Effects of temperature and precipitation on annual variation in breeding phenology of Red-faced Warblers in the Santa Catalina Mountains. Increasesin ambient temperatures have caused advancesin breeding date for birdsin NorthAmerica and Europe. However, empirical data

is lacking for evaluating the potential effects of climate change on breeding phenology of birds in the Madrean region where changes in precipitation may be more important than changes in temperature. We examined the effects of temperature and precipitation on breeding phenology of

red-faced warblers over a 10-year period in the Santa Catalina Mountains. May precipitation was negatively associated with breeding date (t=

-4.89, p = <0.001), but annual variation in temperature had no effect. Additionally, we found a positive relationship between nest initiation date

and year, suggesting that red-faced warblers have initiated nesting later in the season over the past ten years (t= 6.20, p= <0.001). These results

are opposite to those reported in many systems, where results have shown nests are initiated earlier over time. This suggests that in our system,

variation in precipitation may be a more important predictor of phenological variation than temperature; birds delay breeding in dry years.

Efectos de la temperatura y precipitación en la variación anual de la fenología de reproducción del chipe cara roja en la Sierra Santa 

Catalina. Los aumentos de la temperatura ambiental han causado adelantos de la fecha de reproducción de las aves en Norteamérica y Europa.

Sin embargo, los datos empíricos no son suficientes para evaluar los efectos del cambio climático en la fenología de las aves en la región Madrense donde los cambios en precipitación podrían ser más importantes que los cambios en temperatura. Analizamos el efecto de la temperatura

y la precipitación en la fenología de la reproducción del chipe cara roja por un periodo de 10 años en la Sierra Santa Catalina. La precipitación

de mayo tuvo un efecto negativo en la fecha de reproducción (t= -4.89, p = <0.001), pero la variación anual de temperatura no impactó. Se

encontró una relación positiva entre la fecha de inicio del nido y el año, lo cual sugiere que el chipe cara roja ha iniciado la anidación más tarde

en los últimos 10 años (t= 6.20, p= <0.001). Estos resultados son opuestos a los encontrados en muchos sistemas, donde los resultados han

mostrado que los nidos se inician más temprano al pasar los años. Esto sugiere que en nuestro sistema, la variación en precipitación puede ser

un vaticinador de la variación de la fenología más importante que la temperatura; las aves demoran la reproducción en años secos.

DUNCAN, DOUG1, and Rob Clarkson2. 1U.S. Fish andWildlife Service, Tucson,AZ 85745. Doug_Duncan@fws.gov; 2 Bureau of Reclamation,

Glendale, AZ 85306. rclarkson@usbr.gov.

Gila River Basin Native Fishes Conservation Program. The Central Arizona Project Gila River Basin Native Fishes Conservation Program

(CAP Program) conserves native fishes and manages against nonnative fishes after several Endangered Species Act consultations between the

Bureau of Reclamation and the U.S. Fish and Wildlife Service on CAP water transfers to the Gila River basin. Populations of some Gila River

native fish species are extremely rare and appear on the verge of extirpation. The CAP Program funds conservation actionsfor five priority fishes,

and other native fishes. A high priority of the CAP Program is to replicate federally listed species into suitable, protected streams and repatriate

populations of listed and other native fishesinto them.Also funded are the control and eradication of non-native fishes. Over $6 million has been

allocated; and Reclamation will fund about $10.5 million to the Service the next 20 years. One half the funding isfor native fish recovery actions,

and one half for non-native actions. Projects are jointly agreed upon by Reclamation and the Service in consultation with the Arizona and New

Mexico Game and Fish Departments. Numerous conservation and recovery projects have been implemented to date, and will be summarized.

Programa de conservación de peces nativos de la cuenca del río Gila. El programa de conservación de peces nativos de la cuenca del río

Gila del Proyecto Arizona Centro (programa CAP por sus siglas en inglés) conserva los peces nativos y controla peces exóticos. Este acuerdo

se tomó después de que el Servicio Federal de Pesca y Vida Silvestre de E.U. (US FWS por sus siglas en inglés) y la Oficina de Rehabilitación

analizaron la Ley de Especies en Peligro de Extinción y el traspaso de agua CAP a la cuenca del río Gila. Las poblaciones de algunas especies

de peces nativos del río Gila son bastante raras y están en grave peligro de ser extirpadas. El programa CAP apoya proyectos de conservación

de cinco especies de peces prioritarios y otros peces nativos. Para el programa CAP es de suma importancia reproducir y reintroducir especies

con categoría de riesgo en la norma federal y otros peces nativos en ríos adecuados y protegidos. También financia el control y erradicación de

peces exóticos. Se han destinado más de $6 millones de dólares y la Oficina de Rehabilitación destinará cerca de $10.5 millones al Servicio (US

FWS) los próximos 20 años. La mitad del financiamiento es para el rescate de peces nativos y la otra mitad para el control de peces exóticos.

Los proyectos son aprobados tanto por la Oficina de Rehabilitación y el Servicio (US FWS) con asesoramiento del Departamento de Caza y

Pesca de Arizona y Nuevo México. Se presentará un resumen de los numerosos proyectos de conservación y rescate realizados.
526 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

DUNCAN, DOUG. U.S. Fish and Wildlife Service, Tucson, AZ 85745. Doug_Duncan@fws.gov.

Gila opminnow interactions with western mosquitofish: an update. It has long been known, and thoroughly documented the western mosquitofish (Gambusia affinis), has major deleterious effects on individual Gila topminnow (Poeciliopsis occidentalis) and their populations. It

has been many years since information on this negative impact has been updated. Therefore, I will list and discuss additional examples where

mosquitofish have impacted topminnow. In addition, it appears that climate change and long-term drought have a synergistic and negative affect on this relationship. Since the last publications detailing the loss of Gila topminnow populations to mosquitofish, two natural topminnow

populations have been lost in the Santa Cruz River basin: Redrock Canyon and Sheehy Spring. Drought and climate change likely assisted

mosquitofish with extirpating these two natural populations. Lastly, climate change may facilitate expanded mosquito populations or expanded

mosquito seasons within the range of the Gila topminnow. Current literature discusses that existing tropical and subtropical diseases, or novel

diseases, may move north into Gila topminnow range. If mosquitoes and diseases they carry become more prevalent, the desire and demand for

vector control, particularly mosquitofish, will increase. I will discuss tools and programs such as Habitat Conservation Plans and Safe Harbor

Agreements that can allow the use of native fish for vector control.

Interacciones del guatopote del Gila con el guayacón mosquito: información reciente. El efecto nocivo del guayacón mosquito (Gambusia 

affinis) sobre el guatopote del Gila (Poeciliopsis occidentalis) y sus poblaciones se conoce desde hace tiempo y se ha documentado en detalle.

Han pasado muchos años desde la última actualización sobre este impacto negativo, por lo que se enumerará y analizarán ejemplos adicionales

donde el guayacón mosquito impactó al guatopote del Gila. Además parece que el cambio climático y la sequía prolongada tienen un impacto

sinérgico y negativo en esta relación. Desde la última publicación donde se describe la pérdida de poblaciones de guatopote del Gila por guayocón

mosquito, otras dos poblaciones naturales de guatopote han desaparecido de la cuenca del río Santa Cruz: Redrock Canyon y Sheehy Spring. Es

probable que la sequía y el cambio climático contribuyeron con el guayacón mosquito a extirpar estas dos poblaciones naturales. Por último, el

cambio climático quizás ayude a aumentarlas poblaciones del guayacón mosquito o alargue su temporada en el área de distribución del guatopote

del Gila. Según la literatura actual las enfermedades tropicales, subtropicales y raras podrían extenderse al norte en el área del guatopote del

Gila. Si los mosquitos y las enfermedades que transmiten se extienden, la necesidad y demanda por control del vector, en especial el guayocón

mosquito, aumentará. Se analizarán métodos y programas como los Planes de Conservación del Hábitat y Compromiso de Propietarios con las

Especies en Peligro de Extinción que pueden permitir el uso de peces nativos para el control de vectores.

EBLE, JEFF1,2, Wallace Meyer III1,2, Saul Macias, Jason Schaller1, Wendy Moore1,2,3. 1Department of Entomology, University of Arizona, 1140

E. South Campus Dr., Forbes 410, Tucson, AZ 85721-0036; 2Center for Insect Science, University of Arizona, 1007 E. Lowell St., P.O. Box

210106, Tucson, AZ 85721-0106; 3Arizona-Sonora Desert Museum, 2021 N. Kinney Road, Tucson, AZ 85743.

Hidden biodiversity of Sky Island arthropods.Tenebrionid and carabid beetles are conspicuous members of the southwest Sky Island invertebrate community and can be found across a range of habitats, including the oak woodlands and pine forests capping many of the region’s

mountains. Beetles living within these high-elevation habitats typically occur on more than one mountain, though populations on different

mountains have likely been isolated from one another since at least the end of the Wisconsin Glacial Period (10,000 ya). We combine traditional

taxonomy and molecular phylogenetics to show that for many species, geographic isolation has restricted dispersal between mountains, leading

to the establishment of independent evolutionary lineages. The prevalence of genetic isolation in the species examined highlights the unique

diversity contained within the Sky Islands, and the threat climate change poses to regional biodiversity.

Biodiversidad de artrópodos escondida en las Islas Serranas. Los escarabajos tenebriónidos y carábidos son notorios miembros de los invertebrados de las Islas Serranas del Suroeste Norteamericano y se pueden encontrar en un amplio rango de hábitats, incluso en los encinales

y pinales de la zona montañosa. Los escarabajos que habitan este rango de elevación típicamente se distribuyen en más de una montaña, sin

embargo las poblaciones de diferentes montañas se aislaron una de otra desde finales del periodo glacial Wisconsin (10,000 años). Se combinaron técnicas taxonómicas tradicionales y filogenética molecular, para demostrar que el asilamiento geográfico ha limitado la dispersión entre

montañas, generando linajes evolutivos independientes. El predominio de la separación genética de las especies resalta la diversidad única en

las Islas Serranas, y la amenaza que el cambio climático representa.

EDWARDS, TAYLOR1,Mercy Vaughn2, Ma. Cristina Meléndez-Torres3, Robert Murphy4, Alice E. Karl5, Philip R. Rosen6, Kristin H. Berry7. 1

University of Arizona Genetics Core, University of Arizona, Tucson, AZ, taylore@email.arizona.edu; 2Paso Robles, CA; 3CEDES (Comisión

de Ecología y Desarrollo Sustentable del Estado de Sonora), Sonora, México; 4Royal Ontario Museum, Toronto, Canada; 5Alice E. Karl and Associates, Davis, CA; 6

School of Natural Resources and the Environment, University of Arizona, Tucson, AZ; 7U.S. Geological Survey, Western

Ecological Research Center, Riverside, CA.

A biogeographic perspective of speciation among desert tortoises in the genus Gopherus. The distribution of organisms we observe in the

Sonoran Desert has been sculpted by the enduring processes of time, climate, and adaptation. One of these species is Morafka’s desert tortoise,

Gopherus morafkai and we are applying a genomic approach to better understand what evolutionary processes were responsible for shaping

diversity in this species. The “Sonoran” desert tortoise exhibits a continuum of genetic similarity spanning 850 km of Sonoran desertscrub,

from Hermosillo, Sonora to Kingman, Arizona. However, at the ecotone between desertscrub and thornscrub we identify a distinct, “Sinaloan”

lineage that suggests a more complex evolutionary story for this species. By using multiple loci from throughout the tortoise genome, we aim

to determine if divergence between these lineages occurred in allopatry, or if there are signatures of past or current introgression. This international, collaborative project will assist state and federal agencies in developing management strategies that best preserve the evolutionary
USDA Forest Service Proceedings RMRS-P-67. 2013 527

Abstracts/ Resúmenes

potential of the desert tortoise. Ultimately, understanding the evolutionary history of desert tortoises will not only clarify the forces that have

driven the divergence in this group, but it will also contribute to our knowledge of the biogeographic history of the southwestern deserts and

how diversity is maintained within them.

Una perspectiva biogeográfica de especiación de la tortuga del desierto en el género Gopherus. La distribución de organismos que observamos en el Desierto Sonorense ha sido esculpida por los procesos perdurables del tiempo, clima y adaptación. Una de estas especies es

la tortuga del desierto Morafka (Gopherus morafkai) e hicimos un estudio genómico para comprender mejor cuales procesos evolucionarios

son responsables de configurar la diversidad de esta especie. La tortuga del desierto “Sonorense” exhibe un continuo de semejanza genética

abarcando 850 km de matorral del Desierto Sonorense, desde Hermosillo, Sonora a Kingman, Arizona. Sin embargo, en el ecotono entre matorral del desierto y matorral espinoso, identificamos un linaje “Sinaloense” distinto, el cual sugiere una historia de evolución más compleja

para esta especie. Por medio del uso de múltiples loci de todo el genoma de la tortuga, estamos tratando de determinar si la divergencia entre

estos linajes ocurrió en una especiación alopátrica o si son indicaciones de introgresiones del pasado o actuales. Este proyecto de colaboración

internacional servirá a las agencias estatales y federales en la planeación estratégica de manejo para conservar el potencial evolucionario de la

tortuga del desierto. Al final el comprender la historia de la evolución de la tortuga del desierto no sólo ayudará a entender las fuerzas que han

impulsado la divergencia en este grupo, sino que también contribuirá a nuestro conocimiento de la historia biogeográfica de los desiertos del

suroeste y como se sostiene la diversidad en ellos.

ENQUIST, CAROLYN A.F. 1,2, Jake F. Weltzin2, and the USA National Phenology Network National Coordination Office Staff. 1The Wildlife

Society, Bethesda, MD 20814, carolyn@usanpn.org; 2USA National Phenology Network, National Coordinating Office, Tucson, AZ 85721,

jweltzin@usgs.gov.

Phenology as a tool for natural resource management and climate change adaptation in the Southwest and beyond. Phenology is widely

accepted as a robust ecological indicator of the impacts of climate change on plants, animals and people. Climate-induced changes in phenology have been linked to shifts in the timing of allergy seasons and cultural festivals, increases in wildfire activity and pest outbreaks, shifts in

species distributions, declines in the abundance of native species, the spread of invasive species, and changes in carbon cycling in forests. The

breadth of these impacts highlights the potential for phenology to inform management and policy-decisions across sectors. Here, we focus on

using phenology as a tool for natural resource management. We address the importance of climate-smart monitoring in an adaptive management

framework and show how phenology data can and have been used in species vulnerability assessments. In turn, we discuss how these efforts

can inform management planning and implementation, such as in identifying on-the-ground adaptation actions related to invasive species,

fire, and biodiversity conservation. Finally, we highlight how a new, off-the-shelf phenology monitoring program, Nature’s Notebook, is both

operationalizing systematic data collection at protected areas across the United States and is cultivating the public’s natural and climate science

literacy by using a participatory citizen science approach.

Aplicación de la fenología en el manejo de recursos naturales y la adaptación al cambio climático en el Suroeste Norteamericano. La

fenología se reconoce como un fuerte indicador ecológico del impacto del cambio climático en las plantas, animales y la gente. Cambios inducidos por el clima en la fenología se han relacionado con cambios en el inicio de temporadas de alergias y festivales culturales, el incremento

en incendios forestales y el brote de plagas, cambio en la distribución de especies, disminución de especies nativas, propagación de especies

invasoras, y cambios en el ciclo del carbono de los bosques. El alcance de estos impactos resalta la importancia de la fenología en el manejo y

decisiones en diferentes sectores. En este estudio nos enfocamos en la aplicación de la fenología en el manejo de recursos naturales. Abordamos

la importancia del monitoreo eficiente y el manejo adaptable y demostramos como los datos de fenología pueden y son usados en la evaluación

de vulnerabilidad de especies.Asimismo se analiza como esta recopilación de datos puede ser útil en el manejo, como el identificar adaptaciones

relacionadas con especies invasoras, incendios, y conservación de la biodiversidad. Finalmente, se resalta cómo un programa nuevo y práctico

de monitoreo de fenología Nature ́s Notebook (Libreta de la Naturaleza), está basado en operación y sistemática para la colección de datos en

áreas protegidas de Estados Unidos y está influenciando la literatura mediante el uso de metodologías participativas de la ciudadanía con un

enfoque científico.

ESPINOSA-FLORES, EUGENIA1, Nalleli Lara-Díaz1, Helí Coronel-Arellano2, Alejandro González-Bernal2 and Carlos A. López-González1. 1

Laboratorio de Zoología, Universidad Autónoma de Querétaro. Av. de las Ciencias S/N, Juriquilla, Santiago de Querétaro, Querétaro, C.P.

76230; 2Naturalia, A. C., Av. 32 entre Calle 3 y 4 Villa Satélite 3250, Agua Prieta, Sonora, C.P. 84200.

Effects of landscape structure on the abundance of black bear (Ursus americanus) in the Sky Islands of Sonora, México. Landscape

characteristics can be important predictors to understand species presence and abundance. Recently, several studies have highlighted the importance of carnivore management at a landscape level because of their large area requirements. The fundamental unit of any given landscape

is the patch, where size and shape are the two main characteristics. Patch size is correlated to the quantity of resources, while shape to external

effects. Between the years 2009-2011 we estimated abundance of black bear (Ursus americanus) in the sky islands of northeastern Sonora using

trap cameras. Patches were defined from the species point of view using the lowest registered altitude obtain during the samples and cover types

that bears preferred in this part of their distribution. Abundance of bears was highly variable across study sites, ranging from 3 to 34 individuals.

We performed a generalized linear model that shows a significant relationship between abundance and patch features (p=0.05), however we

believe that others factors may be affecting them, such as anthropogenic factors. As black bears are considered endangered in México, there is

an urgent need to develop efficient management conservation strategies.
528 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Efectos de la estructura del paisaje en la abundancia del oso negro (Ursus americanus) en las Islas Serranas de Sonora, México. Las

características del paisaje pueden ser indicadoresimportantes de la presencia y abundancia de especies. Estudiosrecientesresaltan la importancia

del manejo de carnívoros a nivel paisaje por su rango de distribución. La unidad fundamental del paisaje es el parche, las características más

importantes son tamaño y forma. El tamaño del parche está relacionado con la cantidad de recursos, mientras que la forma está ligada a efectos

externos. Entre 2009-2011 estimamos la abundancia del oso negro (Ursus ameriacnus) en las islas serranas del noreste de Sonora empleando

cámaras trampa. Los parches fueron definidos empleando la distribución de la especie, se registró la altitud más baja y el tipo de vegetación

donde se distribuye. La abundancia de osos fue muy variable, de 3 a 34 individuos por sitio. Generamos un modelo linear general que demuestra

la relación entre abundancia y parches(p=0.05),sin embargo creemos que hay otrosfactoresinfluyendo losresultados como factores antropogénicos. Dada la situación de peligro del oso negro en México, existe una necesidad urgente de desarrollar estrategias de manejo y conservación.

FAIRCHILD, ELIZABETH, Ries Lindley, and John Scheuring. Arizona Native Plant Society, Box 41206, Sun Station, Tucson, AZ 85717,

ejfairchild@yahoo.com, Ries.Lindley@gmail.com, jfscheuring@hotmail.com.

The plot thickens: Volunteers, organizations, bufflelgrass control and habitat restoration in the Ironwood Forest National Monument, 

Arizona. Arizona Native Plant Society (AZNPS) volunteers have partnered with numerous organizations to effect measurable invasive plant

removal and native vegetation restoration on an 18-acre, heavily disturbed and buffelgrass (Pennisetum ciliare)-infested site in the Waterman

Mountainsin Ironwood Forest National Monument. In 2010 the Bureau of Land Management performed land reshaping to improve water retention and minimize erosion. Volunteers planted native woody species on the worked ground and repeatedly sprayed buffelgrass as it germinated.

By 2011 over 66 different native plant species had emerged including several thousand native woody plants, some from the existing seed bank

and some from seeds planted by volunteers. Due to poor emergence on about three acres, volunteers continue to install rock media-lunas and

manure-amended planting holes. Annual transect data and photo-point images are gathered to track restoration progress. The long-term objective is to restore this site to a natural foothills paloverde upland landscape.

Voluntarios, organizaciones, control de zacate buffel y restauración de hábitats en el área protegida Ironwood Forest National Monument, Arizona, la historia se complica. Voluntarios de laAZNPS (Arizona Natve Plant Society) han colaborado con numerosas organizaciones

para lograr niveles notables de control de plantas invasoras y restaurar la flora nativa en un sitio de 7 hectáreas bastante alterado e infestado

por zacate buffel (Pennisetum ciliare) en la Sierra Waterman de Ironwood Forest National Monument. La Oficina de Administración de Tierras (BLM) en 2010 reconfiguró la topografía para mejorar la retención de agua y disminuir la erosión. Voluntarios plantaron especies leñosas

nativas en la tierra reconfigurada y fumigaron, en repetidas ocasiones, el buffel recién germinado. Para 2011, más de 66 especies distintas de

plantas nativas habían brotado, incluso varios miles de plantas leñosas nativas, algunas de la reserva natural de semillas en el suelo y otras de

semillas sembradas por los voluntarios. Debido a escasa germinación en aproximadamente una hectárea y media, los voluntarios siguieron

instalando medias-lunas de piedra y hoyos para plantar abonadas con estiércol. Se archivan los datos anuales de transectos e imágenes de puntos

fotográficos para monitorear el avance de la restauración. El objetivo a largo plazo es de restaurar este sitio a un paisaje natural de palo verde.

FALK, DONALD A. School of Natural Resources and the Environment, University of Arizona, Tucson AZ 85721.

Are Sky Island ecosystems approaching tipping points? Severe large-scale disturbances can trigger abrupt ecosystem transitions into novel

configurations. These new configurations can be resilient in their new state and resistant to return to pre-disturbance conditions. Such abrupt

transitions are predicted to become more common under conditions of altered climate, in which the regeneration niche and assembly rules for

species and community responses may preclude a simple linear return to the pre-disturbance distribution. In addition to ecological effects, these

transitions also include potentially persistent alterations to geomorphic, soil, hydrological, and biogeochemical systems. Large-scale, severe

disturbance events appear to play a triggering role that releases accumulated tension between bioclimatic envelopes and ecological communities. In several Sky Island mountain ranges, including the Pinaleños, Huachucas, and Chiricahuas, recent large-scale fires may be setting the

stage for such threshold behavior. Fire regimes in other landscapes, particularly the Gila Wilderness, appear to be less departed from the natural

range of variability and hence more resilient even during relatively severe fire seasons. Ecosystem transitions may be viewed as dysfunctional

system behavior, or as a reflection of system adaptation to novel conditions. We explore both scientific and management implications of these

ecosystem dynamics.

¿Están los ecosistemas de las Islas Serranas alcanzando momentos críticos? Los desastres a gran escala pueden detonar abruptas transiciones en el ecosistema ocasionando nuevas configuraciones. Estas nuevas configuraciones pueden estar mejor adaptadas a su nuevo estado

y resistentes para regresar a las condiciones previas. Estos cambios abruptos serán más comunes en condiciones del cambio climático, en

donde la regeneración de nichos y las reglas de asociación durante la reacción de especies y comunidades impedirá un regreso lineal al estado

previo al disturbio. Aunado a los efectos ecológicos, estas transiciones incluyen alteraciones potenciales y persistentes a la geomorfología,

suelo, hidrología y biogeoquímica del sistema. Disturbios severos y de gran escala juegan un papel detonador que libera la tensión acumulada

entre los factores bioclimáticos y ecológicos en las comunidades. En distintas cordilleras de las Islas Serranas como Pinaleños, Huachucas y

Chiricahuas, incendios recientes de gran escala probablemente estén favoreciendo que se llegue al umbral crítico. Los regímenes de fuego en

otros lugares, en especial el área natural Gila, parecen estar más acorde con el rango natural de variabilidad y por tanto su resistencia es mayor

incluso bajo condiciones severas de incendios. Ecosistemas en transición pueden ser percibidos como sistemas disfuncionales, o como el reflejo

de la adaptación del sistema a nuevas condiciones. Exploramos las implicaciones científicas y de manejo de esta dinámica de los ecosistemas.
USDA Forest Service Proceedings RMRS-P-67. 2013 529

Abstracts/ Resúmenes

FEATHERSTONE, ROGER1, Sky Jacobs2, Sergio Avila-Villegas3. 1Director, Arizona Mining Reform Coalition, PO Box 43565, Tucson, AZ

85733, roger@AZminingreform.org; 2Wild Sonora, P.O. Box 508, Tucson, AZ 85702, skyjacobs@wildsonora.com; 3M.S. Sky Island Alliance,

300 E. University Blvd., Tucson, AZ 85705, sergio@skyislandalliance.org.

Wildlife surveys and monitoring with the use of remote camera traps in the Greater Oak Flat Watershed near Superior, Arizona. In

September, 2011 we initiated a two-year camera survey in the Greater Oak Flat Watershed near Superior, Arizona. Our survey area covers a

total of 25 square miles (6,475 Hectares). The area surveyed is primarily a mixing zone of upper Sonoran Desert and interior chaparral, with

some influences from the Madrean vegetation community ranging from 3,800 feet to 4,800 feet of elevation. Ten cameras were deployed in

early October of 2011 and information gathered and analyzed from that date to April 2012. We located cameras primarily in riparian drainages,

occupying a representative sampling of the total area. Locations were chosen as logical wildlife corridors to obtain a representative sampling

of wildlife while allowing relatively easy access to cameras for data collection. To date, we have identified 11 mammalian species, including

bobcat (Lynx rufus), ringtail (Bassariscus astutus), and coati (Nasua narica).

Evaluación y monitoreo de fauna con cámaras remotas del área de Oak Flat cerca de Superior, Arizona. En septiembre de 2011 iniciamos

un proyecto de 2 años de evaluación de fauna silvestre con cámaras remotas en el área Greater Oak Flat Watershed cerca de Superior, Arizona.

Nuestra evaluación abarca un área de 6,475 hectáreas. La zona de monitoreo es principalmente una mezcla de Desierto Sonorense alto y chaparral

interior, con algunas influencias de vegetación Madrense en elevaciones de 1,200 a 1,500 metros. Diez cámaras fueron instaladas a principios

de octubre de 2011 y la información recopilada y analizada cubre hasta abril de 2012. Las cámaras se ubicaron principalmente en drenajes

ribereños, con una muestra representativa de la superficie total. Las localidades escogidas son corredores de vida silvestre y permiten hacer un

muestreo representativo de fauna, además el acceso es relativamente fácil para la recopilación de datos. Hasta la fecha, hemos identificado 11

especies de mamíferos, incluyendo gato montés (Lynx rufus), cacomixtle (Bassariscus astutus) y cholugo (Nasua narica).

FERGUSON, GEORGE M.1,Thomas R. Van Devender2, andAaron D. Flesch3. 1University ofArizona, Tucson,Arizona 85721, USA, georgef@

email.arizona.edu; 2Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717, USA; 3 Avian Science Center, Division of Biological Sciences,

32 Campus Drive, University of Montana, Missoula, Montana 59812.

Biogeography and diversity of pines in the Madrean Archipelago. Pines are important dominants in pine-oak (POF) and mixed-conifer

forest (MCF) in the Sierra Madre Occidental (SMO) and in the Sky Islands in the United States and México. Of the15 native species of pines

that occur in this region, most (11) have affinities to Madrean biotic communities of the SMO in México; four have affinities with Great Basin

and Rocky Mountains biotic communities in the United States. In general the diversity and density of pines increase with the area and elevation

range of mountain ranges. Lower and smaller ranges have 0-4 species; higher ranges (>1 km2 above 2300 m elevation) have 4-7 species. With

11 species in the Municipio de Yécora eastern Sonora, the diversity of pines in the SMO is higher than any Sky Island. Pinus arizonica and P. 

engelmannii are common in POF, while P. strobiformis is typical of MCF. Several species of pines reach the margins of their geographic ranges

in the region, such as Pinus ponderosa var. scopulorum in the Sierra San José near the Arizona border, and P. cembroides and P. lumholtzii in 

the Yécora area; P. yecorensisin the Sierras de Bacadéhuachi and Huachinera are range extensionsfrom the SMO. The Eurasian Pinus sylvestris

is locally introduced in the Sierra de los Ajos.

Biogeografía y diversidad de pinos en el Archipiélago Madrense. Los pinos son importantes especies dominantes en bosques de pino-encino

(BPE) y bosques de coníferas mixtas (BCM) en la Sierra Madre Occidental (SMO) y en las Islas Serranas de Estados Unidos y México. De

las 15 especies de pinos nativos que ocurren en esta región, la mayoría (11) tiene afinidades con las comunidades bióticas Madrenses de la

SMO en México; cuatro tienen afinidades con las comunidades bióticas de la Gran Cuenca y las Montañas Rocallosas de Estados Unidos. En

general la diversidad y densidad de pinos aumenta con el área y el rango de elevación de las cadenas montañosas. Las sierras más bajas y más

pequeñas tienen de 0 a 4 especies; las sierras más altas (>1 km2 arriba de los 2300 m de elevación) tienen de 4 a 7 especies. Las 11 especies en

el municipio de Yécora en el este de Sonora, demuestran que la diversidad de pinos en la SMO es más alta que en cualquier Isla Serrana. Pinus 

arizonica y P. engelmannii son comunes en BPE, mientras que P. strobiformis es típico de BCM. Varias especies de pinos alcanzan los límites

de sus rangos geográficos en la región, como son Pinus ponderosa var. scopulorum en la Sierra San José cerca de la frontera con Arizona, y

P. cembroides y P. lumholtzii en el área de Yécora; los P. yecorensis en las Sierras de Bacadéhuachi y Huachinera son una extensión desde la

SMO. El pino euroasiático Pinus sylvestris está introducido de forma local en la Sierra de los Ajos.

FERNANDEZ, ERIN1, Gerardo Carreón2, Jim Rorabaugh3, Abigail King4, Susan MacVean5, Christa Weise6, Scott Richardson1, Cat Crawford1,

andTara Sprankle7. 1U.S. Fish andWildlife Service, 201 N.BonitaAve Ste 141,Tucson,AZ 85745, erin_fernandez@fws.gov,scott_richardson@

fws.gov, cat_crawford@fws.gov; 2Naturalia, El Cajon #9, Col Santa Fe, Hermosillo, Sonora 83243 MX, conservacion@naturalia.org.mx; 3U.S.

Fish and Wildlife Service, Tucson, AZ (retired), jrorabaugh@hotmail.com; 4Arizona Game and Fish Department, 5000 West Carefree Highway,

Phoenix, AZ 85086. aking@azgfd.gov; 5Arizona Game and Fish Department, 3500 Lake Mary Road, Flagstaff, AZ 86001, Susan_macvean@

azgfd.gov;6U.S. Fish and Wildlife Service, 9300 E. 28th St, Yuma, AZ 85365, christa_weise@fws.gov; 7The Phoenix Zoo, 455 North Galvin

Parkway, Phoenix, AZ 85008, tsprankle@thephxzoo.com.

Building capacity in Mexico through workshops on research, conservation and monitoring Methods for bats and amphibians in Sonora, 

Mexico. There is an ongoing need to train and support managers and biologistsin Sonora to facilitate conservation, monitoring, and management

of the many listed and sensitive species that occur on both sides of the international border. Beginning in 2008, Naturalia, a Sonora-based nongovernmental organization, the U.S. Fish and Wildlife Service, Arizona Game and Fish Department, Bat Conservation International, and The
530 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Phoenix Zoo conducted workshops on bat and amphibian monitoring and conservation at reserves in Sonora managed by Naturalia. Workshops

were regionally specific to encourage local conservation efforts, and were free of charge to participants. Through lectures, demonstrations, and

field exercises conducted in Spanish, workshop participants learned about species identification, study techniques, study design, survey and

monitoring methods, habitat assessment and management, threats to amphibians and bats, and conservation planning. Workshops have been

extremelywellreceived and there is continued interestforfutureworkshops.Overtime, the primary responsibilities ofidentifying and implementing monitoring, protection and conservation activities and holding training workshops for new participants will be shifted to partners that have

previously been trained in our efforts. We will present the evolution of this program and discuss opportunitiesfor future partnersin this program.

Fortalecimiento de capacidades en México con talleres de métodos de investigación, conservación y monitoreo de murciélagos y anfibios 

en Sonora, México. Existe una necesidad continua de entrenamiento y apoyo a personal de áreas protegidas y biólogos de Sonora para facilitar

la conservación, monitoreo y manejo de las varias especies sensitivas y con categoría de riesgo que ocurren en ambos lados de la frontera internacional. Desde 2008 Naturalia (ONG en Sonora), el U.S. Fish and Wildlife Service, el Arizona Game and Fish Department, Bat Conservation

International, y The Phoenix Zoo realizaron talleres sobre monitoreo y conservación de murciélagos y anfibios en reservas de Sonora a cargo

de Naturalia. Los talleres se centraron en la región para fomentar esfuerzos de conservación local y se ofrecieron de forma gratuita. Por medio

de clases, demostraciones y prácticas de campo en español, los participantes aprendieron como identificar especies, desarrollar un estudio, técnicas de investigación, métodos de monitoreo y censo, evaluación del hábitat y manejo, amenazas para anfibios y murciélagos, y como elaborar

un plan de conservación. Los talleres recibieron muy buena aceptación y hay un interés continuo por más. Con el tiempo, la responsabilidad

de identificar y desarrollar las actividades de monitoreo, protección y conservación, así como organizar talleres de capacitación para nuevos

participantes se pasará a los colaboradores entrenados previamente. Se presentará la evolución de estos talleres y se analizarán oportunidades

para socios futuros de este programa.

FFOLLIOTT, PETER F.1, Gerald J. Gottfried2, and Daniel G. Neary3. 1School of Natural Resources and the Environment, University ofArizona,

Tucson, AZ 85251. ffolpete@ag.arizona.edu; 2U.S. Forest Service, Rocky Mountain Research Station, Tonto National Forest, Phoenix, AZ

85006. ggottfried.fs.fed.us; 3U.S. Forest Service, Rocky Mountain Research Station, Flagstaff, AZ 86001.dneary@fs.fed.us.

Soil erosion and deposition before and after fire in oak savannas of the Southwestern Borderlands. Soil erosion and deposition before and

after prescribed burning treatments and awildfire on the hillslopes oftheCascabelWatershedsin the oak savannas oftheSouthwesternBorderlands

is the focus of this paper. Measurements of soil erosion and deposition were analyzed separately because they represented separate processes

of soil movements on the watersheds. The measurements were obtained in the spring and fall to characterize the soil movements following

winter and summer rainstorms, respectively. Parenthetically, all of the burning events on the watersheds were low severity fires. Relationships

between seasonal soil erosion and deposition following the burning events and the rainfall after these events, physiographic characteristics of

the hillslopes, and post-burn vegetation on the watersheds were analyzed for possible cause-and-effect implications. The information presented

should be useful to managers in developing strategies for the introduction of more natural fire regimes into the oak savannas of the region.

Erosión y deposición de suelo antes y después de incendios en pastizales con encino en la zona fronteriza del Suroeste Norteamericano. 

El objetivo de esta investigación es la erosión y deposición de suelos antes y después de quemas prescritas y de un incendio natural en los

cerros de la cuenca Cascabel en el ecosistema pastizal con encino de la zona fronteriza del Suroeste. Se hicieron análisis por separado de las

medidas de erosión y deposición ya que representan procesos diferentes de movimiento de suelo en la cuenca. Las medidas se tomaron durante

la primavera y el otoño para caracterizar el movimiento del suelo después de las lluvias de invierno y de verano respectivamente. A modo de

aclaración, todos fueron incendios de baja intensidad. Interacciones entre erosión y deposición estacional después de las quemas y la precipitación, características fisiográficas de los cerros, y vegetación posterior al fuego fueron analizadas para entender implicaciones causa y efecto.

Esta información puede servir para desarrollar estrategias para introducir regímenes más naturales de fuego en el ecosistema pastizal con encino.

FLESCH,AARON D.1,2, Richard L. Hutto2, Louise Misztal3, and Carlos Gonzáles-Sánchez4. 1School of Natural Resources and the Environment,

University of Arizona, 325 Bioscience East, Tucson AZ 85721, flesch@ag.arizona.edu; 2Avian Science Center, Division of Biological Sciences,

University of Montana, 32Campus Drive, Missoula, MT59812, hutto@mso.umt.edu; 3Sky IslandAlliance, 300 East UniversityBlvd., Suite 270,

Tucson,AZ 85705, louise@skyislandalliance.org; 4Blvd. Luis Encinas y Rosales S/N, Col. Centro, Hermosillo, Sonora, calvin.cali@gmail.com.

Patterns and drivers of bird species distribution in the Madrean Sky Islands of Sonora, México. The distribution and status of breeding

birds in the Madrean Sky Islands of Sonora, México have not been described since the 1950s. Moreover, the effects of local habitat suitability,

habitat area, movement behavior, and interspecific interactions in driving distribution patterns in the region are unknown. To assess the patterns

and drivers of bird distribution and provide information for conservation planning and management, we began a project in spring 2009 that

will continue through summer 2012. During the past three field seasons, we surveyed birds and quantified environmental conditions at over

800 points along 115 transects between 1,170 and 2,600 m elevation in 17 Sky Islands in Sonora. Thus far, we have observed 122 species of

birds that at least presumably breed (48% confirmed breeding) in Madrean environments in the study area including eight species that had not

been observed previously in the region, and an additional six species that possibly breed yet evidence was insufficient to presume so. We will

describe some notable observations while presenting our investigative framework to assess the drivers of distribution across a range of species

that vary in their habitat use, movement behavior, and biogeographic affinity.
USDA Forest Service Proceedings RMRS-P-67. 2013 531

Abstracts/ Resúmenes

Patrones y factores de distribución de aves en las Islas Serranas Madrenses de Sonora, México. La distribución y el estatus de aves que

se reproducen en las Islas Serranas Madrenses de Sonora, México no se ha descrito desde la década de 1950. Por otra parte, se desconocen los

efectos de la condición y área del hábitat local, el comportamiento de movimiento y las interacciones entre especies que impulsan los patrones

de distribución en la región. En la primavera de 2009 para terminar el verano de 2012 se empezó un proyecto con el fin de evaluar los patrones

y factores principales de distribución de pájaros y generar información para la planeación de su conservación y manejo. En las tres temporadas

de trabajo de campo previas se censaron aves y se cuantificaron las condiciones ambientales en más de 800 puntos en 115 transectos entre los

1,170 y 2,600 m de elevación en 17 Islas Serranas de Sonora. Hasta ahora se han observado 122 especies de pájaros que se piensa que se reproducen (48% reproducción confirmada) en ambientes Madrenses en el área de estudio, entre ellas ocho especies que no se habían observado

anteriormente en la región y otras seis especies que posiblemente se reproducen pero aún no se ha confirmado. Se presentará la estrategia de

investigación para evaluar losfactores de distribución de varias especies con diferente uso del hábitat, comportamiento de movimiento y afinidad

biogeográfica, también se describirán observaciones notables.

FONSECA,JULIAand Mike List. Pima County Office of Conservation and Sustainability and Pima County Information Technology, 201 North

Stone, 6th floor, Tucson, Arizona, 85701.

Ephemeral streams and hydroriparian vegetation in Pima County, Arizona. We compared the distribution of riparian vegetation relative to

water resource availability across diverse elevational gradients for a 2.3 million-acre region for the Sonoran Desert Conservation Plan (SDCP).

We analyzed the distribution of riparian vegetation along the watercourses by intersecting watercourse centerlines with vegetation polygons.

Miles of stream length was summed for ephemeral, intermittent and perennial reaches identified in the SDCP inventory per five hydroriparian categories of riparian vegetation. Most of Pima County’s hydroriparian vegetation occurs along stream reaches that classify as ephemeral.

Ninety percent or more of the mesquite woodlands, riparian scrub and riparian strand classifications of the Brown, Lowe, and Pase system are

associated with ephemeral stream reaches in Pima County. Over half of the Sonoran cottonwood-willow vegetation is associated with stream

reaches that classified as ephemeral.Conservation planning has often focused on protecting perennial stream habitat. Our results support the

importance of identifying and protecting ephemeral and intermittent streams, particularly those having shallow groundwater tables. The SDCP

includes a number of provisions to protect or conserve non-perennial stream environments, including refined floodplain management regulations, new land and water infrastructure review guidelines, and land acquisition.

Arroyos efímeros y vegetación riparia en el Condado de Pima, Arizona. Se comparó la distribución de la vegetación riparia dependiendo

de los recursos hídricos disponibles en un gradiente altitudinal para una región de 930 776 ha. dentro del Plan de Conservación del Desierto

Sonorense (SDCP, Sonoran Desert Conservation Plan).Se analizó la distribución de la vegetación riparia a lo largo de lechos de arroyos por

medio de transectos usando el cruce de las líneas centrales con polígonos de vegetación. Se realizaron largos recorridos en zonas efímeras,

intermitentes o perennes tal como lo indica el inventario SDCP para cinco categorías de vegetación riparia. La mayoría de la vegetación riparia

del Condado de Pima se desarrolla a lo largo de arroyos clasificados como efímeros Noventa por ciento o más de mezquital, matorral ripario y

ramal de las clasificaciones de Brown, Lowe y Pase están asociados con sistemas efímeros. Mas de la mitad de la vegetación sonorense de álamo

y sauce está asociada a zonas efímeras.La planeación para la conservación se ha enfocado en arroyos perennes. Nuestros resultados apoyan la

importancia de identificar y proteger arroyos efímeros e intermitentes, especialmente aquellos con acuíferos someros. La SDCP incluye un gran

número de disposiciones para proteger y conservar zonas ambientales de arroyos no perennes, incluyendo zonas de inundación, adquisición de

tierras e infraestructura y revisión de lineamientos.

FONSECA, JULIA, Pima County Office of Conservation and Sustainability, 201 North Stone, 6th floor, Tucson, Arizona, 85701.

Restoration at Bingham Ciénega, Pima County, Arizona. Bingham Ciénega is located on a 285-acre preserve just north of the town of Redington in the San Pedro River valley. It is owned by the Pima County Regional Flood Control District (District) and managed by The Nature

Conservancy (TNC). During the last century, about 50 acres of the site, including much of the ciénega, were grazed and farmed. Long-time

residents report that the spring and its ciénega has varied considerably in discharge and extent over time, and in 1957, most of the ciénega was

drained and cultivated. After acquiring the site, we began a program of study and analysis, while allowing passive restoration via drainage ditch

abandonment. After 1989, the ciénega expanded in size and became increasingly vegetated with bulrush, cattail, ash, buttonbush and willow. In

1998, an active restoration began, based upon depth-to-water planting zones within the fields, and historical vegetation research. Three planting

zones were identified; deciduous riparian woodland, sacaton grassland, and mesquite woodland, each zone marked by progressively deepening

water levels. Three years later, the Nature Conservancy revegetated 25 acres of farmland using Arizona Water Protection and Flood Control

District funding. Since 1998, water levels have deepened. Spring flow completely ceased in 2003 and has only briefly resumed. The naturally

restored bulrush marsh to the point is now dominated by annuals, though some willow and buttonbush remain. The actively restored sacaton has

been reduced in some area, with the best cover in the year 1 field. A mix of mesquite and sacaton, similar to the natural condition of the former

ciénega outflow, may be the long-term outcome for the ciénega margins. Fire management has been and remains an important focus of effort.

Restauración de la Ciénega Bingham, Pima County, Arizona. La Ciénega Bingham se localiza en una reserva de 115 ha justo al norte

de Redington en el valle del río San Pedro. Pertenece al Distrito de Control de Inundaciones del Condado Pima (Distrito) y la administra la

Conservación de la Naturaleza (Nature Conservancy, TNC). Durante el siglo pasado, aproximadamente 20 ha, incluyendo la ciénega, fueron

utilizadas para ganadería y agricultura. Residentes de antaño reportan que el manantial y la ciénega han variado considerablemente en descarga

y extensión, en 1957, casi toda la ciénega fue drenada y cultivada. Después de adquirir el sitio comenzamos una serie de estudios y análisis,
532 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

permitiendo la restauración pasiva vía abandono de la zanja de drenaje. Después de 1989, la ciénega se expandió y la cubierta vegetal se vio

enriquecida con juncos (Scirpus sp.), tule (Typha domingensis), fresno (Fraxinus velutina), mimbro (Cephalanthus sp.), y sauce (Salís sp.).

El proceso de restauración activa comenzó en 1998, se establecieron zonas de plantación basadas en la profundidad del agua y se investigó la

vegetación histórica. Se identificaron tres zonas para plantación; bosque ribereño caducifolio, pastizal de zacatón, y mezquital; cada zona identificada conforme aumenta la profundidad del agua. Tres años después, Conservación de la Naturaleza reforestó 10 ha con fondos del Distrito

El nivel del agua ha ido bajando desde 1998. El flujo del manantial cesó por completo en el 2003. La zona restaurada de forma natural con

juncos está dominada por anuales con pocos sauces y mimbros. El zacatón se ha visto reducido en algunas zonas. Una mezcla de mezquite y

zacatón, similar a la de las condiciones naturales de la ciénega, puede ser el resultado a largo plazo. El manejo de incendios fue y sigue siendo

un proceso importante de éstos trabajos.

FRANKLIN, KIMBERLY1,3, Wallace M Meyer III1,2, Jeff Eble1,2, Richard C Brusca3, and Wendy Moore1,2,3. 1Department of Entomology, University of Arizona, 1140 E. South Campus Dr., Forbes 410, Tucson, AZ 85721-0036; 2

Center for Insect Science, University of Arizona, 1007 E.

Lowell St., P.O. Box 210106, Tucson, AZ 85721-0106; 3Arizona-Sonora Desert Museum, 2021 N. Kinney Rd., Tucson, AZ 85743.

Diversity and species composition of ant assemblages in the Santa Catalina Mountains. Ants are a dominant faunal group in nearly all terrestrial ecosystems and play important roles in numerous ecosystem processes. In North America, ant diversity peaks in the desert Southwest,

and exceptionally high levels of diversity can be found in the Sky Island region. For example, over 180 species have been documented in the

Chiricahua Mountains of southeastern Arizona. Despite exceptional levels of diversity and great functional importance, ant assemblages of

the sky islands remain poorly studied. We have developed a long-term sampling protocol designed to document and monitor ground-dwelling

arthropod communities in the Santa Catalina Mountains. The design consists of 100 meter transects with 10 pitfall traps deployed at each of

66 sites that span the elevational gradient. Preliminary analysis of the ant data indicates that the diversity of ant assemblages decreases with

elevation, and species turnover across the elevational gradient is high. Transitions between ant assemblages across elevation appear to occur in

congruence with transitions among plant biomes. Understanding which factors structure ant assemblages across elevation gradients in the sky

islands will be essential to predicting the impact of climate change on ant diversity and on the ecosystem services they provide.

Diversidad y composición de especies de los ensambles de hormigas en la Sierra Santa Catalina. Las hormigas son un grupo de fauna

dominante en casi todos los ecosistemas terrestres y desempeñan un papel importante en numerosos procesos del ecosistema. En América del

Norte, la diversidad de hormigas alcanza su máxima diversidad en el Suroeste árido de Estados Unidos y niveles de diversidad bastante altos se

encuentran en la región de las Islas Serranas. Por ejemplo, más de 180 especies se han registrado en la Sierra Chiricahua del sureste de Arizona.

A pesar de los altos niveles de diversidad y su gran importancia funcional, los ensambles de hormigas de las islas serranas aún son muy poco

estudiados. Hemos desarrollado un protocolo de muestreo a largo plazo diseñado para documentar y monitorear las comunidades de artrópodos

que viven en la tierra en la Sierra Santa Catalina. El diseño consiste de transectos de 100 metros con 10 trampas de caída en cada uno de los

66 sitios de muestreo que abarca el gradiente de elevación. Los análisis preliminares de datos indican que la diversidad de los ensambles de

hormigas disminuye con la elevación y que el movimiento de especies es alto en el gradiente de elevación. Las transiciones entre los ensambles

de hormigas en los diferentes rangos de elevación parecen ocurrir en relación con las transiciones entre los biomas vegetales. El entender cuales

factores influyen los ensambles de hormigas en los gradientes de elevación de las islas serranas será fundamental para predecir el impacto del

cambio climático en la diversidad de hormigas y los servicios ecosistémicos que proporcionan.

FREDERICK, GLENN P.1, Brooke S. Gebow2, and Thomas R. Jones3. 1USDA Forest Service, Coronado National Forest, Sierra Vista Ranger

District, 5990 S. Highway 92, Hereford, AZ 85615; 2The Nature Conservancy, 27 Ramsey Canyon Road, Hereford, AZ 85615; 3Arizona Game

and Fish Department, 5000 W. Carefree Hwy, Phoenix, AZ 85086.

Reestablishment and protection of a Chiricahua Leopard Frog population in Scotia Canyon, Huachuca Mountains, Arizona. Bullfrogs

are significant impediments to recovery of native aquatic amphibians and reptiles in riparian systems in the American Southwest. Until recently,

bullfrog control across a landscape with many stock ponds was considered unlikely to succeed. In 2008, however, the Coronado National Forest

launched a multi-year, multi-partner project to eliminate bullfrogs within a 113 km2 area centered on Scotia Canyon in the Huachuca Mountains.

We eliminated all bullfrogs in Scotia Canyon and reestablished Chiricahua leopard frogs. Bullfrogs were removed at 10 stockponds and along

3.7 km of stream within 6 miles of Scotia Canyon. We focused control efforts at perennial ponds and 130-acre Parker Canyon Lake. We found

adult bullfrogs in the Cave Canyon watershed during the summer monsoon season traveling to headwaters with direct connections to Scotia

Canyon. Bullfrogs reinvaded some stock ponds but not Scotia Canyon, where leopard frogs have continued to persist. The Forest Service in

cooperation with the livestock grazing permittee has developed plans to manage stock ponds to disrupt bullfrog dispersal and control their

reproduction. How to simultaneously maintain multiple stock ponds for livestock and native wildlife without perpetuating invasive species is

a management challenge that is likely to increase in complexity with climate change.

Reintroducción y protección de la población de la rana de Chiricahua en Scotia Canyon, Sierra Huachuca, Arizona. Las ranas toro

impiden de manera significativa la recuperación de anfibios y reptiles en los sistemas ribereños del Suroeste Norteamericano. Hasta hace poco

el control de ranas toro en una región con muchos represos de abrevadero se consideraba imposible. Sin embargo en el 2008, el Bosque Nacional Coronado empezó un proyecto de varios años junto con otras instituciones, para eliminar las ranas toro en 113 km2 en Scotia Canyon de

la cordillera Huachuca. Se eliminaron todas las ranas toro de Scotia Canyon y se reintrodujo la rana de Chiricachua. Se eliminaron las ranas

toro de 10 represos y de un tramo de 3.7 km de un arroyo cerca de 10 km de Scotia Canyon. Los trabajos de control se enfocaron en estanques
USDA Forest Service Proceedings RMRS-P-67. 2013 533

Abstracts/ Resúmenes

permanentes y 53 has de Parker Canyon Lake. Se encontraron ranas toro adultas en la subcuenca Cave Canyon durante la estación del monzón

de verano, trasladándose al nacimiento de arroyos con conecciones directas a Scotia Canyon. Las ranas toro reinvadieron algunos represos

pero no Scotia Canyon, donde la rana de Chiricahua aún persiste. El Servicio Forestal a través de su programa de permisos para el pastoreo

de ganado ha desarrollado un programa para manejar los represos para bloquear la dispersión y reproducción de la rana toro. Es un gran reto

mantener simultáneamente múltiples represos para el ganado y fauna silvestre sin perpetuar especies invasoras y este problema probablemente

aumentará bajo las influencias del cambio climático.

GARFIN, GREGG1 and Adrian Quijada-Mascareñas1. 1School of Natural Resources and the Environment, University of Arizona, Tucson, AZ

85721, gmgarfin@email.arizona.edu, aquijada@cals.arizona.edu.

Transborder Climate and experiments in climate communication: Building a foundation for adaptation.In recent years,U.S.-Mexico border

region stakeholders have consistently articulated needs for climate services to provide information, enhance preparedness, and better inform

resource management decisions. We report on a NOAA-funded initiative in the Madrean Archipelago region to not only provide information,

but to build capacity for adaptation. The initiative augments the Border Climate Summary/Resumen del Clima de la Frontera PDF newsletter,

through experiments in climate communication. We focus on Transborder Climate, a brief bulletin that gives a high-level overview of climate

forecasts and research. We also report on experiments in the use of bilingual webinars and social media to increase knowledge exchange and

build a foundation for adaptation to climate change. These experiments are being conducted by a consortium of collaborators from University

of Arizona, UNISON, CIBNOR, CICESE, and the Southern Regional Climate Center. We report on the early stages of these experiments and

the methods and metrics for evaluating the use of these media and their value to border region decision makers.

Clima transfronterizo y programas piloto de comunicación sobre clima: estableciendo bases para la adaptación. Los últimos años, los

sectores interesados de la región fronteriza entre México y E.U. han expresado la necesidad de que el servicio climatológico proporcione mayor

información, medidas de seguridad y fuentes de información útiles en los programas de manejo. Presentamos el proyecto NOAA de la región

del Archipiélago Madrense, que además de informar explora estrategias de adaptación. El proyecto enriquece el boletín en línea “Resumen del

clima de la frontera / Border Climate Summary”, con programas sobre comunicación del clima. Presentamos Clima Transfronterizo, un boletín

con reseñas de investigación y pronóstico de clima. También introducimos el uso de cursos y seminarios bilingües en línea y redes sociales

para facilitar el intercambio de ideas y establecer bases de adaptación al cambio climático. Estos programas piloto se llevan acabo gracias a

la colaboración de University of Arizona, UNISON, CIBNOR,CIECESE y Southern Regional Climate Center. Se presenta el inicio de estos

programas, la metodología y medidas de evaluación del uso de medios electrónicos y su importancia para los directivos de la región fronteriza.

GEBOW, BROOKE1,2, Donald A. Falk3, Christopher Stetson2, and Corrine Dolan3. 1The Nature Conservancy, Southeastern Arizona Preserves,

27 Ramsey Canyon Road, Hereford AZ 85615, bgebow@tnc.org; 2Coronado National Forest, 300 W. Congress Street, Tucson AZ 85701,

chstetson@fs.fed.us; 3School of Natural Resources and the Environment, Biological Sciences East, University of Arizona, Tucson AZ 85721,

dafalk@email.arizona.edu, cdolan@ag.arizona.edu.

FireScape: A program for whole-mountain fire management in the Sky Island Region. Poster.The Coronado National Forest’s (CNF)

FireScape program works to remove barriers to fire playing its natural role on the landscape. A long-term goal is creating landscapes that are

able to survive wildfire with biodiversity intact, especially important in the face of a drier, hotter Southwest. The FireScape team is nurturing

multiple efforts around the Sky Islands—no two projects are alike, but those underway share an approach that includes multiple jurisdictions,

investigations by University of Arizona scientists, public engagement, assessing treatment need at the whole-mountain scale, and creatively

removing implementation barriers when funding is scarce. Clearance for treatments in designated Wilderness is a need across all projects. The

FireScape team has completed a fuels map and an analysis of departure from reference condition for southeastern Arizona that covers the CNF

and partner lands. Partners have worked together to update Landfire data for this 14 million-acre area. These products provide inputs for fire

behavior and effects analyses to support decision-making and outreach. The website www.azfirescape.org is a work in progress that displays

project information, including maps, reports, and vegetation and fuels data.

FireScape: Un programa para el manejo integral de incendios en las Islas Serranas. El programa FireScape del Coronado National Forest

(CNF, Bosque Nacional Coronado) trabaja para quitar barreras que impiden al fuego desempeñar su papel natural en el paisaje. La meta a largo

plazo es crear paisajes capaces de sobrevivir incendios naturalessin afectar la biodiversidad, especialmente ante un escenario seco y más caliente

en el Suroeste Norteamericano. El personal de FireScape está desarrollando varios trabajos en las Islas Serranas, ninguno de los proyectos es

similar, pero los que están en marcha se semejan al incluir varias jurisdicciones, investigaciones científicas de la University of Arizona, participación de la ciudadanía, evaluación de tratamientos en toda la montaña, y creatividad para remover las barreras con presupuesto escaso. En

áreas naturales protegidas se necesita autorización en cualquier proyecto. El personal de FireScape elaboró mapas de áreas propensas a incendios y análisis de puntos de ignición de referencia para el sureste de Arizona, que cubre el CNF y zonas cercanas. Los socios colaboraron para

actualizar la información de estos 5.7 millones de ha. Estos mapas y reportes aportan información sobre el comportamiento del fuego y análisis

que sirven en las decisiones de manejo e información a la ciudadanía. La pagina electrónica www.azfirescape.org se actualiza constantemente

para presentar información de proyectos, mapas, reportes y datos de vegetación y combustible.
534 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

GIBSON, CARLY1, Marc Stamer2, andAnnYouberg3. 1U.S. Forest Service, Stanislaus National Forest, Sonora,CA, 95370; 2U.S. Forest Service,

Mountaintop Ranger District, Fawnskin, CA, 92333; 3Arizona Geological Society, Tucson, AZ, 85701.

Hillslope treatment effectiveness monitoring on Horseshoe 2 and Monument Fires. In 2011, the Coronado National Forest experienced one

of the most active fire seasons in recorded history when fires burned over 360,000 acres between February and July. Burned Area Emergency

Response (BAER) Teams evaluated post-fire watershed conditions and prescribed treatments based on threats to known values at risk. Hillslope

stabilization treatments were prescribed and implemented for areas of high soil burn severity on both Horseshoe 2 and Monument Fires. These

treatments consisted of seeding on the Horseshoe 2 Fire and application of agricultural straw mulch and seed on the Monument Fire. Based on

initial monitoring results one of three seeded species (Hordeum vulgare) emerged in both burned areas, slightly improving effective ground

cover in both treatments. However, seeding treatments failed to meet monitoring success criteria for both fires. Hillslope erosion was reduced

where mulch treatment was applied correctly and where slopes were gentle on the Monument Fire, and appeared to contribute to seeded species

cover. In the Horseshoe 2 Fire hillslope erosion was high in the treatment units and was not sufficiently reduced by seeding alone. A need for

additional monitoring in spring 2012 exists and would improve the current understanding of the effectiveness of hillslope treatments.

Monitoreo de la eficacia de tratamiento en laderas de los incendios Horseshoe 2 y Monument. El Coronado National Forest experimentó

en el 2011 una de las temporadas más activas de incendios forestales en la historia, más de 145,686 ha fueron afectadas entre febrero y julio.

Las brigadas de BAER (siglas en inglés de brigadas de emergencia para incendios) evaluaron la situación posterior al fuego en la cuenca y

prescribieron tratamientos basados en amenazas a recursos en riesgo. Los tratamientos para la estabilización de laderas fueron recomendados e

implementados en área donde se detectó suelo con incendios severos en Horseshoe 2 y Monument. En Horseshoe 2 el tratamiento que se aplicó

fue siembra mientras que en Monument se aplicó un mantillo de paja y siembra. Los monitores preliminares demuestran que una de las tres

especies sembradas (Hordeum vulgare) germinó en las dos zonas afectadas, mejorando ligeramente la cobertura vegetal en ambos tratamientos.

Sin embargo, el tratamiento de siembra no cumple con las expectativas de éxito en ninguno de lossitios. La erosión en laderas disminuyó cuando

el tratamiento de mantillos se aplicó correctamente y con pendientes ligeras en el incendio Monument, ésto contribuyó a la cobertura vegetal de

especies sembradas. En Horseshoe 2 la erosión fue mayor en las áreas tratadas y el sembrado sin mantillo no la disminuyó. Se requieren más

monitoreos en la primavera de 2012 para mejorar nuestro entendimiento de la eficacia de tratamiento de laderas.

GONZÁLEZ-BERNAL, ALEJANDRO1, Helí Coronel-Arellano1, Nalleli E. Lara-Diaz2, Carlos A. López-González2. 1Naturalia A.C. Av. 32

entre Calle 3 y 4 Villa Satélite 3250, Agua Prieta, Sonora, C.P. 84200; 2Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro.

Av. de las Ciencias S/N, Col. Juriquilla, Santiago de Querétaro, Querétaro, C.P. 76230.

Livestock depredation occurrence by mountain lions (Puma concolor) in the Sierra San Luis, Sonora. The ability to predict the occurrence

of wildlife-livestock conflicts would benefit both the livestock producers and the predators that prey upon them. Little has been done to reduce

this conflict using modeling tools. As such, our aim for this study was to create a depredation risk model for livestock in the Sierra San Luis,

Sonora. We modeled the probability of occurrence of pumas and livestock using program PRESENCE. Covariatesinvolved in the model include

elevation, slope, aspect, NDVI. Point locations for pumas and livestock were obtained using camera trap surveys obtained between 2009 and

2011. Using ArcGIS we combined the occurrence of pumas and livestock and developed a low to high probability of depredation where areas

of overlap occur. We considered the highest probability of depredation to occur at piedmont, rugged areas, and closed cover. The results from

this study can be used to improve livestock management and to recommend livestock rotation in the area.

Depredación de ganado por pumas (Puma concolor) en la Sierra San Luis Sonora. La habilidad de predecir conflictos entre fauna silvestre

y ganado puede ser útil tanto a los productores de ganado como a los predadores. Se ha hecho muy poco para resolver este conflicto empleando

modelos computarizados. El objetivo de este estudio fue generar un modelo de riesgo de depredación en la Sierra San Luis, Sonora. Modelamos

la probabilidad de encuentros entre pumas y ganado empleando el programa PRESENCE. Las covarianzas usadas en el modelo son elevación,

pendiente, aspecto, NDVI. Localidades puntuales para pumas y ganado se obtuvieron empleando cámarastrampa entre 2009 y 2011. Empleando

ArcGis combinamos la presencia de pumas y ganado y desarrollamos la probabilidad de depredación de menor a mayor en las áreas donde

coexisten. Consideramos que la mayor probabilidad de depredación existe en el piedemonte, zonas escarpadas y de cobertura cerrada. Los

resultados pueden ser empleados para mejorar el manejo de ganado y recomendar la rotación del ganado en el área.

GONZÁLEZ-ELIZONDO, M. S., M. González-Elizondo, L. Ruacho-González, I. L. López-Enríquez, F. I. Retana-Rentería, and J. A. TenaFlores. CIIDIR I.P.N., Sigma 119 Fracc. 20 de Noviembre II, Durango, Durango, 34220, Mexico, herbario_ciidir@yahoo.com.mx.

Ecosystems and diversity of the Sierra Madre Occidental. The ecosystems and vegetation of the Sierra Madre Occidental (SMO) were

mapped usingArcView, based on Landsat images and field verification. Data on the composition, distribution and ecological determinism of the

vegetation are presented. The Sierra Madre Occidental is the biggest continuous ignimbrite plate on Earth. With a complex geological history

and a high biological and cultural diversity, it is a biological corridor, a barrier for the surrounding elements and an active center of speciation,

yet ecologically not well known. We describe the vegetation of the SMO above 1,800 m asl. Ecosystems are described along a gradient from

tropical deciduous forests to high montane communities. The most widespread communities are the Madrean pine-oak forests and woodlands.

Among the dramatic changes occurring—besides fragmentation and deforestation driven by humans—are the effects of bark beetle (Dendroctonus) infestations that have killed extensive areas of pines (and also affected fir, Douglas fir and spruce), which were likely already stressed

by drought. Other changes included the expansion of chaparral driven by disturbance and the dwindling of oak woodlands, which are being

replaced by Dodonaea viscosa, an invader from warmer areas.
USDA Forest Service Proceedings RMRS-P-67. 2013 535

Abstracts/ Resúmenes

Diversidad y ecosistemas de la Sierra Madre Occidental. Los ecosistemas y la vegetación de la Sierra Madre Occidental (SMO) fueron

cartografiados usando ArcView, con base en imágenes Landsat y verificación de campo. Se presentan datos sobre la composición, distribución

y determinismo ecológico de su vegetación. La Sierra Madre Occidental es la placa continua de ignimbrita más grande sobre la Tierra. Con una

compleja historia geológica y una gran diversidad biológica y cultural, la SMO es un corredor biológico, una barrera para los elementos que

la rodean y un centro activo de especiación; sin embargo, biológica y ecológicamente todavía no se conoce bien. En este trabajo se describe la

vegetación de la SMO por encima de los 1,800 m snm a lo largo de un gradiente desde los bosques tropicales deciduos hasta las comunidades

de alta montaña. Las comunidades más ampliamente distribuidas son los bosques madrenses de pino-encino y los bosques bajos abiertos. Entre

los dramáticos cambios que están ocurriendo—además de la fragmentación y la deforestación antropogénicas—están los efectos de las infestaciones por descortezadores (Dendroctonus) que han destruido extensas áreas de pinares (y afectado también a Abies, Pseudotsuga y Picea),

previamente estresados y debilitados por sequía. Otros cambios incluyen la expansión de chaparrales favorecidos por disturbio y la reducción

de encinares, los cuales están siendo reemplazados por Dodonaea viscosa, un arbusto invasor proveniente de áreas más cálidas.

GOTTFRIED, GERALD J.1 and Peter F. Ffolliott2. 1 U.S. Forest Service, Rocky Mountain Research Station, Tonto National Forest, 2324 E.

McDowell Road, Phoenix,AZ 85008, ggottfried@fs.fed.us; 2School of Natural Resources and the Environment, University ofArizona, Tucson,

AZ 85721, ffolpete@ ag.arizona.edu.

Ecology and management of oak woodlands and savannas in the southwestern borderlands region. Management of the Madrean oak

woodlands and the less dense and ecologically different oak savannas has become more complicated in recent years and, therefore, must be

based on sound ecological information. However, relatively little is known about the Madrean oak ecosystems in spite of the fact that they cover

about 80,000 km2 in the southwestern United States and northern Mexico. Emory oak (Quercus emoryi) is the dominant tree in most stands and

is usually associated with other oak species and juniper. Trees are utilized for firewood, fence posts, and construction materials and acorns are

gathered as food by local inhabitants. The woodlands and savannas are important for wildlife habitat, including habitats for listed and sensitive

species, watershed protection, and livestock grazing. Tree regeneration and water consumption are important considerations. Tree harvesting has

been restricted because of heavy utilization in the past but coppice management could ease the supply situation. Recreational demands and fire

management concerns are increasing as populations grow in southern Arizona. This presentation reviews the ecology and management of oak

woodlands and savannas based on recent research findings along the border and relevant literature from the United States and northern Mexico.

Ecología y manejo de encinales y pastizales en la zona fronteriza del Suroeste Norteamericano. En el Archipiélago Madrense el manejo

del encinal y del pastizal con encino, menos denso y con ecología diferente, se ha complicado en los últimos años, por lo tanto debe basarse en

información ecológica sólida. Sin embargo existe muy poca información de estos ecosistemas a pesar de que cubren 80,000 km2 del suroeste de

Estados Unidos y noroeste de México. La bellota (Quercus emoryi) por lo general es el árbol dominante del arbolado y comúnmente asociado

con otras especies de encino y táscate. Los árboles son utilizados como leña, postes y materiales de construcción, y el fruto es colectado para

alimento por lugareños. El encinal y el pastizal son importantes hábitat para animales, incluyendo especies sensibles y en peligro, protección

de la cuenca y pastoreo. La regeneración de árboles y consumo de agua son factores importantes. La extracción de madera se ha prohibido

debido a la explotación que se dio en el pasado, sin embargo los planes de manejo pudieran ayudar en esta situación. Las demandas por zonas

recreativas y el manejo de fuego son problemas cada vez mayores debido al aumento de la población en el sureste de Arizona. Esta presentación

revisa la ecología y manejo de ecosistemas de encino basados en estudios en zonas fronterizas y literatura de Estados Unidos y norte de México.

GOTTFRIED, GERALD J.1, Peter F. Ffolliott2, and Daniel G. Neary3. 1U.S. Forest Service, Rocky Mountain Research Station, 2324 E. McDowell Road, Phoenix, AZ 85006, ggottfried@fs.fed.us; 2

School of Natural Resources and the Environment, University of Arizona, Tucson,

AZ 85251, ffolpete@cals.arizona.edu; 3U.S. Forest Service, Rocky Mountain Research Station, 2500 S. Pine Knoll Drive, Flagstaff, AZ 86001,

dneary@fs.fed.us.

Effects of prescribed fire and wildfire on biological resources of oak savannas in the Peloncillo Mountains, New Mexico. Private and

public land managers are attempting to reintroduce fire into the ecosystems of the Peloncillo Mountains in order to reduce the density of woody

species, increase the herbaceous plant cover, and improve the area’s ecological diversity. The Cascabel Watershed Study, which includes 12

small, gauged watersheds, was started in 2000 to evaluate the impacts of cool-season (November-April) and warm-season (May-October)

prescribed fires on a number of physical and biological components of the oak (Quercus spp.) savannas that are common to the Southwestern

Borderlands Region. A wildfire in 2008 modified the original experimental design. All three fire treatments resulted in low severity burns. The

effects of the different fire treatments on tree overstories, herbaceous production, ground cover,selected wildlife species and on bird populations

are presented. In general, measurements did not indicate many significant differences among treatments or with pre-fire conditions. Almost 78

percent of the overstory trees on the 12 watersheds, for example, survived the fires. The main exception was a large increase in grass and total

herbaceous plant production of early-growing species after the fires. The fires also produced statistically significant increases in the production

of late-growing grass species but the changes were not as large.

Efectos de quemas prescritas e incendios naturales en los recursos biológicos del pastizal de encino en las montañas Peloncillo, Nuevo 

México. Personal de manejo de recursos de áreas naturales del gobierno y privadas de las montañas Peloncillo están intentando reintroducir

el fuego en el ecosistema con la finalidad de reducir la densidad de especies leñosas, incrementar la cobertura vegetal herbácea y mejorar la

diversidad biológica del área. El estudio de la Cuenca Cascabel, que incluye 12 subcuencas con calibradores,se comenzó en el 2000 para evaluar

el impacto de quemas prescritas en las estaciones frías (noviembre-abril) y cálidas (mayo-octubre) sobre componentes físicos y biológicos del
536 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

pastizal con encino (Quercus spp.) común en la zona fronteriza del Suroeste Norteamericano. En el 2008 un incendio natural modificó el diseño

experimental inicial. Las tres quemas prescritas fueron de baja intensidad. Se presenta el efecto de los tratamientos con fuego sobre el dosel

arbóreo, producción herbácea, cubierta vegetal, especies selectas de fauna y poblaciones de aves. En general los datos no muestran diferencias

significativas entre tratamientos o condiciones previas al incendio. Casi el 78% del dosel arbóreo en las 12 subcuencassobrevivieron las quemas.

La excepción fue un gran aumento de zacates y producción total de herbáceas de especies tempraneras después de los incendios. Los incendios

también causaron aumentos estadísticos significativas de zacates de crecimiento tardío, sin embargo los cambios no fueron tan grandes.

GRAY, KATHERINE M. and Robert J. Steidl. School of Natural Resources and the Environment, University of Arizona, Tucson AZ, 85721,

graykat@email.arizona.edu, steidl@ag.arizona.edu.

Effects of buffelgrass on habitat use of Sonoran Desert tortoises. Buffelgrass (Pennisetum ciliare) is a nonnative grass that is increasing

markedly in distribution throughout the southwestern United States and Mexico. By altering ecosystem structure and function, buffelgrass has

the potential to alter the quantity and quality of habitat for animalssuch asthe Sonoran desert tortoise (Gopherus agassizii), which inhabits areas

being invaded by buffelgrass. We examined effects of buffelgrass on habitat use of desert tortoises by surveying tortoises on 50 4-ha plots that

we established across the gradient of buffelgrass cover in Saguaro National Park in 2010 and 2011. On each plot, we contrasted characteristics

of vegetation and substrate in 5-m-radius plot centered on 186 tortoises with areas thought available to tortoises as measured along a systematic

array of transects established across each plot. Our results will help biologists understand the threat that buffelgrass poses to this rare vertebrate

in the southwestern United States.

Efectos del zacate buffel en el hábitat de la tortuga del desierto Sonorense. El zacate buffel (Pennisetum ciliare) es un pasto introducido

que está notablemente aumentando su distribución en el suroeste de Estados Unidos y noroeste de México. El zacate buffel tiene el potencial

de alterar la cantidad y calidad del hábitat cambiando la estructura y función del ecosistema, afectando a especies como la tortuga del desierto

Sonorense (Gopherus agassizii), ya que vive en zonas donde el buffel está invadiendo. Examinamos el efecto del buffel en hábitat de la tortuga

del desierto mediante un muestreo en 50 cuadrantes de 4 ha en un gradiente con buffel en el Parque Nacional Saguaro durante el 2010 y 2011.

En cada cuadrante comparamos características de la vegetación y el substrato en cuadrantes de 5 m de radio centrados en 186 tortugas, con áreas

probables de tortugas, conforme a los datos de transectos de arreglo sistemático en cada cuadrante. Los resultados servirán para comprender la

amenaza que representa el buffel a este vertebrado escaso del Suroeste Norteamericano.

GUTIÉRREZ-GONZÁLEZ, CARMINA1, Miguel Gómez-Ramírez1, and Carlos López-González2. 1Naturalia A. C./ Northern Jaguar Project

Inc. Hermosillo, Sonora, México, reservajaguar@gmail.com, miguelgomezzz@hotmail.com; 2Facultad de Ciencias Naturales, Universidad

Autónoma de Querétaro, Santiago de Querétaro, Querétaro, México, cats4mex@aol.com.

Jaguar permanency and density in the Northern Jaguar Reserve in Sonora, México. years 1999 – 2010. The Northern Jaguar Reserve was

created in order to protect the northernmost population of jaguars in the continent. Camera traps have monitored the reserve for more than 10

years. This long term monitoring effort has allowed us to evaluate jaguar population turnover. We estimated the density of jaguars by year as

well as their survival rate and capture probability. We identified a total of 32 jaguar individuals over 10 years of sampling through each jaguar’s

unique spot pattern. The maximum time spent in the area was six years by a female. The overall residence time was 4 years. The maximum

number of different individuals detected per year was 8. We calculated a home range of 84.6 km2 for the male jaguar with the highest number

of records since 2006. The average density of jaguars on the site was variable but averaged 1.6 (± 1.6) individuals/100 km2/year. According to

the results, the detection probability varies with time and not with the technique used for monitoring. We consider that the use of camera traps

for jaguar studies, besides being a noninvasive technique, provide sufficient information for long-term monitoring. This is the first study that

involves both density and permanency for a time period of 10 years for the northern jaguar population so it cannot be compared with studies in

similar habitats. However, the results of permanence agree with those obtained in long-term studies in the Amazon.

Permanencia y densidad poblacional de jaguares en la Reserva del Jaguar del Norte en Sonora, durante el período de 1999 a 2010. La

Reserva delJaguar del Norte fue creada con la finalidad proteger la población más norteña de jaguares en el continente. Ha sido monitoreada con

trampas cámara por mas de 10 años, incluso desde antes de su establecimiento como reserva. Este monitoreo ha permitido evaluar los cambios

poblacionales de los jaguares en el área de estudio. Para evaluar estas tendencias, se estimó la densidad de jaguares por año, así como la tasa de

supervivencia y la probabilidad de captura. En total se identificó a 32 individuos de jaguar a lo largo de los 10 años de muestreo por medio de su

patrón de manchas único. El tiempo máximo de permanencia en el área de estudio fue de seis años para una hembra pero en general el tiempo

de permanencia resultó de 4 años. El número máximo de individuos diferentes detectados por año fue de 8. Se calculó un ámbito hogareño de

84.6 km2 para el jaguar macho con el mayor número de registros a partir de 2006. La densidad de jaguares varió cada año, con un promedio de

1.6 individuales/100 km2/año. La probabilidad de detección de los jaguares varía con el tiempo y no con la técnica usada para el monitoreo por

lo que se considera que el estudio con trampas cámara además de ser una técnica no invasiva para el estudio de estas poblaciones proporciona

información suficiente para monitoreos a largo plazo de esta especie.
USDA Forest Service Proceedings RMRS-P-67. 2013 537

Abstracts/ Resúmenes

GUTIÉRREZ-GONZÁLEZ,CARMINA1, Miguel Gómez-Ramírez1 andCarlos López-González2. 1NaturaliaA.C. / Northern Jaguar ProjectInc.

Hermosillo, Sonora, México, miguelgomezzz@hotmail.com; 2Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Santiago

de Querétaro, Querétaro, México, cats4mex@aol.com.

Ocelot density in Northern Jaguar Reserve and surrounding areas in Sahuaripa, Sonora, Mexico, using an open population model.

The Northern Jaguar Reserve was established as a private reserve to provide a refuge for the northernmost population of jaguars in the continent. However, the protection afforded by the reserve has been extended to other species under some category of protection, such the ocelot,

who is in danger of extinction according the Mexican legislation. In this study we calculated the density of ocelots in the area bounded by the

Northern Jaguar Reserve and 10 cattle ranches adjacent to it with an agreement to protect wildlife. We sampled monthly with camera traps in

the study area between 2009 and 2011. We obtained 188 ocelot photographs that we could identify to individual level. With all captures, we

calculated the home range of a female in 18.84 km2 and 37.14 km2 for a male. We calculated the density for each month of sampling and the

average density was 4.11 (± 1.42) individuals / 100 km2. The average survival probability (phi) was 0.71 (± 0.16). The use of open population

models for calculating the density of a species can include births and deaths, which gives more reliable results, allowing to study strategies and

conservation of the species in the long term.

Densidad de ocelotes, usando un modelo de población abierta, en la Reserva del Jaguar del Norte y zonas aledañas en Sahuaripa, Sonora, México. La Reserva del Jaguar del Norte fue establecida como reserva privada para proveer un refugio para la población más norteña

de jaguares del continente, sin embargo, la protección brindada por la Reserva se ha extendido a otras especies bajo alguna categoría de

protección, como el ocelote que se encuentra en peligro de extinción en la legislación mexicana. En este estudio calculamos la densidad de ocelotes en el área comprendida por la Reserva del Jaguar del Norte y 10 ranchos ganaderos aledaños a ésta con convenio de protección a la fauna

silvestre. Hicimos muestreos mensuales con trampas cámara en la zona de estudio entre 2009 y 2011. Obtuvimos 188 fotografías de ocelote

identificables a nivel de individuo. Con los datos de capturas fotográficas, calculamos el ámbito hogareño de una hembra en 18.84 km2 y de un

macho en 37.14 km2. Calculamos la densidad para cada mes de muestreo y la densidad promedio fue de 4.11 (±1.42) individuos/ 100 km2. La

probabilidad de supervivencia (phi) promedio fue de 0.71 (± 0.16). El uso de modelos de poblaciones abiertas para el cálculo de densidad de

una especie permite incluir muertes y nacimientos, lo cual da mayor confiabilidad a los resultados, y sirve para generar estrategias de estudio

y conservación de la especie a largo plazo.

GUTÍERREZ-GARCÍA, DANIELA1, Miguel Gómez-Ramírez 1, Carmina Gutiérrez-González 1, and Carlos A. López-González2. 1Naturalia A.

C./ Northern Jaguar Project Inc. Hermosillo, Sonora, México, beluga128@hotmail.com, miguelgomezzz@hotmail.com; 2Facultad de Ciencias

Naturales, Universidad Autónoma de Querétaro, Santiago de Querétaro, Querétaro, México, cats4mex@aol.com.

Mammalian community comparisons related to cattle presence. Livestock husbandry is an activity that can affect mammal distribution and

community structure. To test such effect we considered a site with and without livestock. The Northern Jaguar Reserve is a privately owned with

no cattle beginning in 1999, surrounded by working cattle ranches. Using camera traps we compared mammalian abundance and dominance,

both at the reserve and 10 adjacent ranchesfor three years. We recorded 17 species of medium and large mammals. Only peccary (Pecari tajacu)

showed higher abundance values on cattle ranches than on the reserve. The ringtail (Bassariscus astutus) and the spotted skunk (Spilogale 

gracilis) had similar abundances inside and outside the reserve. The 14 remaining species showed higher abundances in the reserve, including

jaguar and ocelot. The feline abundance also showed differences between sites with abundances in the reserve significantly higher than outside.

These results serve as a basis for understanding the effects of livestock and ecosystem recovery after exclusion, and livestock management

strategies that are compatible with the habitat recovery of mammal species

Comparación de la comunidad de mamíferos en relación con la presencia de ganado. La ganadería es una actividad que puede llegar a

desplazar las poblaciones de mamíferos o alterar la estructura de la comunidad de éstos. Un modelo para estudiar este efecto necesita por lo

menos dos factores, un sitio sin ganado y uno con presencia del mismo. La Reserva del Jaguar del Norte es una propiedad privada cuya exclusión de ganado ha sido gradual y está rodeada de ranchos ganaderos. Mediante trampeo fotográfico se ha monitoreado mensualmente tanto la

reserva como 10 ranchos aledaños a esta por 4 años. Se hicieron comparaciones de la abundancia relativa y la dominancia de especies en ambas

zonas con la finalidad de determinar si la presencia de ganado afecta la estructura de la comunidad. El estudio se llevó a cabo con 17 especies

de mamíferos medianos y grandes. Del total de especies, el jabalí (Pecari tajacu) presentó mayor abundancia en los ranchos ganaderos. El

cacomixtle (Bassariscus astutus) y el zorrillo manchado (Spilogale gracilis) tuvieron abundancias similares dentro y fuera de la reserva. El

resto de las especies presentaron abundancias mayores en la reserva. Las abundancias de felinos dentro de la reserva fueron significativamente

mayores que fuera de ella, estando el jaguar y ocelote poco representados en la zona con ganado. Estos resultados sirven como base para entender los efectos del ganado y la recuperación del ecosistema después de su exclusión y las estrategias de ganado que sean compatibles con la

recuperación de hábitat para las especies de mamíferos.

HALE, SARAH L.1 and John L. Koprowski2. 1School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721,

shale@email.arizona.edu; 2SchoolofNaturalResources andtheEnvironment,UniversityofArizona,Tucson,AZ85721,squirrel@ag.arizona.edu.

A review of black-tailed prairie dog reintroduction strategies and site selection. The black-tailed prairie dog was once widely distributed

throughout the Western United States; however, anthropogenic influences have reduced their numbers to 2% of what they were historically.

Black-tailed prairie dogs are keystone species in the grassland ecosystem, and provide many services that other species cannot, including

providing burrows for other species (e.g. burrowing owls [Athene cunicularia] and rattlesnakes [Crotalus sp.]), excavating nutrient rich soil
538 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

that, in turn, provides rich vegetation for grazers, and serving as food for many carnivores and birds of prey. Several efforts have been made

to reestablish this keystone species to its historical range; however, there has not been a comprehensive review outlining site selection, methodology, and success of these efforts. In southeastern Arizona, a recent reestablishment effort was built upon work of scientists that identified

potential suitable areas with characteristics similar to existing prairie dog colonies in Mexico. Prairie dogs were first translocated to the sites in

2008, and individuals still remain on the landscape today. We compare this to other reestablishment efforts, and provide suggestions on ways

to increase success of future reintroductions.

Análisis de las estrategias de reintroducción y selección de sitios del perrito de la pradera de cola-negra. El perrito de la pradera de colanegra anteriormente tuvo una distribución amplia en el oeste de Estados Unidos; sin embargo actividades antropogénicas lo han reducido a 2%

de la población histórica. En ecosistemas como pastizales los perritos de la pradera son especies clave y aportan innumerables servicios que

otras especies no, incluyendo madrigueras para otras especies (Ej. tecolote llanero [Athene cunicularia] y víbora de cascabel [Crotalus sp.],

excavan suelo rico en nutrientes que suple con vegetación diversa a los herbívoros, y son alimento de muchos carnívoros y aves rapaces. Se han

realizado grandes esfuerzos para restablecer esta especie clave a su rango histórico; sin embargo no se cuenta con una publicación detallada de

la selección de sitios, metodología, y el éxito de estos trabajos. En el sureste de Arizona, una reintroducción reciente se basó en investigaciones

científicas que identificaron áreas adecuadas con características similares a las colonias en México. Los perritos de la pradera se llevaron a éstos

sitos en el 2008, y aún quedan individuos como parte del paisaje. Comparamos estos resultados con otros trabajos, y aportamos sugerencia para

aumentar el éxito de futuras reintroducciones.

HANSON, MOLLY. The Nature Conservancy, 1510 E Ft. Lowell, Tucson AZ 85719, mhanson@tnc.org.

Reduction of erosion and sedimentation along the lower San Pedro River through hydrologic restoration of modified ephemeral washes. 

The Nature Conservancy received funding from the Arizona Water Protection Fund Commission to restore the two ephemeral washes tributary

to the Lower San Pedro River back to their natural state. The purpose of this wash restoration was to rectify considerable erosion problems

across the San Pedro River floodplain. Sheet flow erosion rates at the site increased after a berm that directed wash flows around the agricultural

field was breached during the 2006 flooding event and created chronic erosion and sedimentation conditions across lands cleared and leveled

for agricultural purposes. The project involved decommissioning of a ditch and berm, re-contouring the transition between the uplands and the

terrace, reconstructing two historic washes, and revegetation of native plant communities to stabilize all construction areas. Approximately 9

acres of stream channel were created and ~95 acres surrounding the new washes were hydroseeded with native seed mix. The seed was worked

into the soil with a tractor pulled culti-packer in the uplands and with an OHV pulled flex harrow along the stream channels and banks. The

site will be irrigated, mowed and monitored for at least three years to promote native vegetation establishment, to reduce competition from

non-natives, and to document changes in erosion.

Disminución de erosión y sedimentación en la parte baja del río San Pedro mediante restauración hidrológica de arroyos efímeros 

modificados . La organización Nature Conservancy obtuvo financiamiento de Arizona Water Protection Fund Commission para restaurar dos

arroyos efímeros tributarios del río San Pedro a su estado natural. El propósito de la restauración fue corregir problemas de erosión de la llanura

aluvial del río San Pedro. El índice de erosión laminar delsitio aumentó después que el bordo que permitía el riego del arroyo en la zona agrícola

se lo llevó la corriente del 2006 causando bastante erosión y sedimentación afectando los terrenos agrícolas. El proyecto consistió en destruir

el bordo y la zanja, definir de nuevo la transición entre la altiplanicie y el terraplén, reconstrucción de arroyos históricos, y reforestación de

comunidades de plantas nativas para estabilizar todas las áreas de construcción. Aproximadamente 3.4 ha. del cauce del arroyo fueron creadas

y ca. 40 ha. de los alrededores fueron sembradas con una mezcla de semillas nativas. La semilla se introdujo al suelo con un tractor empleando

un quebra terrones “cultipacker” en las zonas elevadas y un OHV arrastrado a lo largo del canal del arroyo y bancos. El sitio será irrigado,

podado y monitoreado por lo menos tres años para establecer especies nativas, reducir competencia de especies introducidas y documentar los

cambios en erosión.

HARE, TREVOR1, Sarah Williams1, Van Clothier2, Craig Sponholtz3, and Bill Zeedyk4. 1Sky Island Alliance Restoration Program, 300 E

University Blvd, Ste 270, Tucson, AZ 85705, trevor@skyislandalliance.org; 2Stream Dynamics Box 2721, Silver City, NM 88062; 3Drylands

Solutions Inc.551 W. Cordova Rd. #832, Santa Fe, NM 87505; 4Zeedyk Ecological Consulting, LLC.

Peloncillo Mountains ciénega restoration project. Cloverdale Ciénega was a vast desert marsh irrigated by a 10,000-acre watershed that is

now in excellent condition. It was so large, in relation to its watershed size, that it was able to harvest the bulk of the runoff from a typical storm

by super charging its alluvial storage. Down valley, wetland swales brought water to Cloverdale Creek, creating a perennial stream reach. This

wetland complex has been degraded due to an old concrete dam, dozens of berms and a deep gully. The only perennial creek in the Peloncillo

Mountains is now an arroyo and the ciénega is greatly reduced. The project goals were to restore a more natural hydrology to the watershed;

restore riparian vegetation, and to protect and enhance the population of the frog. After careful planning we commenced work in spring 2010.

We removed over 6000 cubic yards of dirt from levees and with 250 tons of boulders we constructed four diversions along the man-made

gully. These structures were designed to allow storm flows to spread out over the entire width of the dried ciénega surface. The re-wetting of

this surface will allow wetland plant communities to re-colonize what is now grassland and will provide enhanced habitat for the Chiricahua

leopard frog (Rana chiricahuensis).
USDA Forest Service Proceedings RMRS-P-67. 2013 539

Abstracts/ Resúmenes

Proyecto de restauración de una ciénega en la Sierra Peloncillo. La Ciénega de Cloverdale fue un extenso humedal del desierto, alimentado

por una cuenca de 5,000 has, que actualmente se encuentra en una condición excelente. Su gran extensión en relación con el tamaño de su cuenca

le permitía aprovechar la mayor parte de los escurrimientos de las lluvias y sobre cargar su almacenaje aluvial. Valle abajo, las depresiones

naturales llevaron agua al Cloverdale Creek, creando un riachuelo perenne. Este complejo de humedales se degradó debido a un antiguo dique

de concreto, docenas de bordos y un gran barranco. El único riachuelo perenne en las Sierra Peloncillo es ahora un arroyo seco y la ciénega

se ha reducido en gran medida. Las metas del proyecto son devolverle una hidrología más natural a la cuenca; restaurar la vegetación de la

ciénega y proteger y aumentar la población de rana. Después de una planeación cuidadosa se empezó a trabajar en la primavera de 2010. Se

removieron más de 5000 metros cúbicos de tierra de los diques y con 250 toneladas de rocas se construyeron cuatro desviaciones para el agua

a lo largo del barranco excavado. Estas estructuras fueron diseñadas para permitir a las corrientes de las lluvias dispersarse a lo largo y ancho

de la superficie seca de la ciénega. Agua en la superficie permitirá que las comunidades de plantas de la ciénega recolonicen lo que ahora es un

pastizal y se mejore el hábitat para la rana chiricahua (Rana chiricahuensis).

HARE, TREVOR.Sky Island Alliance Restoration Program, 300 E University Blvd, Ste 270, Tucson, AZ 85705, trevor@skyislandalliance.org

The future of restoration in the Sky Islands: Bringing volunteers, landowners, practitioners and agencies together to benefit wildlife and 

habitats in a global biodiversity hotspot.The Sky Island region ofthe Southwestern US and Northwestern México isrecognized internationally

as a biodiversity hotspot. Sky Island Alliance is a grassroots organization dedicated to the protection and restoration of this rich natural heritage

of unique and rare species and habitats. We bring volunteers, landowners and agency personnel together to plan and implement riparian and

upland restoration projects to protect desert, grassland and riparian wildlife and habitats. Four case studies of restoration projects are presented

that can inform similar projects. First is a study of a planning project in the headwaters of the Santa Cruz River where a lack of communication

between the project manager, adjacent landowners, and wildlife managers doomed its completion. Second is a study on an ongoing project in

the Huachuca Mountains where man-made impoundments allowed bullfrogs to breed and along with un-natural rates of erosion have extirpated

native aquatic species. Third is a study of non-native species control project in the Pajarito Mountains where in 2009 we initiated a collaborative

landscape-level inventory and control program. Fourth is a study on upland restoration focusing on road density reductions and arroyo erosion

control to protect the oak savannas and riparian areas of the Ciénega Creek Watershed.

El futuro de la restauración en las Islas Serranas: unir voluntarios, propietarios, profesionales y agencias para beneficiar la fauna y el 

hábitat en un área de importancia global por su biodiversidad. La región de las Islas Serranas del Suroeste de E.U. y Noroeste de México

es reconocida internacionalmente como un área de importancia mundial por su biodiversidad. Sky Island Alliance es una organización local

dedicada a la protección y restauración de esta rica herencia natural de hábitats y especies únicas y raras. Reunimos voluntarios, propietarios

y personal de agencias para planear e implementar proyectos de restauración de ríos y tierras adyacentes para proteger desiertos, pastizales,

hábitats y fauna ribereña. Se presentan cuatro casos de estudio de proyectos de restauración que pueden ilustrar proyectos semejantes. El

primero es el estudio de un proyecto de planeación en el nacimiento del río Santa Cruz donde la falta de comunicación entre el encargado del

proyecto, los propietarios vecinos y los encargados de fauna causó que éste fracasara. El segundo es un estudio de un proyecto en curso en la

Sierra Huachuca donde la construcción de represos permitió la reproducción de la rana toro que junto con índices de erosión anormales han

extirpado las especies acuáticas nativas. El tercero es un estudio de un proyecto de control de especies introducidas en la Sierra Pajarito donde

en 2009 empezamos un inventario biológico y un programa de control. El cuarto es un estudio de restauración de tierras altas concentrado en

reducir el número de caminos y la erosión de arroyos para proteger los pastizales, encinales y áreas ribereñas de la cuenca de Ciénega Creek.

HARE, TREVOR1 and Van Clothier2 . 1Sky Island Alliance Restoration Program, 300 E University Blvd, Ste 270, Tucson, AZ 85705, trevor@

skyislandalliance.org; 2 Stream Dynamics Box 2721, Silver City, NM 88062.

Walking the ciénegas of the Sky Island Region: Current condition and future potential. We describe current conditions and discussrestoration and long-term protection potential of a selected ciénegas of the region based on preliminary assessments. There is a scarcity of information,

current or historical, on the ciénegas of the region and we hope to add to the renewed interest in and growing knowledge of their ecology, and their

protection and restoration. Sky Island Alliance staff, volunteers, and collaborators have visited many historic ciénega sites and have performed

assessments that ranged from intensive planning to short (1-2 days) investigations. These assessments have focused on two issues, what are the

impacts affecting the system and what is the restoration potential? We have assessed, to one degree or another, a range of ciénega types from

large (>200 acres) to small (<1 acre) associated with rivers, creeks, and spring and seep systems. Impacts include aquifer drawdown, draining,

un-managed livestock grazing, invasive species of plants and animals, flow capture by roads, water developments, erosion such as arroyo cutting, and lack of management. Ciénegas discussed include upper and lower Cloverdale, Saracachi, Los Fresnos, St. David, San Simon, Animas

Creek, Canelo Hills, Bog Hole, Ciénega Creek, plus othersitesin the Burro, Peloncillo, Huachuca, Pinaleño, Galiuro, and Santa Rita Mountains.

Caminando por las ciénegas de la Región de la Islas Serranas: condiciones actuales y potencial. Según una evaluación preliminar se seleccionó un grupo de ciénegas para describir las condiciones actuales, discutir su restauración y el potencial de protección a largo plazo. Debido

a que existe muy poca información actual o histórica de las ciénegas de la región, esperamos aumentar el renovado interés en las mismas y el

conocimiento de su ecología, protección y restauración. Personal de Sky Island Alliance, voluntarios y colaboradores visitaron varias ciénegas

históricas e hicieron evaluaciones que varían desde trabajosintensivos a investigaciones cortas de 1 o 2 días. Estas evaluacionesse concentraron

en dosinquietudes: ¿Cuálesson losimpactos afectando elsistema? y ¿Cuál es el potencial de restauración? Se han evaluado de una u otra forma,

una gran variedad de ciénegas desde grandes (>80 has) a pequeñas (<0.5 has) asociadas con ríos, riachuelos, aguajes y sistemas de filtración.

Los impactos incluyen descenso del acuífero, drenado, sobrepastoreo del ganado, especies invasoras de plantas y animales, corrientes de agua
540 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

en los caminos, infraestructura para uso del agua, erosión de las paredes de los arroyos y falta de manejo. Se discuten las ciénegas de Cloverdale

alto y bajo, Saracachi, Los Fresnos, St. David, San Simón, Ánimas Creek, Canelo Hills, Bog Hole, Ciénega Creek, además de otros sitios en

las Sierras Burro, Peloncillo, Huachuca, Pinaleño, Galiuro y Santa Rita.

HARTMANN, HOLLY C.1, Kiyomi Morino2, and Kimery Wiltshire3. 1Arid LandsInformation Center, University ofArizona, Tucson,AZ, holly.

hartmann@arizona.edu; 2Laboratory of Tree-Ring Research, University of Arizona, Tucson, AZ, kmorino@ltrr.arizona.edu; 3Carpe Diem West,

325 Pine Street, Sausalito, CA 94965, kimery@carpediemwest.org.

An organized toolbox: Choosing the appropriate tools and methods to support climate change adaptation. Resource managers face a

confusing and often overwhelming plethora of evolving tools and methods for considering climate change in planning and management. Many

require substantial investments in data gathering, analysis, or stakeholder engagement. Many address only pieces of the climate change adaptation challenge without clear interconnection. Irreducible uncertainties associated with climate change call for an iterative approach to risk

management using a mix of tools and methods matched to the stage of risk management being addressed, the specific issues and time scales

of relevance, the resources available for planning, and desired characteristics for stakeholder engagement. We integrated several organizing

frameworks into an interactive ‘toolbox’, the Carpe Diem West Academy (carpediemwestacademy.org). It reviews and characterizes over 100

extant tools, training resources, and best practices useful for planners and managers. While developed specifically for water resources planning,

it has broad applicability for management of natural areas, because water-related stresses pervasively affect achievement of natural and cultural

resource objectives, and because many tools, themselves, have broad applicability. Shared learning is supported through a series of engaging

webinars addressing specific questions of planners and managers.The toolbox also is useful for identifying gaps where additional tools, methods,

or professional development training are needed.

Protocolo organizado: Cómo escoger las técnicas y métodos adecuados para la adaptación al cambio climático. El personal de manejo

de recursos se enfrenta con complicadas y abrumadoras técnicas y metodologías siempre cambiantes al tratar de incluir el cambio climático

en sus planes de manejo. Muchos requieren inversiones considerables para obtener información, analizarla, o la participación de los sectores

interesados. Muchos sólo abarcan fracciones de la adaptación al cambio climático sin una clara interconexión. La incertidumbre inapelable

asociada con el cambio climático requiere de una aproximación reiterativa en el manejo de riesgos empleando una mezcla de técnicas y métodos a la altura del problema que se está abordando, temas específicos y escala de tiempo relevante, los recursos disponibles para la planeación,

y características ideales de los distintos sectores. Integramos varios marcos de referencia en un “protocolo” interactivo: la Carpe Diem West

Academy (carpediemwestacademy.org). Aquí se compilan y caracterizan más de 100 técnicas, cursos de capacitación, y las mejores prácticas

útiles para los encargados del programa de manejo. Aunque se diseñó para el manejo de recursos hídricos, se puede aplicar al manejo de áreas

naturales, porque el estrés relacionado con el agua afecta en gran medida los objetivos del resto de los recursos naturales y culturales, además

muchas técnicas se pueden aplicar independientemente a distintos problemas. Se promueve el aprendizaje compartido a través de seminarios

por internet abordando preguntas específicas de los encargados de manejo. Este “protocolo” también es útil para identificar lagunas donde se

necesiten nuevas técnicas, metodologías o capacitación profesional.

HEILMAN, PHILIP1, Timothy O. Keefer,2 and David C. Goodrich3. 1USDA-ARS Southwest Watershed Research Center, 2000 E. Allen Rd.

Tucson AZ 85719, phil.heilman@ars.usda.gov; 2USDA-ARS Southwest Watershed Research Center, 2000 E. Allen Rd. Tucson AZ 85719, tim.

keefer@ars.usda.gov; 2USDA-ARS Southwest Watershed Research Center, 2000 E. Allen Rd. Tucson AZ 85719, dave.goodrich@ars.usda.gov.

The utility of PRISM precipitation estimates within the Madrean Archipelago. Precipitation is the primary driver for ecological processes

across the Madrean Archipelago. Often researchers and managers must interpret monitoring observations, or assess potential climate change,

across large areas based on a limited number of rain gauges. A potential additional source of information is the PRISM (Parameter-elevation

Relationships on Independent Slopes Model) dataset (http://www.prism.oregonstate.edu/) available in the U.S. at both 4 km (free) and 800 m

(priced) spatial and monthly temporal resolutions. Because so much of the precipitation in the Madrean Archipelago falls as short-duration,

high-intensity convective thunderstorms during the North American Monsoon, it is unclear how to interpret the PRISM estimates, which do

not reflect the spatial variability inherent in those storms. We compared measured values for a single gauge located in Tombstone and PRISM

estimates of annual precipitation to the measured precipitation from 88 rain gauges for the 149 sq. km Walnut Gulch Experimental Watershed,

1999-2008. The mean error of area-weighted annual precipitation from a single gauge was -10%, with the greatest single year error of -37%.

PRISM had corresponding errors of 9 and 30%. This preliminary analysis indicated that PRISM could complement limited observations to

improve spatial and temporal understanding of precipitation for some research and management purposes.

Estimación de precipitación en el Archipiélago Madrense usando PRISM. El factor principal en procesos ecológicos del Archipiélago

Madrean es la precipitación. Investigadores y administradores deben interpretar las observaciones del monitoreo, o evaluar el cambio climático

potencial de grandes áreas basándose en un número limitado de pluviómetros. Una fuente adicional es la información de PRISM (por sus siglas

en inglés, modelo de relación de parámetro-elevación en pendientes independientes http://www.prism.oregonstate.edu/) disponible en Estados

Unidos en dos presentaciones 4 km(gratis) y 800 m (costo) de resolución espacial y con una resolución temporal mensual. Debido a que mucha

de la precipitación en el Archipiélago Madrense cae en periodos cortos durante tormentas convectivas de gran intensidad durante el monzón

norteamericano, la información del PRISM es difícil de interpretar, ya que no refleja la variabilidad espacial de las tormentas. Comparamos

valores de un pluviómetro en Tombstone, además de estimaciones de precipitación anual del PRISM y las medidas de precipitación de 88 pluviómetros de los 149 km2 de la estación experimental Walnut Gulch, de 1999 a 2008. El error medio de la precipitación anual ponderada de un
USDA Forest Service Proceedings RMRS-P-67. 2013 541

Abstracts/ Resúmenes

solo pluviómetro fue de -10%, mientras que el error mayor de un solo año fue de -37%. Los errores de PRISM fueron 9 y 30% respectivamente

Este análisis preliminar sugiere que PRISM podría complementar las observaciones limitadas para mejorar la compresión de precipitación

espacial y temporal, con fines de investigación y manejo.

HEMSTROM, MILES1, Jessica Halofsky2, Joshua Halofsky3, David Conklin4, Dominique Bachelet, and Becky Kerns5. 1Portland Forestry

Sciences Laboratory, 620 SW Main, Portland, Oregon 97205, mhemstrom@fs.fed.us; 2Portland Forestry Sciences Laboratory, 620 SW Main,

Portland, Oregon 97205; 3Portland Forestry Sciences Laboratory, 620 SW Main, Portland, Oregon 97205; 4Conservation Biology Institute, 136

SW Washington Avenue, Suite 202, Corvallis, Oregon 97333; 5 Corvallis Forestry Sciences Laboratory, 3200 SW Jefferson Way, Corvallis,

Oregon 97331.

Potential climate change effects on vegetation types in the Sky Islands area. Climate change may have strong effects on the vegetation and

associated habitats and human values in the Sky Islands area of southeastern Arizona. We connected a dynamic global vegetation model (MC1)

to local state and transitions models (STMs) for major Sky Island vegetation types to better understand how changes in climatically driven

vegetation potential and wildfire regimes might affect the local vegetation in the Sky Islands area. MC1 produces projections of potential future

vegetation types and wildfire regimes under alternative climate scenarios, but does not generally consider detailed vegetation dynamics such

as succession, growth, fire resistance, competition, and other factors. On the other hand, while available local STMs track the effects of disturbances, vegetation dynamics, and management on individual vegetation cover and structure combinations, STMs assume vegetation potential

is static. We derived changes in vegetation potential and wildfire from MC1 under three different climate scenarios and incorporated them in a

suite of local STMs. Our results suggest that climate-driven changes over the next 100 years could be dramatic, including substantial declines

in coniferous forest area and large increases in warm-season (C4) grasslands. Our models also allow land managers and others to examine the

possible effects of different vegetation management activities on future vegetation conditions given the three climate change scenarios.

Efectos potenciales del cambio climático en la vegetación de las Islas Serranas. El cambio climático puede afectar en granmedida la vegetación

y hábitats asociados e inmuebles de las Islas Serranas del sureste de Arizona. Conectamos un modelo de vegetación mundial dinámico (MC1)

con modelos locales y en transición (STMs) para los principales tipos de vegetación de las Islas Serranas, para entender como los cambios en

vegetación causados por el clima, y cambio en el régimen de incendios forestales pueden afectar la vegetación local de las Islas Serranas. Los

MC1 proyectan tipos de vegetación potencial en el futuro y el régimen de incendios bajo diferentes escenarios climáticos, pero no consideran

procesos dinámicos detallados de la vegetación como sucesión, crecimiento, resistencia a fuego, competencia y otros factores. Por otro lado,

aunque los modelos locales STMs siguen los efectos de disturbio, dinámica de la vegetación, manejo de cobertura vegetal por especie y combinaciones de estructura, los STMs asumen que el potencial vegetal es estático. Derivamos cambios en la vegetación potencial y los incendios de

MC1 bajo tres diferentes escenarios climáticos y los incorporamos a los STMs locales. Los resultados sugieren que los cambios causados por

el clima en los próximos 100 años pueden ser drásticos, con considerable disminución del área de bosques de coníferas y un gran incremento

en pastizales (C4) de estación cálida. Nuestros modelos permiten que el personal de manejo de recursos y otros, examinen los posibles efectos

de actividades de manejo bajo condiciones futuras de vegetación en tres escenarios del cambio climático.

HERNÁNDEZ-RODRÍGUEZ, ANA LILIA, María de la Paz Montañez-Armenta, Hugo Silva-Kurumiya, and Gertrudis Yanes-Arvayo. Universidad de la Sierra, División de Ciencias Biológicas, Moctezuma, Sonora, México.

Floristic analysis of a vegetation island and biological corridor in Sonora, Mexico. We performed a floristic study of the vegetation islands

on the mountain town of Montezuma, Sonora. From August to December 2011, the relevé method was used on ranches El Rodeo and Basora.

The sites are located ca. 900 m elevation in foothills thornscrub. We collected a total of 85 species in 70 genera and 30 families. The species

were identified through expert advice, literature, and comparision with specimens in the Herbarium of the Universidad de Sonora Herbarium

(USON) in Hermosillo. Specimens were deposited into USON. The best represented families are Asteraceae, Convolvulaceae, Fabaceae, and

Solanaceae. The study area is a transition zone between subtropical thornscrub and Sierra Madrean forest elements, making it an important

biological corridor for the Sky Islands. It was found that the loss of flora is high due to changing land use, mainly the invasion of buffelgrass

(Pennisetum ciliare). Representative species of the area as Zinnia zinnioides, Ipomoea cristulata, I. hederacea, and Abutilon can be replaced

by species that have adapted to local new conditions.

Análisis florístico de las islas de vegetación y corredor biológico de la sierra media de Sonora, México. Se realizó un estudio florístico en

las islas de vegetación de la sierra media del municipio de Moctezuma, Sonora. De agosto a diciembre de 2011, se realizaron tres recorridos

de campo usando el método de relevé en los ranchos El Rodeo y Básora. Los sitios son ubicados ca. 900 msnm en matorral espinoso. Se recolectaron un total de 85 especies en 70 géneros y 30 familias. Las especies fueron identificadas mediante la asesoría de expertos, bibliografía

y herbario USON. Ejemplares fueron depositados en el herbario de la Universidad de Sonora. Las familias mejor representadas son Asteraceae,

Convolvulaceae, Fabaceae y Solanaceae. El área de estudio es una zona de transición del matorralsubtropical y elementos del bosque madrense,

que lo convierten en un importante corredor biológico para las Islas Serranas. Se encontró que la pérdida de flora es alta debido al cambio de

uso de suelo, principalmente por la invasión de zacate buffel (Pennisetum ciliare). Especies representativas de la zona como Zinnia zinnioides, 

Ipomoea cristulata, I. hederacea e Abutilon pueden ser desplazadas por especies que se han ido adaptando a las condiciones nuevas del lugar.
542 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

HINOJO-HINOJO, CÉSAR1, Carolina Trujillo- López1, Luis Omar Calva-Pérez1, Onésimo Galaz-García1, and Alejandro E. CastellanosVillegas1

. 1Departamento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Blvd. Luis Encinas y Rosales S/N, Col. Centro,

Hermosillo, Sonora, México, carolatrujillo@hotmail.com.

Association between nurse plants and saguaros (Carnegiea gigantea) in the western Sonora. The objectives are to determine nurse species

associated with the saguaro, as well as the plants in the patch, and the mortality of nurse plants. Because the presence of termites on the green

tissue of the saguaro has been linked, the existence of protection against termites in three sites, located in the western Sonora, was studied. In

each site a transect using the quadrant point method was performed, where data was taken on the closest saguaros, organic material coverage

around it, termite activity in them, the conditions of its nurse plants, the floristic composition, and the dimensions of the associated patch of

vegetation. Data on 56 saguaros showed the most common nurse plants were Larrea tridentata, Olneya tesota, Ambrosia deltoidea, and Parkinsonia microphylla. Twenty-one (21) species were found distributed in different patches associated with the saguaro, where O. tesota and P. 

microphylla showed the mostspecies diversity. The mortality of the nurse plantsis associated with the number of plantslocated under its canopy

and its size. The termite activity was in saguaros with little organic material. The positive interactions permit for the saguaro populations in this

region to be healthy. It is the first work to document the decrease of termite invasion of live saguaro tissue.

Asociación de las plantas nodrizas con el Sahuaro (Carnegiea gigantea) en la región oeste de Sonora. Los objetivos son determinar especies nodrizas asociadas al sahuaro, así como la diversidad de plantas en el parche y la mortalidad de plantas nodrizas. Dado que se ha asociado

la presencia de termitas sobre tejido verde de sahuaro, se estudió la existencia de protección contra termitas en tres sitios ubicados en la región

oeste de Sonora. En cada sitio se realizó un transecto utilizando el método del punto cuadrante, donde se tomaron datos de los sahuaros más

cercanos, cobertura de materia orgánica alrededor, actividad de termitas en sahuaros, condición de su nodriza, composición florística y dimensiones del parche de vegetación asociado. Se muestrearon 56 sahuaros, las nodrizas más comunes fueron Larrea tridentata, Olneya tesota, 

Ambrosia deltoidea y Parkinsonia microphylla. Se encontraron 21 especies distribuidas en diferentes parches asociados al sahuaro, donde O. 

tesota y P. microphylla presentan mayor diversidad de especies. La mortalidad de la nodriza está asociada al número de plantas de sahuaro establecidas bajo su dosel y tamaño. La actividad de termitas estaba en sahuaros con poca materia orgánica. Las interacciones positivas permiten

que poblaciones del sahuaro en esta región estén saludables. Es el primer trabajo que documenta la disminución de la invasión de termitas a

los tejidos vivos de sahuaro.

HINOJO-HINOJO, CÉSAR1, Jose M. Llano-Sotelo1, and Alejandro E. Castellanos-Villegas1 1Departamento de Investigaciones Científicas y

Tecnológicas, Universidad de Sonora, Blvd. Luis Encinas y Rosales S/N, Col. Centro, Hermosillo, Sonora, México

Wide range of functional attributes in the Central Sonora flora: a diversity yet to be explored. Even though the Sonoran desert is not

considered the desert with more species richness, it has been documented in multiple occasions that it has the most diversity of growth forms

of North America. It is unknown if in addition to high growth form diversity there could also be high functional diversity in this Desert. We

characterized the ecophysiological functional attributes related to photosynthetic capacity, stomata conductance, transpiration, and resources

(water and nitrogen) use efficiency, as well as leaf area morphostructural attributes and nitrogen content in leaves of 51 species of the arid and

semiarid region of central Sonora. We found a wide range in the studied attributes even higher in comparison with species studied worldwide,

which is confirmation of high functional diversity. This study presents evidence of high functional diversity in the Sonoran Desert that is waiting to be described and documented.

Gran amplitud en el rango de atributos funcionales encontrada en la flora de la Región Central de Sonora: una diversidad por ser 

explorada . En el Desierto Sonorense, aunque no sea este el desierto con mayor riqueza de especies de plantas, se ha documentado en repetidas

ocasiones que éste posee la mayor diversidad en cuanto a formas de crecimiento en Norteamérica. Se desconoce si además de esta alta diversidad,

este desierto pudiera también albergar una alta diversidad funcional. En este trabajo se caracterizaron los atributos funcionales ecofisiológicos

de capacidad fotosintética, conductancia estomática, transpiración y eficiencia de uso de recursos (agua y nitrógeno), así como los atributos

morfoestructurales de área foliar específica y contenido foliar de nitrógeno en 51 especies que habitan en las zonas áridas y semiáridas de la

Región Central de Sonora. En tan solo estas especies se encontró en los rangos de todos los atributos medidos una amplitud similar e incluso

mayor a la reportada para todas las especies del mundo en las que han sido estudiados dichos atributos, lo cual refleja una alta diversidad funcional. Esta gran amplitud no es algo comúnmente encontrada en otras regiones del mundo. Este trabajo presenta evidencia de la alta diversidad

funcional del Desierto Sonorense en espera de ser descrita y documentada.

HODGES, DAVID1 and Jeff Williamson2. 1Cuenca Los Ojos Foundation, 12626 E. Turkey Creek Road, Pearce AZ 85625, dhodges73@gmail.

com; 2Arizona Zoological Society, 455 N Galvin Parkway, Phoenix, AZ 85008-3431, jwilliamson@thephxzoo.com.

Planning the future on the Coronado National Forest. Most U.S. Sky Island mountain ranges are public lands managed by the U.S. Forest

Service.The majority (20) are found on 12 Ecosystem Management Units oftheCoronado National Forest.TheCoronado is currently engaged in

two multi-year processes - Land and Resource Management Planning, and Travel Management Planning - as are all U.S. national forests. These

public processes will determine future management of, and impacts to, these Sky Island ranges and their hydrology, wildlife, and opportunities

for quite recreation, as well as cultural, geologic, scenic, historic, and vegetation resources. Major challenges to good management planning

outcomesinclude shrinking Forest Service budgets, laws and regulationsthat do not keep pace with science or community values, climate change,
USDA Forest Service Proceedings RMRS-P-67. 2013 543

Abstracts/ Resúmenes

and a 20th century paradigm that pushes Multiple Use into as many areas as possible. Some recent initiativessuch as FireScape and the Pinaleno

Ecosystem Restoration Project provide a useful model for agency response to public input and the use of science to solve complex challenges.

Planificar para el futuro del Bosque Nacional Coronado. La mayoría de las Islas Serranas de Estados Unidos son áreas naturales manejadas

por el Servicio Forestal de Estados Unidos. La mayoría (20) se encuentran en 12 Unidades de Manejo de Ecosistemas del Bosque Nacional

Coronado. El Coronad, igual que todos los bosques nacionales en EU, está trabajando en dos proyectos plurianuales: Plan de Manejo de Tierra

y Recursos y Plan de Manejo de Visitantes. Estos procesos con la ciudadanía determinarán el manejo futuro e impacto en estas Islas Serranas

su hidrología, y flora y fauna, así como oportunidades de recreación, culturales, geológicas, escénicas, históricas, y de uso maderable. Los

grandes retos para obtener buenos resultados con el plan de manejo son: reducción del presupuesto del Servicio Forestal, leyes y normativas no

actualizadas con la ciencia o intereses de la comunidad, cambio climático y el paradigma del siglo XX que pide usos múltiples para la mayoría

de áreas. Algunos programas nuevos como FireScape y el proyecto de restauración del ecosistema Pinaleño son modelos de cómo la atención

de las agencias a la aportación del público y el uso de la ciencia pueden resolver problemas complejos.

HODGES, DAVID. Cuenca Los Ojos Foundation, 12626 E. Turkey Creek Road, Pearce AZ 85625, dhodges73@gmail.com.

Si se puede! A decade of restoration, research, and science on cross-border private lands. The Cuenca Los Ojos Foundation owns and/or

manages more than two hundred thousand acres in southeast Arizona and northeast Sonora. Much of the foundation’s focus has been on restoring landscapes and wildlife populations that have been damaged to varying degrees by over utilization of natural resources and the cascading

effects of this over utilization. Initial work on these lands was focused on both passive and active restoration. Over the past decade, Cuenca Los

Ojos lands have attracted almost a hundred individual researchers from dozens of universities, NGO’s, as well as private investigators. These

lands have become outdoor laboratories for the study of natural processes, specific species, and ecosystem services. In an attempt to better

understand these processes and services, we recently conducted follow-up surveys and interviews with scientists who had worked here. Survey

results support the status of these lands as biodiversity hotspots and providers of valuable ecosystem services. Results that follow restoration

activities document the return of several locally extirpated species, resurgence of grass cover, and re-wetting of streams and ciénegas.

¡Si se puede! Una década de restauración e investigación científica en tierras de propiedad privada a través de la frontera. Más de 80,

937 has pertenecen o son manejadas por la fundación Cuenca Los Ojos en el sureste de Arizona y noreste de Sonora. La mayor parte del trabajo

de la fundación se dedica a restaurar el paisaje y poblaciones de fauna y flora que han sido degradadas en varios niveles por la sobreexplotación

de los recursos naturales y efectos cascada de la misma. Los trabajos iniciales incluyeron tanto métodos pasivos como activos de restauración.

Casi cien investigadores de docenas de universidades, ONG’s y del sector privado han visitado Cuenca Los Ojos, durante la última década,

Estas tierras se han convertido en un laboratorio para el estudio de procesos naturales, especies específicas y servicios ecosistémicos. Con el

intento de entender mejor estos procesos y servicios, entrevistamos a los investigadores que han realizado sus proyectos aquí. El resultado de

estas encuestas confirma el estatus de estas tierras como centro de biodiversidad y fuente de invaluables servicios ecosistémicos. Resultados

del trabajo de restauración incluyen el regreso de varias especies extirpadas localmente, resurgimiento de la cobertura de zacates, y agua en

arroyos y ciénegas.

HODGSON, WENDY and Andrew Salywon. Desert Botanical Garden, 1201 N. Galvin Parkway, Phoenix, AZ 85008, whodgson@dbg.org,

asalywon@dbg.org.

Pre-Columbian Agaves in Arizona: a new way of looking at species and their landscapes. The importance of agaves to Mesoamerica and

its cultures has long been recognized, providing food, fiber and beverage. However, their significance to these cultures has overshadowed and

distorted the plants’role for indigenous peoples north of the U.S. - Mexico border. Pre-Columbian farmers cultivated several species of agave in

Arizona dating to at least A.D. 600, including Agave murpheyi, A. delamateri, A. phillipsiana, A. parryi var. parryi, A. parryi var. huachucensis,

and two undescribed species. Because of their longevity and primarily asexual reproduction, relict agave clones have persisted in the landscape

to the present, providing an opportunity to study pre-Columbian nutrition, trade, migration and agricultural practices.Additionally, these remnant

clones present a rare opportunity to examine cultivars virtually unchanged since they were last cultivated within a prehistoric cultural context.

At least three, A. murpheyi, A. delamateri, and A. phillipsiana, are most likely not native to Arizona, probably having originated in northern

Mexico and traded asfar north as Grand Canyon. These discoveries underscore the necessity of viewing landscapes and some plantspeciesfrom

a cultural, rather than “natural,” perspective that may help discern potential cryptic species veiled by more traditional taxonomic treatments.

Understanding these plants and their ecological/cultural roles requires interdisciplinary collaboration between botanists and archaeologists.

Agaves precolombinos en Arizona: un nuevo enfoque en especies y su paisaje. La importancia del agave en la cultura mesoamericana es

ampliamente reconocida por ser fuente de alimento, fibra y bebida. Sin embargo, esa importancia cultural a opacado y distorsionado el papel

de estas plantas para los indígenas al norte de la frontera entre México y E.U.. Labradores precolombinos cultivaban varias especies de agave

en Arizona por lo menos desde el año 600 DC., como son: Agave murpheyi, A. delamateri, A. phillipsiana, A. parryi var. parryi, A. parryi var.

huachucensis y dos especies no descritas. Debido a su longevidad y reproducción asexual, relictos de clones de estos agaves han persistido en el

paisaje, permitiendo estudiar nutrición,intercambio,migración yprácticas agrícolas de culturasprecolombinas.Aunadoa ésto los clones presentan

la oportunidad de examinar cultivares virtualmente prístinos ya que fueron cultivados por última vez en el contexto prehistórico. Al menos A. 

murpheyi, A. delamateri y A. phillipsiana, no son nativos de Arizona, tal vez son del norte de México y se intercambiaron entre culturas hasta

el Gran Cañón. Estos descubrimientos enfatizan la necesidad de estudiar el paisaje y algunas especies de plantas con una perspectiva cultural
544 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

en lugar de “natural” y así identificar especies crípticas encubiertas por tratamientos taxonómicos tradicionales. El estudio de éstas plantas así

como su papel ecológico y cultural requiere de colaboración interdisciplinaria entre botánicos y arqueólogos.

HOWE, MARK L. USDA Sequoia National Forest, Kern River Ranger District, Lake Isabella, CA. 93240, Mark.Howe@ibwc.gov.

The 2011 Wallow Fire: An archeologists perspective on BAER work. The 2011 Wallow Fire in Arizona and New Mexico burned over

553,456 acres on the Apache-Sitgreaves and Gila National Forests with associated private and Tribal lands. Archeological work was to first

protect cultural resources then move into BAER (Burned Area Emergency Rehabilitation) work for saving cultural resources after the fire. Our

work on BAER was to first find sites affected by the Wallow Fire then determine damage and assess mitigation for protecting them. This was

to protect from damage by debris flows or other natural factors with the upcoming monsoon season in Northeastern Arizona. Since all fires are

different, the Wallow Fire allowed us to work on what we can do to protect sites quickly with little resources on a fire of unprecedented size

using skills learned in BAER work from previous large fires across the country.

Incendio Wallow 2011: una perspectiva arqueológica de BAER. En el 2011 el incendio forestal Wallow en Arizona y Nuevo Mexico afectó

más de 224,000 ha. en los Bosques Nacionales Apache-Sitgreaves y Gila, así como terrenos privados y pertenecientes a tribus. El trabajo arqueológico primero se enfocó en proteger los recursos culturales y segundo realizar el trabajo de BAER (por sus siglas en inglés, rehabilitación

de emergencia de áreas quemadas) de proteger recursos culturales después del incendio. El trabajo en BAER consistió en encontrar zonas

afectadas por el incendio Wallow, determinar el daño y evaluar la mitigación necesaria para protegerlas. La protección fue prevenir derrumbes

u otros factores naturales que pudieran causar las lluvias del monzón en el noreste de Arizona. Ya que todos los incendios son diferentes, el

incendio Wallow permitió que se trabajara rápido en la protección de sitios con recursos escasos empleando técnicas adquiridas en incendios

anteriores en BAER.

INIGUEZ, JOSE M.1, Stephen R. Yool2, and Geoffrey G. Towne3. 1Rocky Mountain Research Station, 2500 Pine Knoll Dr., Flagstaff, AZ

86001, jiniguez@fs.fed.us; 2Department of Geography and Regional Development, University of Arizona, Tucson, AZ 85721, yools@email.

arizona.edu; 3Department of Geography and Regional Development, University of Arizona, Tucson, AZ 85721, gtowne@email.arizona.edu.

A pre and post-fire comparison of vegetation in the Santa Catalina Mountains. In 1995 vegetation plots were sampled along a series of

transects within five of the largest mountain ranges in the sky islands. Since then many of these areas have burned. To take advantage of this

unique opportunity, in 2009 the vegetation plots in the Santa Catalina Mountains were re-measured. These plots burned in 2002 and 2003 as

part of the Bullock and Aspen fires, respectively. The objectives of this study were to (1) asses how stand structure affected burn severity: (2)

determine the impact of burn severity on stand structure and (3) asses post-fire succession patterns. Stand structure is critical in forest and fire

ecology. Stand structure influences fire behavior particularly crown fires and is also directly related to succession and therefore a major part of

state-transition models. Despite the importance of stand structure, little is known about the relationship between fire and stand structure in the

sky islands. Our results suggest that some previously forested areas have been converted to shrub fields as a result of high burn severity; however, the majority of the plots sampled in 1995 experienced low to moderate severity fire. This information will provide valuable information

that managers can use in managing post-fire landscape mosaics.

Una comparación de vegetación antes y después de incendios en la Sierra Santa Catalina. Se hicieron muestreos en cuadrantes de vegetación en una serie de transectos ubicados en 5 de las cordilleras más grandes de las Islas Serranas en 1995. Desde entonces muchas de estas

áreas se han quemado. Para aprovechar esta oportunidad única, en 2009 los cuadrantes de vegetación en la Sierra Santa Catalina se midieron

de nuevo. Estas parcelas se quemaron en 2002 y 2003 en los incendios Bullock y Aspen, respectivamente. Los objetivos de esta investigación

fueron (1) evaluar como la estructura forestal afectó la severidad del incendio: (2) determinar el impacto de la severidad del incendio en la

estructura forestal y (3) evaluar patrones de sucesión después de incendios. La estructura forestal es crítica en la ecología forestal y del fuego.

La estructura forestal influye el comportamiento del fuego, especialmente fuegos de copas, y también se relaciona directamente con la sucesión

y por tanto es una parte importante de los modelos de transición. A pesar de la importancia de la estructura forestal, se conoce poco sobre la

relación entre el fuego y la estructura forestal en las islas serranas. Nuestros resultados sugieren que algunos rodales forestales se han convertido

en matorrales por causa de incendio muy calientes; sin embargo, la mayoría de las parcelas muestreadas en 1995 experimentaron incendios de

intensidad baja o moderada. Esta información será muy útil en el manejo de paisajes afectados por el fuego.

JIMÉNEZ-MALDONADO, ROSA ELENA1, Christa Weise2, Mario Cirett-Galán1, Guadalupe Flores1, Manuel Munguia1, E. Isaías Ochoa3. 1

CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos-Bavispe, Segunda este final y Av. Sinaloa S/N, C.P. 86120. Cananea,

Sonora, elena.jimenez@conanp.gob.mx; 2KOFA National Wildlife Refuge. U.S. Fish and Wildlife Service, Tucson AZ, Christa_weise@fws.

gov; 3NATURALIA, A.C., El Cajón #9, Col. Santa Fe, C.P. 83249 Hermosillo, Sonora.

Preliminary List of flying mammals in the Ajos-Bavispe National Forest Reserve and Wildlife Refuge, Sonora: synthesis. The generation

of information on bat communities, including their composition, abundance, distribution and ecology, can support management programs in

protected areas, and also provide information and initiativesfor the designation of new protected areas. From the year 2010 and 2011 monitoring

was done within a Sky Island, called Sierra de los Ajos, which is part of the Ajos-Bavispe reserve. During this period of time, we had recorded a

total of eightspecies(Myotis californicus, Myotis auriculus, Myotis volans, Myotis thysanodes, Lasiurus cinereus, Lasiurus blossevillii, Eptesicus 

fuscus, and Choeronycteris mexicana) in riparian and oak-pine forests. This preliminary list can speculate about the good health of the ecosystem
USDA Forest Service Proceedings RMRS-P-67. 2013 545

Abstracts/ Resúmenes

in this protected area and provides good guidelines for carrying out monitoring in other sky islands within the same protected area: La Madera,

El Tigre, Buenos Aires, and La Púrica. The intention of this paper is to use the bat as a tool to assess the degree of conservation of these Sky

Islands, because they are abundant, diverse, relatively easy to monitor and there are low-cost materials used for study. In the end, presence and/

or absence of bat communities will allow conservation and management decisions in the short and long term in this important protected area.

Listado preliminar de mamíferos voladores en la Reserva Forestal Nacional y Refugio Fauna Silvestre Ajos-Bavispe: síntesis. La generación de información de las comunidades de murciélagos,su composición, abundancia, distribución y ecología, permite sustentar los programas

de manejo en áreas naturales protegidas, y proporcionan datos e iniciativas para la designación de nuevas áreas de protección. De 2010 a 2011

se realizó un monitoreo en la Sierra de Ajos que forma parte de Ajos Bavispe, registrándose un total de ocho especies (Myotis californicus, 

Myotis auriculus, Myotis volans, Myotis thysanodes, Lasiurus cinereus, Lasiurus blossevillii, Eptesicus fuscus y Choeronycteris mexicana), en

bosques de galería, y bosques de encino-pino. Este listado preliminar permite especular sobre el buen estado de salud del ecosistema en esta

área natural protegida y da pauta para llevar a cabo monitoreo en otras Islas Serranas en la misma área natural protegida:Sierra La Madera,

El Tigre, Buenos Aires y La Púrica. La intención de este trabajo es utilizar a los quirópteros como una herramienta para evaluar el grado de

conservación de estas Islas Serranas; ya que son abundantes, diversos y relativamente fáciles de monitorear, con costos bajos en el material

para su estudio. La presencia y/o ausencia de comunidades de murciélagos permitirá tomar decisiones de manejo y conservación a corto y largo

plazo en ésta área natural protegida.

JONES, LAWRENCE C. Coronado National Forest, 300 West Congress, Tucson, Arizona 85701, ljones02@fs.fed.us.

Changes in a lizard assemblage at an ecotone in southeastern Arizona. The Madrean Archipelago and its associated valleys have the highest

diversity of lizards in the United States. This is due to a convergence of ecoregions in an area that provides excellent environmental conditions

for life history needs of terrestrial ectotherms. The study area, near Safford, Arizona, is known to have about 20 species of sympatric lizards,

although only about one-half are common. The lizard community is typical of Sonoran and Chihuahuan deserts, semi-desert grasslands, and

lower Madrean and boreal woodlands. It has recently been suggested that lizard species are expected to decline globally due to climate change

and other factors. The study site is located at an ecotone between desert and grasslands in a foothills situation, as ecotones are expected to be

sensitive to environmental change. Study objectives were to (1) assess the baseline lizard community and (2) detect changes in the lizard assemblage over time. In 2003, 2010, and 2011, about 4,000 lizards were recorded on transects. Early data returns suggest that adaptable desert

and grassland species are common, while other, more specialized or mesic species appear to be trending toward a decline. Lizards may be better

bio-indicators of environmental change than the plant communities in which they occur.

Cambios en un ensamble de lagartijas en un ecotono del sureste de Arizona. El Archipiélago Madrense y sus valles tienen la más alta

biodiversidad de lagartijas en Estados Unidos. Esto se debe a la convergencia de eco regiones en una zona que posee condiciones ambientales

excelentes para el ciclo de vida de organismos ectotérmicos terrestres. El área de estudio, cerca de Safford, Arizona, tiene aproximadamente

20 especies de lagartijas simpátricas, aunque sólo alrededor de la mitad son comunes. La comunidad de lagartijas es típica de los Desiertos

Sonorense y Chihuahuense, praderas semidesérticas y bosques Madrenses y boreales más bajos. Recientemente se publicó que se espera una

disminución mundial de lagartijas por el cambio climático y otros factores. El sitio de estudio está ubicado en un ecotono entre desierto y

pastizal en el piedemonte, ya que se espera que los ecotonos sean sensibles al cambio climático. Los objetivos del estudio fueron (1) evaluar la

comunidad de lagartijas y (2) detectar cambios en el ensamble de lagartijas. En 2003, 2010 y 2011, cerca de 4.000 lagartijas se documentaron

en los transectos. Los primeros datos sugieren que las especies adaptables de los desiertos y los pastizales son comunes, y parece ser que especies más especializadas o mésicas van en declive. Las lagartijas pueden ser mejores bioindicadores del cambio ambiental que las comunidades

vegetales donde ocurren.

JONES, ROBERTW. Facultad de Ciencias Naturales, UniversidadAutónoma de Querétaro,Avenida de las Ciencias,s/n,Juriquilla, C.P. 76230,

Queretaro, México, rjones@uaq.mx.

What do we know of the Insects of Sonora, México? The insects reported from the Mexican state of Sonora are limited given its area and

diversity, with over-representation of several orders. Of the records in the Madrean Archipelago Biodiversity Assessment (MABA) database,

much of which isfrom records of the Mexican National Commission for the Knowledge and Use of Biodiversity (CONABIO), there are reported

a total of 10,341 insect specimen records. Of these records, 65% are Lepidoptera, 12% are Odonata and 9 % are Hymenoptera; a typical collection bias(butterflies and dragonflies) for poorly collected regions. Two of most diverse insect orders, Coleoptera and Diptera, together represent

less than 5% of the present database, of which 70% are scarabs and aquatic beetles.Alatitudinal analysis of the butterfly fauna will be presented;

the trends of which, may be indicative of other insect taxa. Clearly, international cooperation between scientists in México and the United States

is necessary to coordinate new collection efforts and share systematic knowledge of the insects of Sonora, and specifically their Sky Islands. 

¿Qué sabemos sobre los insectos de Sonora, México? Losinsectosreportados del estado de Sonora son insuficientes dado su área y diversidad,

y sobre cargados en varios órdenes. De los registros de la base de datos MABA (Evaluación de la biodiversidad del Archipiélago Madrense),

la mayoría de CONABIO (Comisión Nacional para el Conocimiento y Uso de la Biodiversidad) se reportan un total de 10,341 registros de

ejemplares de insectos. De estos registros, 65% corresponden a Lepidoptera, 12% a Odonata y 9% a Hymenoptera, un sesgo de colecta típico

(mariposas y libélulas) para regiones muy poco colectadas. Coleoptera y Diptera, dos de los órdenes más diversos, juntos representan menos

del 5% de la actual base de datos y de éstos el 70% corresponde a escarabajos e invertebrados acuáticos. Se presentará un análisis latitudinal de

la fauna de mariposas, donde las tendencias pueden ser indicadoras de otros taxones de insectos. Es evidente que se necesita una cooperación
546 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

internacional entre investigadores de México y E.U. para coordinar nuevos esfuerzos de colecta y compartir el conocimiento sistemático de los

insectos de Sonora, y en especial de sus Islas Serranas.

KAPLAN, MATTHEW E. Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721; Division of Biotechnology, Arizona Research Laboratories, University of Arizona, Tucson, Arizona 85721, mkaplan@email.arizona.edu.

Phylogeography of Yarrow’s Spiny lizard: What didn’t happen in the Holocene. Estimates from radiocarbon-dated packrat middens indicate that the high elevation woodland communities of the Madrean Sky Islands were continuous as recently as 8,000 to 12,000 years ago. A

number of population studies on a diverse collection of taxa have investigated the extent to which the Madrean Sky Island system has limited

gene flow among mountain ranges. The results of several of these studies indicate that population divergences may be more ancient than the

Holocene. Yarrow’sspiny lizards, Sceloporus jarrovii, were sampled from eightsitesrepresenting seven mountain ranges. DNAsequencesfrom

the lizards were used to reconstruct their evolutionary relationships and estimate the ages of the populations. The findings of these analyses

indicate that the sky island populations of S. jarrovii have been isolated for hundreds of thousands of years and did not experience gene flow

during the last woodland expansion.

Filogeografía de la lagartija Sceloporus jarrovii: Lo que no pasó en el Holoceno. Estimaciones de paleomadrigueras de roedores del género

Neotoma fechadas con radiocarbono indican que las comunidadesforestales en elevaciones altas de lasIslas Serranas Madrensesfueron continuas

hasta hace como 8.000 a 12.000 años. Varios estudios de población en diversos grupos taxonómicos han investigado hasta qué punto las Islas

Serranas Madrenses ha limitado el flujo genético entre cordilleras. Resultados de estos estudios indican que las divergencias de las poblaciones

son más antiguas que el Holoceno. Se estudiaron lagartijas Sceloporus jarrovii en ocho sitios que representan siete cordilleras. Las secuencias

de ADN de las lagartijas se usaron para reconstruir sus relaciones evolutivas y calcular las edades de las poblaciones. Los resultados de los

análisis indican que las poblaciones de S. jarrovii de las Islas Serranas han estado aisladas por cientos de miles de años y no experimentaron el

flujo genético durante la última expansión de los bosques.

KENNEDY, LINDA1 and Dan Robinett2. 1National Audubon Society, Appleton-Whittell Research Ranch, Elgin, AZ; 2Robinett Rangeland

Resources, Elgin, AZ.

Vegetation monitoring on semi-arid grasslands ungrazed by domestic livestock. The Research Ranch is an 8000 acre sanctuary and research

facility in the semi-arid grasslands of southeastern Arizona, USA. Cattle were removed from the property in 1968 to provide a reference area

by which various land uses such as grazing and exurbanization could be evaluated. Vegetation transects were established in 2000 and 2003

on several ecological sites in Major Land Resource Area 41. This monitoring program has tracked changes after wildfires and during drought.

Trends associated with non-native, invasive species, particularly Eragrostis lehmanniana (Lehmann lovegrass) and E. curvula (Boer lovegrass)

have been documented. For example, in an 8-year span, the frequency of Lehmann lovegrass on one loamy upland site grew from 1% to 60%

at the expense of the native Eragrostis intermedia (Plains lovegrass) and Bouteloua gracilis (Blue grama). Results from monitoring provides

feedback to Audubon personnel in the management of the property and are shared with other land managers to help tease apart the effects of

grazing and other land use actions from the effects of climate.

Monitoreo de vegetación en praderas semiáridas libres de ganado. La estación biológica Appleton-Whittell es una reserva de 3,237 has

y un centro de investigación en las llanuras semiáridas del sureste de Arizona, E.U. Se removió el ganado en 1968 para tener una referencia

para evaluar cambios de uso del suelo como el pastoreo y la urbanización previos. Se establecieron transectos de vegetación en varios sitios

ecológicos de Major Land Resource Area 41, durante 2000 y 2003. Este programa de monitoreo ha registrado cambios después de incendios

y durante las sequías. Se han documentado los efectos de las especies introducidas e invasoras, especialmente Eragrostis lehmanniana y E. 

curvula. Por ejemplo, en un lapso de 8 años, la frecuencia de E. lehmanniana en una altiplanicie con suelo franco aumentó de 1% a 60% a

expensas de los zacates nativos Eragrostis intermedia y Bouteloua gracilis. Los resultados del monitoreo sirven al personal de Audubon en la

administración de la parcela y se comparten con otros administradores de tierras para ayudar a distinguir los efectos del pastoreo y otros usos

del suelo de los efectos del cambio climático.

KETCHAM, SHARI L.1 and John L. Koprowski2 . 1School of Natural Resources and the Environment, University of Arizona, Tucson, AZ

85721, sketcham@email.arizona.edu; 2School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721, squirrel@

ag.arizona.edu.

Impacts of wildfire on wildlife in Arizona: a synthesis. Due to a century of fire suppression practices, the Madrean Archipelago regions

in Arizona have accumulated excessive fuel loads, which increase wildfire sizes, intensities, and frequencies. These drastic changes in forest

ecosystems can either benefit or adversely affect wildlife species. Therefore, it is imperative to understand how wildlife species react to such

ecosystem changes after wildfires in both the short-term and long-term time periods. I will examine scientific literature to determine wildlife

distribution, abundance, elevational migration, and behavioral changes (i.e. how wildlife use resources post-fire). Understanding the impacts of

wildfire-induced habitat fragmentation and creation of edge effects on wildlife species will provide us information about overall forest condition and forest management practices.
USDA Forest Service Proceedings RMRS-P-67. 2013 547

Abstracts/ Resúmenes

Síntesis del impacto de incendios en la fauna de Arizona. Debido a un siglo de supresión de incendios, las regiones del Archipiélago Madrense en Arizona han acumulado combustible excesivo que aumenta el tamaño, intensidad y frecuencia de incendios. Estos cambios drásticos

en los ecosistemas forestales pueden beneficiar o perjudicar la fauna silvestre. Por eso, es imperativo entender como la fauna reacciona a

tales cambios ecosistémicos después de los incendios a largo y corto plazo. Analizaré la literatura científica para determinar la distribución, la

abundancia, la migración altitudinal y los cambios de conducta (ej. uso de recursos después del incendio) de la fauna. Entender los impactos

de la fragmentación y la creación de efectos de borde inducidos por los incendios en la fauna silvestre aportará información sobre la condición

general del bosque y prácticas de manejo forestal.

KOPROWSKI, JOHN L., Melissa J. Merrick, Brittany Oleson, Erin E. Posthumus, Nicolas Ramos-Lara, Timothy G. Jessen, and R. Nathan

Gwinn. University of Arizona School of Natural Resources and the Environment, 1311 E. 4th St., Tucson, AZ 85721, squirrel@ag.arizona.edu;

mmerrick@email.arizona.edu; brit@email.arizona.edu; eposthum@email.arizona.edu; ramosln@yahoo.com.mx; tjessen@email.arizona.edu;

jessnate2001@yahoo.com.

It’s lonely at the top: Endemic biodiversity at risk to loss from climate change. Climate change is a serious immediate and long-term threat

to wildlife species. State and federal agencies are working with universities and non-government organizations to predict, plan for, and mitigate such uncertainties in the future. Endemic species may be particularly at risk as climate-induced changes impact their limited geographic

ranges. The Madrean Archipelago is characterized by high levels of endemism, natural fragmentation, and increasingly poor connectivity

among the mesic montane islands within an arid matrix of low elevation deserts and grasslands. The region already has experienced an increase

in temperature and this trend is predicted to accelerate over the remainder of the century. We assessed patterns of endemism and elevational

distribution of terrestrial vertebrates within the Madrean Archipelago. We examined incremental temperature increases and determined how

much endemic wildlife biodiversity is at risk to be lost from montane islands due to elevational and geographic restrictions. Endemism of nonvolant and smaller bodied species is greater than for more vagile forms. Taxa also differ in their susceptibility to the impacts of climate change.

We review the levels of risk, assess patterns of concern among the vertebrates, and discuss the implications of climate change on the endemic

vertebrate fauna of the region.

Soledad en la cima: La biodiversidad endémica en riesgo de perderse por el cambio climático. El cambio climático es una amenaza seria

y a largo plazo para las especies de flora y fauna..Agencias estatales y federales están trabajando con universidades y organizaciones no- gubernamentales para predecir, prepararse, y mitigar tales incertidumbres en el futuro. Las especies endémicas particularmente estarán en riesgo

cuando cambios en el clima afecten suslimitadas áreas de distribución. ElArchipiélago Madrense se caracteriza por niveles altos de endemismo,

fragmentación natural y una baja conexión entre las sierras montañosas mésicas en la matriz árida de pastizales y desiertos bajos. La región ya

ha experimentado un aumento de temperatura y se predice que ésto se acelere durante el resto del siglo. Evaluamos patrones de endemismo y

distribución en un rango de elevación de vertebrados terrestres del Archipiélago Madrense. Analizamos incrementos graduales de temperatura

y determinamos cuanta biodiversidad de fauna endémica está en peligro de perderse de las islas montañosas por restricciones altitudinales y

geográficas. El endemismo es más común en las especies no-volantes y más pequeñas que en las formas más móviles. Los taxones también

difieren en su vulnerabilidad a los impactos del cambio climático. Examinamos y evaluamos los niveles de riesgo de los vertebrados y analizamos las implicaciones del cambio climático en la fauna vertebrada endémica de la región.

LABORÍN-SIVIRIÁN, ERNESTO ALONSO1, Hugo Silva-Kurumiya1 , Rosa Elena Jiménez-Maldonado2, Gertrudis Yanes-Arvayo1. 1Universidad de la Sierra, División de Ciencias Biológicas, Moctezuma, Sonora. 2Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos

Bavispe, Cananea, Sonora.

Feline monitoring in Sierra La Madera, Municipio de Moctezuma, Sonora, Mexico. This work was completed in the region known in

Sierra La Madera, Fracción 5 of the Reserva Forestal Nacional y Refugio de. Fauna Silvestre Sierra de Ajos-Bavispe Spanningan area of 97,

542.74 has, which in recent years has been influenced by ranching and illegal hunting, it is necessary to know the current biological state and

abundance of the felines of the area. Monitoring efforts rely on the use of 20 camera traps that were distributed in several places in La Sierra,

for three different periods from August to December 2011. During the first period three records of mountain lion (Puma concolor), and three

records of bobcat (Lynx rufus) were obtained. During the second period,seven records of mountain lion and two records of bobcat were obtained.

When monitoring is complete it will be necessary to obtain prey abundance and feline activity patterns.

Monitoreo de felinos en Sierra La Madera, Municipio de Moctezuma, Sonora, México. El presente trabajo se realizó en la región conocida

como Sierra la Madera, Fracción V de la Reserva Forestal Nacional y Refugio de. Fauna Silvestre Sierra deAjos-Bavispe.Abarca una superficie

de 97, 542.74 ha, que en los últimos años han tenido influencia por ganadería y cacería ilegal, es necesario conocer el estado biológico actual

y la abundancia de felinos del área. El monitoreo está basado en el uso de 20 cámaras trampa, distribuidas en varios puntos de la Sierra, en tres

campañas de agosto a diciembre de 2011. Durante la primera campaña se obtuvieron tres registros de puma (Puma concolor) y tres registros de

gato montés (Lynx rufus). En la segunda campaña se obtuvieron siete registros de puma y dos registros de gato montés.Al finalizar el monitoreo

se estima obtener abundancia de presas y patrones de actividad de felinos.
548 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

LACROIX, KELLY MOTT, Joanna B. Nadeau, Brittany Choate, and Sharon B. Megdal. Water Resources Research Center, University of

Arizona, Tucson,AZ 85721, klacroix@email.arizona.edu; jnadeau@cals.arizona.edu; bchoate@email.arizona.edu;smegdal@cals.arizona.edu.

Assessing and addressing ecological water needs in Arizona. The University of Arizona Water Resources Research Center (WRRC) has

published two technical documents from a project to assess Arizona’s environmental water needs. The Arizona Environmental Water Needs

Assessment (AzEWNA) Report and theArizona Environmental Water NeedsAssessment Methodology Guidebook for the first time explore and

synthesize efforts to quantify the water needs of river systems in Arizona. The Report summarizes the state of knowledge about environmental

water needs, explores the information gaps and contains maps and graphics to help readers visualize the extent of the inventory. The Guidebook

describes the methods employed to quantify Arizona’s environmental flows and provides decision trees for those designing environmental

flow studies. The WRRC is now working to improve understanding about ecological water needs and build upon continuing efforts to address

increasing water demands around the state. Regionally focused materials developed from the AzEWNA work and other resources are being

made available for water planning through a series of outreach presentations throughout the state. Our presentation at the Madrean Archipelago

conference will highlight our findings on water needs of Arizona’s environment and initial progress made with those involved in regional and

state processes exploring what it means to consider the environment in water planning.

Evaluación de la situación ecológica de agua en Arizona. El Centro de Investigación de Recursos Hídricos (WRRC, por sus siglas en inglés)

de la Universidad de Arizona tiene dos publicaciones técnicas de un proyecto de evaluación de la necesidad hídrica del medio ambiente en

Arizona. El Informe y la Guía sobre las necesidades hídricas medioambientales de Arizona (AzEWNA por sus siglas en inglés) por primera

vez exploran y sintetizan los trabajos para cuantificar las necesidades hídricas de los sistemas de ríos en Arizona. El Informe compila todo el

conocimiento sobre las necesidades hídricas medioambientales, analiza las lagunas de información e incluye mapas y diagramas para ayudar a

los lectores a visualizar el alcance del estudio. La Guía describe los métodos utilizados para cuantificar los flujos medioambientales de Arizona

y contiene árboles de decisión para quienes estén diseñando estudios de los flujos medioambientales. El WRRC está trabajando para mejorar

la comprensión de las necesidades ecológicas de agua y fortalecer el trabajo continuo para enfrentar la creciente demanda de agua en el estado.

Materiales para la región elaborados en base a los resultados de AzEWNA y otros recursos se están proporcionando para la planificación del

uso de agua por medio de presentaciones al público del estado. Esta presentación hará hincapié en nuestras conclusiones sobre las necesidades

hídricas del medioambiente de Arizona y el progreso inicial logrado con quienes trabajan en los procesos regionales y estatales, y que tienen

interés en considerar el medioambiente en la planificación hídrica.

LAMBERTON, JESSICAA., and SergioAvila-Villegas. Sky IslandAlliance, 300 E. University Blvd., Tucson,AZ 85705, jessica@skyislandalliance.org, sergio@skyislandalliance.org.

A decade of wildlife tracking in the Sky Islands. In 2001 Sky Island Alliance developed a citizen science program that uses track and sign

identification and count surveys to monitor potential wildlife corridors throughout southeastern Arizona and southwestern New Mexico. The

goal of the Wildlife Linkages Program is to protect and advocate for an interconnected landscape where wildlife, based on their ecological

needs, can move easily between core habitats, the Sky Island mountain ranges. Currently, we train and engage volunteers in the monitoring of

fifty 1.5 mile-long transects within seven priority linkage areas; the majority of these study areas are located on public lands. To date we have

conducted over 1,000 track count surveys and documented over 4,100 records for more than 40 different animal species in the region. Sky

Island Alliance has successfully applied the resulting species presence data to land-use policy and permanent land conservation, incorporating

wildlife data and corridor priorities into the Sonoran Desert Conservation Plan, the Santa Cruz County Comprehensive Plan, the Pima County

Wildlife Connectivity Assessment and the Arizona Wildlife Linkages Assessment.

Una década de rastreo de fauna en las Islas Serranas. En 2001 Sky Island Alliance desarrolló un programa que usa la identificación de

huellas y marcas de animales y cuenta los transectos para monitorear corredores potenciales de fauna en el sureste de Arizona y suroeste de

Nuevo México. La meta del Programa de Enlaces de Fauna Silvestre es proteger y abogar por un paisaje interconectado donde la fauna, con

base en sus necesidades ecológicas, pueda moverse fácilmente entre sus hábitats principales, las cordilleras de las Islas Serranas. Actualmente,

se capacita y trabaja con voluntarios en el monitoreo de cincuenta transectos de 2.5 kilómetros de longitud ubicados en siete áreas prioritarias;

la mayoría en áreas naturales del gobierno. A la fecha hemos realizado más de 1,000 muestreos usando huellas y documentado más de 4,100

registros de más de 40 diferentes especies animales en la región. La información resultante sobre la presencia de especies y su análisis, ha sido

aplicada exitosamente a políticas de uso de suelo y conservación permanente de tierras, incluyendo la incorporación de información sobre fauna

y corredores prioritarios en el Plan de Conservación del Desierto Sonorense, el Plan Comprehensivo del Condado Santa Cruz, la Evaluación

de Conectividad de Fauna del Condado Pima y la Evaluación de Enlaces para la Fauna Silvestre de Arizona.

LAPORT, ROBERT G. and Robert L. Minckley. Department of Biology, University of Rochester, 213 Hutchison Hall, River Campus, Rochester,

NY 14627, robert.laport@rochester.edu.

Ecological divergence of Larrea tridentata at the Chihuahuan-Sonoran Desert ecotone. The Chihuahuan and Sonoran Desert floras have

evolved largely in isolation, separated by the Sierra Madre Occidental. Biogeographers have debated the location of the present-day boundary

between these deserts, although Shreve, among others, suggested this transition occurred in the low elevation areas of the Gila and San Pedro

river drainages of Arizona. Now desert scrub, this area is thought to have been grassland approximately 100 years ago. The dominant and

widespread shrub creosote bush (Larrea tridentata (DC.) Coville; Zygophyllaceae) occurs throughout the warm deserts of North America, with

three cytotypes that are broadly distributed in the Chihuahuan (diploid), the Sonoran (tetraploid and hexaploid), and the Mojave (hexaploid)
USDA Forest Service Proceedings RMRS-P-67. 2013 549

Abstracts/ Resúmenes

Deserts. Here, we describe an abrupt (~10 km) transition between diploid and tetraploid L. tridentata in the Gila and San Pedro River drainages.

This cytotype transition, and predictions from ecological niche modeling, suggests a persistent ecotone between the Chihuahuan and Sonoran

Deserts and raises interesting questions about the historical extent of these biogeographic provinces. Future efforts to obtain packrat (Neotoma)

midden data from the low elevations of the eastern Catalina and northwestern Galiuro Mts. would be particularly informative for determining

ancient botanical communities of the area and further elucidate the desert boundaries.

La divergencia de Larrea tridentata en el ecotono de los desiertos Chihuahuense y Sonorense. La flora de los desiertosChihuahuense y Sonorense ha evolucionado en gran parte aislada, separada por la Sierra Madre Occidental. Los biogeógrafos han debatido la ubicación de la frontera

actual entre estos desiertos, aunque Shreve, sugirió que esta transición ocurre en las zonas de baja altitud de los drenajes de los ríos Gila y San

Pedro de Arizona. Esta área actualmente es matorral desértico, pero se cree que hace aproximadamente 100 años fue pastizal. La gobernadora

(Larrea tridentata (DC.) Coville; Zygophyllaceae), común y con distribución amplia, ocurre en todoslos desiertos cálidos de Norteamérica, con

tres citotipos distribuidos extensamente en el Desierto Chihuahuense (diploide), el Desierto Sonorense (tetraploide y hexaploide) y el Desierto

Mohave (hexaploide). Aquí describimos una transición abrupta (~10 km) entre L. Tridentata diploide y tetraploide en los drenajes del río Gila

y río San Pedro. Esta transición de citotipo, y las predicciones de modelos de nichos ecológicos, sugiere que hay un ecotono persistente entre

los desiertos Chihuahuense y Sonorense y hace pensar sobre el alcance histórico de estas provincias biogeográficas. Estudios futuros con datos

de paleomadrigueras de roedores del género Neotoma de elevaciones bajas del este de la Sierra Catalina y del noroeste de la S. Galiuros serían

especialmente informativos para identificar las comunidades botánicas antiguas de la zona y aclarar más las fronteras desérticas.

LARA-DÍAZ,NALLELIE.1,CarlosA.López-González1,HelíCoronel-Arellano2,AlejandroGonzález-Bernal2. 1Facultad deCienciasNaturales,

Universidad Autónoma de Querétaro, Av. de las Ciencias S/N, Col. Juriquilla, Santiago de Querétaro, Querétaro, C.P. 76230; 2Naturalia A.C,

Av. 32 entre Calle 3 y 4, Villa Satélite 3250, Agua Prieta, Sonora, C.P. 84200.

Black Bear population and connectivity in the Sky Islands of Mexico/United States. The Sky Island region is a mountainous region surrounded by grasslands, deserts and intermountain valleys, located between Mexico and the United States. However, different land management

and human impact can have an effect on its wildlife populations. Currently, the border wall poses an immediate threat to the survival of black

bears (Ursus americanus), considered an endangered species in Mexico. Our aim was to determine the conservation status of black bear in the

Sierra San Luis as affected by the border fence. We determined population size through camera trap and radio-telemetry, and modeled population

occupancy using PRESENCE. We documented a bear population with more than 500 individuals. Surveys along the border failed to detect bears

crossing it, but we identified linkages between the two countries important for future landscape planning. Increased vehicular traffic, migration

and drug traffic have a negative effect on bear populations, exacerbated by an increase in anthropogenic activities resulting from the construction and maintenance of the border wall. We recommend modifications to the structure of the border wall, and to increase wildlife monitoring

by the United States authorities to reduce the potential impact this structure has on black bear and other wildlife populations.

Población de oso negro y su conexión en las Islas Serranas de México y Estados Unidos. La región de las Islas Serranas es un área montañosa rodeada por pastizales, desiertos y valles ubicada en México y Estados Unidos. Sin embargo, un manejo e impacto humano diferente

pueden afectarsus poblaciones de fauna.Actualmente, el muro fronterizo representa una amenaza inmediata para la supervivencia del oso negro

(Ursus americanus), una especie en peligro en México. El objetivo de este trabajo fue determinar como el muro fronterizo afecta el estado

de conservación del oso negro en la Sierra San Luis. Determinamos el tamaño de la población mediante cámaras trampa y radio-telemetría, y

se hizo un modelo de población usando PRESENCE. Registramos una población de osos con más de 500 individuos. Monitoreos cerca de la

línea fronteriza no detectaron osos cruzándola, pero identificamos corredores entre los dos países importantes para el manejo futuro del paisaje.

Aumento del tráfico vehicular, migración humana y tráfico de drogas tienen un efecto negativo en las poblaciones de osos, empeorado por un

aumento de actividades antropogénicas por la construcción y mantenimiento del muro. Recomendamos modificaciones a la valla, y un aumento

del monitoreo de fauna por parte de Estados Unidos para reducir el posible impacto de esta estructura en las poblaciones de oso negro y otras

especies de fauna.

LARA-DÍAZ, NALLELI E.1, CarlosA. López-González1, Helí Coronel-Arellano2,Alejandro González-Bernal2 , Gerardo Carreón-Arrollo2 y

Rosa E. Jiménez-Maldonado3. 1Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Av. de las Ciencias S/N, Col. Juriquilla,

Santiago de Querétaro, Querétaro, C.P. 76230; 2Naturalia A.C. Av. 32 entre Calle 3 y 4 Villa Satélite 3250, Agua Prieta, Sonora, C.P. 84200;

3CONANP, Reserva Forestal Nacional y Refugio de Fauna Silvestre Ajos Bavispe, Calle 2a Este final y Sinaloa. Col Centro. Cananea, Sonora,

C.P. 84620.

Mammalian diversity in the Sky Islands, Sonora, México. The sky island system of northwestern Mexico has been known as a center for

biodiversity, nevertheless land tenure and management of populations may create alternative patterns of diversity. The objective of this study

was to estimate the mammalian species richness and diversity of the sky islands of northeastern Sonora, Mexico. During 2009-2011, we used

camera trapping (between 21 to 40 per location) at 13 sites, at mountain ranges and lower elevations. We determined species richness through

species accumulation curves. We documented a total of 27 species, with a range of seven to 21 per site, finding significant differences between

sampled areas. Jaccard ́s similarity index yield a variation between 32 to 95% among sites. Our results show a different composition of species,

with relevant populations of endangered species such as black bears (Ursus americanus), and also important connectivity for species such as

badger (Taxidea taxus), jaguar (Panthera onca), and ocelot (Leopardus pardalis).
550 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

La diversidad de mamíferos en las Islas Serranas de Sonora, México. El sistema de las Islas Serranas del noroeste de México se conoce

como centro de biodiversidad, sin embargo la tenencia de tierras y el manejo de poblaciones puede resultar en patrones diferentes de diversidad.

El objetivo de este estudio fue estimar la riqueza y diversidad de mamíferos de las Islas Serranas del noreste de Sonora, México. De 2009 a

2011, usamos cámaras trampa (de 21 a 40 por lugar) en 13 sitios, en cordilleras y altitudes más bajas. Determinamos riqueza de especies usando

curvas de acumulación. Registramos 27 especies, variando de 7 a 21 por sitio, con diferencia significativa entre áreas. El índice de similitud de

Jaccard da una variación entre 32 y 95% entre los sitios. Nuestros resultados muestran una composición variada de especies, con poblaciones

relevantes de especies en peligro como el oso negro (Ursus americanus), y también conectividad importante para especies como tejones(Taxidea

taxis), jaguar (Panthera onca) y tigrillo (Leopardus pardalis).

LENART, MELANIE1 and Christopher Jones2. 1University of Arizona, Department of Soil, Water and Environmental Science, 429 Shantz

Building, 1177 E. Fourth Street, Tucson, AZ 85721, mlenart@email.arizona.edu; 2University of Arizona, Gila County Cooperative Extension,

5515 S Apache Avenue, Suite 600, Globe, AZ 85501, ckjones@cals.arizona.edu.

Perceptions about climate change influence support for forest adaptation measures. Land managers can draw from a toolbox of potential

management responses in an effort to adapt to climate change and mitigate some of its impacts. In order to consider the level of acceptance for

various adaptation responses among land managers and scientists, we administered a detailed anonymous survey. It was distributed at a May

2011 National Workshop on Climate and Forests held in Flagstaff, Arizona, and electronically by the Society of American Foresters and the Association of Natural Resource Extension Professionals. Results compiled from the 1,029 U.S.-based respondents will be described. The research

was designed to consider whether participants’ perceptions about climate change influenced their perceived acceptance of various climate and

proxy records and their willingness to consider different forest adaptation and mitigation management techniques. Another research goal was

to foster a greater understanding of how researchers and educators can better provide for the needs of forest managers. The results also provide

an indication of the acceptability of specific management techniques to the group as a whole and by specific sector. Results will be considered

for the nation as a whole and, using a subset of respondents who provided geographic information, for the Southwest specifically.

Percepciones sobre el cambio climático influyen el respaldo a medidas de adaptación forestal. El personal de manejo de recursos puede

escoger entre posibles formas de manejo con el fin de adaptarse al cambio climático y mitigar algunos de sus impactos. Para considerar el nivel

de aceptación de varios métodos de adaptación entre personal de manejo y científicos, aplicamos un cuestionario anónimo. Se distribuyó en el

Taller Nacional de Clima y Bosques en Flagstaff, Arizona en mayo de 2011, y electrónicamente por la Sociedad de Silvicultores Americanos y

la Asociación de Profesionales de Extensión de los Recursos Naturales. Los resultados recopilados de 1,029 encuestados en E.U. se describirán

a continuación. La investigación se diseño para considerar si las percepciones de los participantes sobre el cambio climático influyeron su aceptación de varios registros y estimaciones climáticas y su disposición para considerar técnicas diferentes de adaptación forestal y de manejo de la

mitigación. Otra meta de la investigación fue entender mejor cómo losinvestigadores y educadores pueden apoyar al personal de manejo forestal.

Los resultados también indican la aceptación de técnicas de manejo específicas por todo el grupo y por sectores específicos. Los resultados se

considerarán para la nación en conjunto y usando los cuestionarios que incluyen información geográfica, para el Suroeste, específicamente.

LEVICK, LAINIE1, Phillip Guertin1, David Goodrich2, Joel Murray3, Russell Lyon1, Samantha Hammer1, Brian Bledsoe3, and Melinda Laituri3. 1

UniversityofArizona,Tucson,AZ; 2USDA-ARS/SouthwestWatershedResearchCenter,Tucson,AZ; 3ColoradoStateUniversity,FortCollins,CO.

A classification of intermittent and ephemeral streams on Department of Defense land. Ephemeral and intermittent streams are the predominant fluvial forms in arid and semiarid environments. Various studies have shown biodiversity and habitat values in these lands to be

considerably higher along ephemeral and intermittent stream corridors in comparison to adjacent uplands. Yet, knowledge of how hydrologic

properties of these systems are linked to their ecological function is limited, leading to difficulty in making well-informed management decisions. To improve decision making, this research will develop an ecohydrologically-based classification scheme that distinguishes ephemeral

and intermittent channel types by a set of biotic and abiotic attributes that are directly related to their hydrologic regime. Using the stream type

classification, we will develop an assessment methodology that allows Department of Defense managersto evaluate the impacts of perturbations

(e.g., climate change, military activities) on the hydrologic regimes of these systems and the species that depend on them. Methods include

analysis of GIS and high-resolution remote sensing data, and hydrologic modeling (AGWA/KINEROS/SWAT). Statistical approaches such as

classification and regression trees will be used to classify ephemeral and intermittent streams along a xeric to mesic gradient.

Una clasificación de arroyos intermitentes y efímeros en las tierras del Departamento de Defensa. Los arroyos intermitentes y efímeros

son las formas fluviales predominantes en los ambientes áridos y semiáridos. Varios estudios han mostrado que los valores de la biodiversidad

y el hábitat en estas tierras son considerablemente más altas en los corredores de los arroyos efímeros e intermitentes comparados con las tierras

altas colindantes. Pero el conocimiento de cómo las propiedades hidrológicas de estos sistemas están vinculadas a sus funciones ecológicas es

limitado, haciendo difícil que se tomen decisiones de manejo bien informadas. Para mejorar la toma de decisiones, esta investigación desarrollará una clasificación ecohidrológica que identifica las corrientes intermitentes y efímeras por un grupo de atributos bióticos y abióticos

directamente relacionados al sistema hidrológico. Utilizando la clasificación de tipo de arroyo, desarrollaremos una metodología de evaluación

para que los administradores del Departamento de Defensa puedan evaluar los impactos de las perturbaciones (ej. cambio climático, actividades

militares) en los regímenes hidrológicos de estos sistemas y las especies que dependen en ellos. Los métodos incluyen análisis de datos de SIG,

sensores remotos de alta resolución y modelos hidrológicos (AGWA/KINEROS/SWAT). Métodos estadísticos como diagramas de regresión y

clasificación se utilizarán para identificar los arroyos intermitentes y efímeros en un gradiente de árido a mésico.
USDA Forest Service Proceedings RMRS-P-67. 2013 551

Abstracts/ Resúmenes

LLANO-SOTELO. JOSÉ M.1and Alejandro E. Castellanos-Villegas1. 1DICTUS, Universidad de Sonora, Hermosillo, Sonora, CP 83000, llanojm@correom.uson.mx; acastell@guaymas.uson.mx.

Ecologic aspects related with the potential of species of the Sonoran Desert for the use of biofuels. The use of biofuels is proposed for an

alternative to decrease the negative effects related with the climatic change for the use of fossil fuels. In arid lands there are a great proportion

of species than produce important quantities of oil in the seeds. In this study we determine the content and composition of oils (fatty acids) of

different families of arid land plants in Mexico. We did determinations in the laboratory with gas chromatography and bibliographic values to

their potential as a possible source for biofuels. The ease of combustion of a biofuel is determined by the cetane number, which is related to

the proportion of unsaturated and saturated fatty acids. The proportion of unsaturation of fatty acids is related to the environmental conditions

of place of origin of the species, which is higher in colder places. We found lower cetane numbers for a high proportion of unsaturated fatty

acids in the Brassicaceae family, followed by species in the families Asteraceae, Cucurbitaceae, Euphorbiaceae, Oleaceae, Malvaceae, and

Fabaceae. This reflects the capacity of the family Brassicaceae for growth in habitats colder than other families and the origin more tropical of

Fabaceae. This study propose a greater analysis of the content and composition of fatty acids in species of arid lands for the determination of

the potential of possible source of biofuels.

Aspectos ecológicos relacionados con el potencial de especies del Desierto Sonorense para su uso como biocombustibles. El empleo de

biocombustibles ha sido propuesto como alternativa a la disminución y los efectos negativos relacionados con el cambio climático derivados

de los combustibles fósiles. En zonas áridas existe una gran proporción de especies que producen cantidades importantes de aceites en sus

semillas. En este estudio se determinaron los contenidos y composición de aceites (ácidos grasos) de diversas familias de plantas de las zonas

áridas de México. Tanto determinaciones en el laboratorio por cromatografia de gases como valores obtenidos de la bibliografía mostraron un

potencial de fuentes posibles para uso como biocombustible. La facilidad con que un biocombustible efectúa su combustión esta determinada

con su número de cetano, el cual se ha relacionado a su vez con la proporción de ácidos grasos insaturados y saturados. La proporción de

insaturación de los ácidos grasos se ha encontrado estar relacionada con las condiciones ambientales de los lugares de origen de las especies,

siendo mayor esta proporción en lugares más fríos. Se encontró un menor número de cetano debido a una mayor proporción de ácidos grasos

insaturados en la familia Brassicaceae, seguido de especies en las familias Asteraceae, Cucurbitaceae, Euphorbiaceae, Oleaceae, Malvaceae

y Fabaceae. Lo anterior refleja también la capacidad de la familia Brassicaeae para crecer en habitats mas templados que las otras familias y

el orígen más tropical de Fabaceae. El estudio propone un mayor análisis del contenido y composición de ácidos grasos en especies de zonas

áridas para determinar su potencial como fuentes posibles de biocombustibles.

LYNCH, ANN M.1,2 and Christopher D. O’Connor3,2. 1U.S. Forest Service, Rocky Mountain Research Station; 2University of Arizona, Laboratory of Tree-Ring Research, 105 W Stadium, Tucson AZ 85721; 3

University of Arizona, School of Natural Resources and the Environment,

Tucson AZ 85721

An unprecedented mountain pine beetle outbreak in the Pinaleño Mountains. Mountain pine beetle (MPB, Dendroctonus ponderosae

Hopkins) has incurred an unprecedented outbreak in the Pinaleño Mountains of southeast Arizona. MPB has rarely been observed in southern

Arizona, is not recorded from the Pinaleños, and has no record as a significant damage agent south of Flagstaff. In recent years, though, it

has been killing southwestern white pine (SWWP, Pinus strobiformis) and occasional ponderosa pine (P. ponderosa) in small patches in the

Pinaleños. Mortality each year is minor, but the outbreak has persisted for at least 12 years and cumulative effects have reached ecologically

significant levels that affect resource values. We sampled MPB-infested plots and compared them to a random sample of SWWP-inhabiting

plots. Average stocking and SWWP dominance on infested plots were each approximately twice what was found in the random sample. Trees

as small as 13 cm dbh were attacked. Cross dating the attack years of killed trees indicates that the outbreak has persisted at least 12 years.

MPB population dynamics are temperature-responsive, and in this situation the life cycle appears to be univoltine. The ultimate course of the

outbreak is unknown, but is likely to significantly affect the character of the Pinaleño mixed-conifer forest.

Una plaga sin precedentes del descortezador de pinos en la Sierra Pinaleño. El descortezador (Dendroctonus ponderosae) ha provocado

una plaga sin precedentes en la Sierra Pinaleño del sureste de Arizona. El descortezador rara vez se ha observado en el sur de Arizona, no se ha

registrado en Pinaleños y no tiene antecedentes de ser un agente de daño significativo al sur de Flagstaff. Pero en años recientes ha matado el

pino blanco del suroeste (Pinus strobiformis) y ocasionalmente pino ponderosa (P. ponderosa) en zonas pequeñas en Pinaleños. La mortalidad

cada año es menor, pero la plaga ha persistido por lo menos 12 años y los efectos acumulativos han llegado a niveles ecológicamente significativos que afectan los recursos. Se hicieron muestreos en parcelas infestadas por descortezadores y se compararon con una muestra aleatoria de

parcelas con pino blanco. Tanto el promedio del arbolado como la dominancia de pino blanco en las parcelas infestadas fueron casi el doble de

la muestra aleatoria. Incluso árboles de 13 cm DBH fueron infestados. Al determinar los años de ataques de árboles muertos se concluye que

la plaga ha persistido por lo menos 12 años. La dinámica de la población del descortezador es sensible a la temperatura, y en esta situación el

ciclo de vida parece ser univoltino. Se desconoce como terminará esta plaga, pero es probable que afectará significativamente el bosque mixto

de coníferas de los Pinaleños.
552 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

LYNCH, ANN M. U.S. Forest Service, Rocky Mountain Research Station, 105 W Stadium, Tucson AZ 85721, alynch@fs.fed.us.

Insects as agents of change: Contemporary forest insect outbreaks in the Arizonan Sky Islands. Predicted responses of forest insect populations to warming climate include altered distribution of key species, especially movement to higher elevation and higher latitudes; acceleration

of herbivory rates; altered frequency and intensity of outbreaks; emergence of previously innocuous insects as pests; unpredictable interactions

with natural enemy populations (parasites, predators, pathogens); and invasion of new habitats. Many of the predicted responses have already

commenced in the Sky Islands. Recent forest insect activity in the Sky Islands has been remarkable for the multitude of insect species incurring

outbreaks, the severity of damage, the appearance of new species as pests, and the temperature-sensitive population dynamics ofseveralspecies.

Nine species have experienced severe or unprecedented outbreaks in the last 20 years. Other damaging insects and pathogens have been active

as well, especially after fire events and during drought years. Activity by so many species in such a short period of time in high elevation Sky

Island forests indicates that insect populations in low-latitude high-elevation forests may be particularly responsive to climate change. Though

the precise timing and nature of altered insect outbreak regimes is not predictable, it is certain that insects will play a major role in future change

in the Madrean Sky Island forest ecosystems.

Los insectos como agentes de cambio: Brotes de plagas de insectos forestales en las Islas Serranas de Arizona. Las respuestas previstas de

poblaciones de insectosforestales al calentamiento climático incluyen alteración en la distribución de especies claves, especialmentemovimiento

a elevaciones y latitudes más altas, incremento del índice de herbívoros, alteración de la frecuencia e intensidad de brotes,surgimiento de insectos

inofensivos como plagas, interacciones imprevisibles con poblaciones enemigas naturales (parásitos, predadores, patógenos), y la invasión de

nuevos hábitats. Muchas de las reacciones predichas ya han comenzado en las Islas Serranas. La actividad reciente de los insectos forestales en

las Islas Serranas ha sido notable por la multitud de insectos plaga, la severidad del daño, el surgimiento de nuevas especies como plagas, y la

dinámica de la población sensible a la temperatura en algunas especies. Nueve especies han experimentado brotes severos o sin precedentes en

los últimos 20 años. Otros insectos y patógenos dañinos han estado activos también, especialmente después de incendios y en años con sequía.

La actividad de tantas especies en tan corto tiempo en bosques de elevaciones altas de las Islas Serranas indica que las poblaciones de insectos

en bosques de latitud baja y elevación alta podrían ser más sensibles al cambio climático. Aunque el momento y condiciones específicos de

ataques de plagas no es predecible es seguro que los insectos tendrán un papel importante en los cambios futuros de los ecosistemas forestales

de las Islas Serranas Madrenses.

LYTLE, DAVID A. Department of Zoology, Oregon State University, Corvallis, OR 97331, lytleda@oregonstate.edu.

Floods, droughts, and aquatic biodiversity in the Madrean archipelago. The Madrean Sky Island region harbors a rich variety of aquatic

habitats, ranging from mountain streamsto lowland rivers and ciénegas. Compared with other regions, MSI aquatic habitats are unusual because

they are patchily distributed acrossthe landscape and strongly influenced by seasonal flooding and drought. Thisfragmented spatial arrangement

coupled with strong disturbance dynamics has shaped both the evolution and ecology of MSI aquatic organisms. Many aquatic insects, fish, and

riparian plants are now known to possess adaptations for surviving extreme events such as flash flooding or prolonged drought. Additionally,

population genetic studies have revealed marked evolutionary divergence of aquatic populations at fine spatial scales. In one case, populations

of an aquatic invertebrate predator have evolved unique behaviors to escape flash floods, and these behavioral adaptations differ in response

to local canyon-scale flood regimes. Disturbance dynamics and spatial isolation also facilitate high aquatic invertebrate biodiversity within a

surprisingly small amount of habitat, with largely distinct communities coexisting within meters of each other. This fine-scale partitioning of

genetic and species diversity is a double edged sword.Although changes in hydrologic regime due to climate change, groundwater pumping and

other factors have eliminated unique populations and communities, the diversity and adaptability of these organisms could provide resilience

to ecological change.

Inundaciones, sequías, y biodiversidad acuática en el Archipiélago Madrense. La región de las Islas Serranas Madrenses alberga una gran

variedad de hábitats acuáticos, desde los arroyos de montañas a los ríos y ciénegas de la tierra baja. Comparados con otras regiones, los hábitats

acuáticos de la región Madrense son diferentes por estar fragmentados y bastante influenciados por inundaciones y sequías de estación. Esta

fragmentación espacial combinada con fuertes dinámicas de disturbio ha influenciado la evolución y ecología de los organismos acuáticos de

la región Madrense. Ahora se sabe que muchos insectos acuáticos, peces y plantas ribereñas están adaptados para sobrevivir eventos extremos,

como son las crecidas o sequías prolongadas. Además, estudios genéticos poblacionales han revelado una marcada divergencia evolutiva de las

poblaciones acuáticas en escalas espaciales pequeñas. En un caso, las poblaciones de un depredador invertebrado acuático han evolucionado

comportamientos únicos para escapar de las crecidas, y estas adaptaciones conductuales difieren en repuesta a los regimenes locales de inundaciones a escala del cañón. Las dinámicas de perturbación y aislamiento espacial también facilitan una alta biodiversidad de invertebrados

acuáticos en un hábitat sorpreseivamente pequeño, con comunidades muy distintas coexistiendo separadas por unos cuantos metros. Esta división en una escala pequeña de diversidad genética y de especies es una espada de doble filo. Aunque los cambios en el régimen hidrológico

debido al cambio climático, extracción de agua subterránea y otros factores han eliminado poblaciones y comunidades únicas, la diversidad y

adaptabilidad de estos organismos podría aumentar la capacidad de recuperación del cambio ecológico.
USDA Forest Service Proceedings RMRS-P-67. 2013 553

Abstracts/ Resúmenes

MacEWEN, REBECCA. Saguaro National Park, 3693 S. Old Spanish Trail, Tucson, AZ 85730, Becky_MacEwen@nps.gov.

Analyzing changes in the spatial distribution of saguaros (Carnegiea gigantea) in Saguaro National Park. Results of the 2010 Saguaro

Census at Saguaro National Park were used to analyze changes in spatial distribution of the saguaro cactus both within plots and park-wide.

This was done by quantitatively comparing 11 historic 1-ha saguaro plots where individual saguaros were mapped in 1975 (by hand) and again

in 2010 (by GPS) with 45 standard 2-ha census plots. Recently collected LiDAR data and aerial imagery were used in conjunction with the

data from the saguaro census plots and additional high-elevation plots to evaluate elevation limitations of the existing population and create a

suitable habitat model. The derived saguaro habitat was then also projected 50 years into the future using predicted growth rates and climatic

factors to examine potential distribution of the species in the park. Results indicate a wide variety of expected changes in saguaro distribution

related to age class structure, landscape, presence of external influences, and overall population dynamics.

Análisis de cambios en la distribución espacial de sahuaros (Carnegiea gigantea) del área protegida Saguaro National Park. El censo de

sahuaros de 2010 de Sahuaro National Park se usó para analizar cambios en la distribución espacial de sahuaros tanto en parcelas de muestreo

como en todo el parque. Se compararon cuantitativamente 11 parcelas antiguas de 1 ha donde cada sahuaro se ubicó en un mapa en 1975 (a

mano) y de nuevo en 2010 (con GPS) en 45 parcelas de censo estándares de 2 ha. Se usaron datos recientes de LiDAR, fotografías aéreas, datos

del censo de sahuaros, y se incluyeron parcelas de elevaciones altas para evaluar las limitantes de elevación de la población existente y crear un

modelo del hábitat adecuado. El hábitat de sahuaro obtenido se extrapoló 50 años en el futuro usando tasas de crecimiento y factores climáticos para examinar la distribución potencial de la especie en el parque. Los resultados indican una amplia variedad de cambios probables en la

distribución de sahuaro relacionado con distribución por edad, paisaje, presencia de factores externos y dinámica de la población en general.

MAKINGS, ELIZABETH.Arizona State University School of Life Sciences, PO Box 874501, Tempe,AZ 85257, elizabeth.makings@asu.edu.

Vegetation and flora of the St. David Ciénega. In this talk I will explore the St. David and Lewis Springs Ciénegas along the San Pedro River

in Cochise County, Arizona – with an emphasis on the flora. In the Sky Island region, ciénegas are rare marshlands amidst arid surroundings

where groundwater perennially intersects the surface. Because of their unique soil and hydrological properties, ciénegas maintain a trajectory

toward graminoid wetlands. Evaporation usually causesthe water to be alkaline, and vegetation around a ciénega commonly includes halophytes

and other unusual species. Depending on their age and size, they may also harbor high levels of endemism. Ciénegas are far from pristine, and

like many wetlands and riparian areas, they have probably been exploited for millennia. However, in the American Southwest, these habitats

are hypothesized to have been in severe decline over the past 150 years due to a cycle of erosion and arroyo cutting. In light of their biological

and cultural importance, it is important to learn as much as we can about their current condition and conservation potential.

Vegetación y flora de la Ciénega de San David. En este trabajo presentaré la ciénega de San David y el aguaje Lewis del río San Pedro en

el condado de Cochise, Arizona – con énfasis en la flora. En la región de las Islas Serranas, las ciénegas son humedales escasos en ambientes

áridos donde el agua subterránea sube a la superficie de forma constante. Debido a su suelo y propiedades hidrológicas únicas, las ciénegas

mantienen un humedal graminoide. La evaporación usualmente causa que el agua sea alcalina, y la vegetación de la ciénega frecuentemente

incluye halófitas y otras especies inusuales. Y dependiendo de su edad y tamaño, también pueden albergar niveles altos de endemismo. Las

ciénegas distan mucho de ser prístinas, e igual que muchos humedales y zonas ribereñas, probablemente se han explotado por milenios. Sin

embargo, en el Suroeste Norteamericano, se asume que estos hábitats han estado en declive severo los últimos 150 años debido a un ciclo de

erosión y fragmentación por arroyos. Debido a su importancia biológica y cultural, es importante estudiar todo lo posible sobre su condición

actual y potencial de conservación.

MALUSA, JIM1, Larry Laing2, Don Falk1, and Brooke Gebow3. 1School of Natural Resources and the Environment, University of Arizona,

Tucson,AZ85721malusa@email.arizona.edu, dafalk@email.arizona.edu; 2EnvironmentalConsultant, GoldCanyon,AZ85118 desertsoilguy@

gmail.com; 3The Nature Conservancy, Southeastern Arizona Preserves, 27 Ramsey Canyon Road, Hereford AZ 85615. bgebow@tnc.org.

Mapping vegetation and landscapes utilizing a multiple-scale approach in southeastern Arizona. To advance a Forest Service standard

for ecosystem mapping at multiple scales, the USFS in 2010 issued a directive mandating the use of “The National Hierarchical Framework of

Ecological Units.” (Cleland et al.1997). Fortunately the Coronado NF and other agency partners had been moving in this direction since 2007,

when they supported a pilot project to map its lands at landscape scales, resulting in a first approximation of Landtype Associations (LTAs) of

southeastern Arizona. Criteria used in mapping LTA’s include bedrock and superficial geology, topography, elevation, potential and existing

vegetation, key soil properties, and local climatic variables. The mapping effort has benefited the Coronado’s FireScape program, in which

multiple partners utilize this mapping as a framework for fire planning. Second generation LTAmaps for the Catalina-Rincon mountains, which

are available at www.azfirescape.org, are used for managing ecological units (e.g., mixed conifer on granitic soils) typically no smaller than a

thousand acres, and often much larger. Not surprisingly, the success of the project depends on an accurate depiction of vegetative and physical

setting reality, not just interpretations of remote imagery. To that end, FireScape supports extensive on-the-ground data collection, often with

surprising revelations about places we assumed we already knew.

Trazado de mapas de vegetación y paisaje utilizando una escala múltiple en el sureste de Arizona. Para mejorar el método del Servicio

Forestal para trazar mapas de ecosistemas con escalas múltiples, en 2010 el Servicio Forestal emitió una circular exigiendo el uso del “Marco

Nacional Jerárquico de Unidades Ecológicas.” (Cleland et al. 1997). Afortunadamente el Bosque Nacional Coronado y colaboradores ya habían

empezado desde 2007, cuando apoyaron un proyecto piloto para trazar mapas de sus montes a escala del paisaje, resultando en una primera
554 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

aproximación de Asociaciones de los Tipos de Tierras (ATT,) del sureste de Arizona. Los criterios usados en el trazado de mapas ATT incluyen:

lecho rocoso y geología, topografía, altitud, vegetación actual y potencial, tipo de suelo y datos meteorológicos locales. Estos mapas son usados

en el programa FireScape delBosque NacionalCoronado y lossocios utilizan estos mapas como referencia para programarincendios. La segunda

edición de mapas ATT de la Sierra Catalinas y S. Rincón, (disponibles en www.azfirescape.org) se usan para el manejo de unidades ecológicas

(e.g., bosques mixtos de coníferas en suelos de granito) de 400 has o más grandes. Como es de esperarse el éxito del proyecto depende de una

representación exacta del entorno vegetal y físico, no solamente de las imágenes de satélite. Con este objetivo, FireScape apoya la toma de datos

de campo exhaustiva, frecuentemente con descubrimientos sorprendentes sobre lugares supuestamente conocidos.

MANN, ROY S.1, Philip Heilman,2 and Jeffry Stone3. 11301 Tamarisk Dr., West Linn, OR 97068, rsmann@teleport.com; 2USDA Agricultural

Research Service, 2000 E. Allen Rd, Tucson, AZ 85719, pheilman@tucson.ars.ag.gov; 3USDA Agricultural Research Service, 2000 E. Allen

Rd, Tucson, AZ 85719, jeff.stone@ars.usda.gov.

Potential for extending major resource areas into northern Mexico. There is a significant history of cooperative efforts between Mexico

and the United States on natural resource management issues including rangeland management. Mexico and the U.S. have jointly conducted

research and developed range management technologies. Bringing these technologies together and improving technical communications are an

ongoing process. This paper discusses a potential tool that can provide a common denominator for both countriesto more easily frame, describe,

and share data relative to rangeland resources. The objective is to present possibilities for utilizing current data and provide a vehicle that can

facilitate technical communications. Existing mapsincluding climate and elevation were used to define probable areas of Chihuahua and Sonora

that would be similar enough to Major Land Resource Areas (MLRAs) of Arizona to consider them an extension of those MLRAs. Reconnaissance surveys were made to compare soils and vegetation to those described in Arizona. Comparisons were also made between Ecological

Site Descriptions (ESDs) used in the U.S. with those developed in Mexico by COTECOCA. A preliminary map was developed that represents

probable boundaries of MLRA 41 if extended from the U.S. border into the states of Chihuahua and Sonora. Some sites were mapped to test

application of U.S. ecological site descriptions with on ground conditions in Chihuahua. The potential for extending MLRA boundaries from

the U.S. into Mexico is feasible and realistic. This would facilitate the direct use of Ecological Site Descriptions across borders and improve

exchange of rangeland data between the two countries.

Potencial de extender las áreas claves de recursos en el norte de México. México y Estados Unidos comparten una historia de colaboración

en temas del manejo de recursos naturales, incluyendo el manejo de agostaderos. México y E.U. han realizado investigaciones y desarrollado

tecnologías para el manejo de agostaderos conjuntamente. Unir estas tecnologías y mejorar la comunicación técnica son procesos en desarrollo. Este artículo presenta una herramienta potencial que puede proveer un punto en común para que ambos países puedan formular, describir

y compartir datos en relación con los recursos del agostadero más fácilmente. El objetivo es presentar las posibilidades de utilizar los datos

actuales y proveer un vehículo que pueda facilitar las comunicaciones técnicas. Los mapas existentes que incluyen el clima y la elevación se

usaron para definir zonas de Chihuahua y Sonora que probablemente serían muy similares a las Áreas Claves de Recursos Terrestres (ACRT) de

Arizona para considerarlas una extensión de tales ACRT. Se hicieron inspecciones terrestres para comparar suelo y vegetación con los descritos

en Arizona. También se hicieron comparaciones entre las Descripciones de los Sitios Ecológicos (DSE) usados en E.U. con los desarrollados en

México por COTECOCA. Se desarrolló un mapa preliminar que representa los límites probables de ACRT 41 si se extendieran de la frontera

E.U. a los estados de Sonora y Chihuahua. Algunos sitios se trazaron en una mapa para probar la aplicación de las descripciones de los sitios

ecológicos estadounidenses con las condiciones sobre el terreno en Chihuahua. El potencial de extender los límites de los ACRT de los E.U. a

México es realizable y realista. Esto facilitaría el uso directo de los Descripciones de los Sitios Ecológicos a través de la frontera y mejoraría

el intercambio de datos del agostadero entre los dos países.

MARSHALL, ROB, Marcos Robles, Dan Majka, and Dale Turner. The Nature Conservancy, 1510 E. Fort Lowell Road, Tucson, AZ 85719,

rmarshall@tnc.org, mrobles@tnc.org, dmajka@tnc.org , dturner@tnc.org.

Energy by Design: A tool for achieving no net conservation loss with transmission line and power plant siting. The Madrean region has

recently been the focus of several large energy-related development proposals, including power generating plants and long-distance electrical

transmission lines. The implementation of such projects could have significant and widespread conservation impacts, and yet environmental

considerations are often only considered late in the regulatory process when the cost of reconciling tradeoffs is high. We present an approach

called Energy by Design that was developed to minimize the impacts of energy infrastructure by integrating the best available science early into

regulatory and planning processes. The premise of the energy by design framework is to make tradeoffs explicit and identify a range of options

that would help achieve a standard of no net loss of conservation values. The standard is met through use of the mitigation hierarchy where

options are identified to avoid, minimize, restore, and offset major project impacts to habitat and species at large landscape and local scales. We

will use the SunZia Southwest Transmission Project in Southeastern Arizona as an example of the application of Energy by Design principles.

Energía por Diseño: Cero pérdida de la conservación con el emplazamiento del cableado y plantas generadoras. La región Madrense

recientemente ha sido el foco de varias propuestas de desarrollo relacionadas con la energía, incluyendo las plantas generadoras de electricidad

y cableados de larga distancia. La implementación de tales proyectos podría tener impactos considerables y extensos en la conservación, aún

así las consideraciones medioambientales frecuentemente sólo se consideran tarde en el proceso normativo cuando el costo de reconciliar los

compromisos es alto. Presentamos el enfoque Energía por Diseño que se desarrolló para minimizar los impactos de la infraestructura al integrar

la mejor ciencia disponible a los procesos normativos y de planificación. La premisa del marco de la Energía por Diseño es hacer explícitos
USDA Forest Service Proceedings RMRS-P-67. 2013 555

Abstracts/ Resúmenes

los compromisos e identificar una gama de opciones que ayudarían a lograr un estándar de cero pérdida neta de los valores de conservación. El

estándar se alcanza por el uso de la jerarquía de mitigación en la cual las opciones se identifican para evitar, minimizar, restablecer y compensar

los impactos importantes al hábitat y las especies en escalas grandes de paisaje y locales. Utilizaremos el Proyecto de Transmisión Suroeste

SunZia en el sureste de Arizona como ejemplo de la aplicación de los principios de Energía por Diseño.

MCCOY1,A. L., C.A. Zugmeyer2, C. L. McIntyre2, E. M. Brott2, and B. J. Orr3. 1Ecosystem Economics, Tucson,AZ; 2Sonoran Institute, Tucson

Arizona; 3Arizona Remote Sensing Center & Office of Arid Lands Studies, University of Arizona, Tucson, AZ.

Impacts of effluent subsidies on riparian resilience. In 2005, a significant and sudden vegetation die-off event occurred in the riparian corridor of the Upper Santa Cruz River in southern Arizona. Analysis of past and current vegetation along the river suggests that this event can be

linked to two competing factors relating to the discharge of treated municipal effluent into the river. Examination of records of riparian vegetation indicates an increase in the extent and density of vegetation in the 1980s and 1990s; this increase coincided with the presence of effluent

subsidies and favorable climate conditions that encouraged growth. However, while water quantities in the river supported vigorous vegetative

growth, poor effluent water quality was slowly hindering hydrologic functions. Our analysis of the riparian ecosystem suggests that the 2005

event was partially due to the eventual extreme degradation of the river’s hydrologic functions coupled with drought conditions of the 2000s.

The die-off, along with the results of our analysis, highlight the uncertainties and unknowns of effluent use in the Upper Santa Cruz River and

further highlight the need to develop scientific and legal mechanisms to ensure that effluent bolsters, rather than degrades, riparian function and

the provision of riparian ecosystem services in the semi-arid southwestern United States.

Impactos de aguas residuales en la resiliencia de hábitats ribereños. Una mortandad generalizada de plantas ocurrió en 2005 en el corredor

ribereño del alto río Santa Cruz en el sureste de Arizona. Análisis de la vegetación histórica y actual del río sugiere que este evento puede vincularse a dos factores relacionados con descarga de aguas residuales municipales tratadas al río. Los registros de vegetación ribereña indican un

aumento de la extensión y densidad de la vegetación en la década de 1980 y 1990, este aumento coincidió con la presencia de descarga de aguas

residuales y un clima favorable que fomentaron el crecimiento. Sin embargo, mientras las cantidades de agua en el río ayudaron al crecimiento

vegetal vigoroso,lamala calidad de aguasresiduales gradualmente entorpeció lasfunciones hidrológicas.Nuestro análisis del ecosistema ribereño

sugiere que la mortandad de 2005 se debió en parte a la degradación extrema de las funciones hidrológicas del río, aunado a las condiciones de

sequía de la década de 2000. La mortandad, junto con los resultados de nuestro análisis, resalta las incertidumbres y las incógnitas del uso de

aguas residuales en el alto río Santa Cruz y enfatiza la necesidad de desarrollar mecanismos científicos y legales para asegurar que se refuerce,

en vez de degradar, el funcionamiento ribereño y suministro de los servicios ecológicos ribereños en el suroeste semiárido de Estados Unidos.

MERRICK, MELISSA1,John L. Koprowski2, and Craig Wilcox3. 1The University ofArizona School of Natural Resources and the Environment,

1311 E. 4th St., Tucson, AZ 85721, mmerrick@email.arizona.edu; 2The University of Arizona School of Natural Resources and the Environment, 1311 E. 4th St., Tucson, AZ 85721, squirrel@ag.arizona.edu; 3

USDA Forest Service, Coronado National Forest, 711 S. 14th Ave., Suite

D, Safford, AZ 85546, cpwilcox@fs.fed.

Into the third dimension: Benefits of incorporating LiDAR data in wildlife habitat models. LiDAR (Light detection and ranging) is an

increasingly useful tool for characterizing wildlife habitat, providing detailed, three-dimensional landscape information not available from other

remote sensing applications. The ability to accurately map structural components such as canopy height, cover, woody debris, tree density,

and ground surface can greatly improve wildlife habitat models as animals interact and respond to three-dimensional habitat features. Prior

to LiDAR, accurate measurements of structural features were difficult to obtain over large areas and other remotely sensed data is based on

two-dimensional spectral responses. The southwest harbors an incredible diversity of unique vegetation communities, each with an associated

wildlife assemblage with various management needs. LiDAR is a unique tool, managers can use to accurately characterize vegetation structural

characteristics for entire districts or management units. Data surfaces derived from the LiDAR point cloud can be readily incorporated into

species-specific or multispecies habitat models.Although LiDAR hasreceived much attention in characterizing foreststructure, few studies(n =

28) have suggested orincorporated thistechnology to improve wildlife habitatmodels. Herein we provide a review of currentLiDARapplications

in wildlife habitat models, provide future directions, and detail how LiDAR can increase our ability to represent the world’s animal experience.

A la tercera dimensión: Beneficios de incorporar los datos de LiDAR en los modelos del hábitat de fauna. El LiDAR (Light Detection and

Ranging; detección y medición a través de la luz) es una herramienta cada vez más útil para evaluar el hábitat de fauna, aportando información

tridimensional detallada que no se puede obtener con otros sistemas de detección remota. La capacidad de trazar en mapas con exactitud los

componentes estructurales como la altura del dosel, la cobertura, el detrito leñoso, la densidad arbórea y la superficie del suelo pueden mejorar

los modelos de hábitat ya que los animales interactúan y responden a las características tridimensionales del hábitat. Antes del LiDAR, las

mediciones exactas de las características estructurales eran difíciles de obtener en áreas grandes y otros datos de sensores remotos se basan

en las respuestas espectrales bidimensionales. El suroeste norteamericano alberga una diversidad increíble de comunidades vegetales únicas,

cada uno con ensambles de fauna asociada con varias necesidades de manejo. El LiDAR es una herramienta que los encargados de manejo

pueden usar para evaluar con exactitud las características estructurales de la vegetación por regiones completas o unidades de manejo. Los

datos derivados de LiDAR pueden incorporarse fácilmente a modelos de hábitat para una o varias especies. Aunque el LiDAR se conoce como

una herramienta para definir la estructura forestal, pocos estudios (n = 28) han utilizado esta tecnología para mejorar los modelos del hábitat

de fauna. Se presenta una síntesis de las aplicaciones actuales del LiDAR en modelos de hábitat faunísticos, señalamos futuros usos y cómo el

LiDAR puede aumentar nuestra capacidad de representar la experiencia del mundo animal.
556 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

MEYER, WALLACE M., III1,2,Jeff Eble1,2, Kimberly Franklin1,2,3, Paul E Marek1,2, Garret B Hughes1, Richard C Brusca3, and Wendy Moore1,2,3. 1

Department of Entomology, University of Arizona, 1140 E. South Campus Dr., Forbes 410, Tucson, AZ 85721-0036; 2Center for Insect Science, University of Arizona, 1007 E. Lowell St., P.O. Box 210106, Tucson, AZ 85721-0106; 3

Arizona-Sonora Desert Museum, 2021 N. Kinney

Road, Tucson, AZ 85743

Do plant biomes along the elevation gradient in the Santa Catalina Mountains harbor unique arthropod communities? The southwestern sky islands represent one of the most diverse regions in North America for arthropods. However, land managers are ill equipped to make

informed decisions concerning the preservation of arthropods or predict how arthropods will respond to climate change because fundamental

questions about species distributions and diversity remain unanswered. The lack of data for most arthropod groups means that management

decisions are often based on extrapolations from the few taxa studied, despite it being unclear if such patterns can be generalized across taxa.

To address these shortcomings we are examining whether patterns of community structure and diversity among seven evolutionarily distinct

ground-dwelling arthropod groups (ants, beetles, soil mites, grasshoppers, millipedes, isopods, scorpions/ pseudoscorpions) covary in the Santa

Catalina Mountains. Collections were made using pitfall traps at 66 sites that, taken together, span the elevational gradient of the Catalinas,

ranging from desertscrub, chaparral, and grasslands at low elevations to oak, pine, and mixed conifer forests at higher elevations. Preliminary

data on ant and beetle assemblages indicate that different plant biomes harbor unique communities. In addition to providing valuable baseline

data, this study will help identify vulnerable high elevation arthropod species that could be “pushed off” the mountaintop as climates warm.

Los biomas vegetales del gradiente altitudinal de la Sierra Santa Catalina, ¿albergan comunidades únicas de artrópodos? Las Islas

Serranas del suroeste norteamericano representan una de las regiones con fauna de artrópodos más diversas de Norteamérica. Sin embargo, los

encargados de manejo no están preparados para hacer decisiones acerca de la conservación de artrópodos o predecir cómo responderán al cambio climático ya que su distribución y diversidad se desconocen. La falta de datos para la mayoría de los grupos de artrópodos significa que las

decisiones administrativasfrecuentemente se basan en la extrapolación de los pocostaxones estudiados, aunque se desconoce si tales patronesse

pueden generalizar entre taxones. Para encontrar una solución estamos investigando si los patrones de estructura y diversidad de la comunidad,

entre siete grupos evolutivamente distintos de artrópodos epiedáficos(hormigas, escarabajos, ácaros delsuelo, chapulines, miriápodos, isópodos,

alacranes /seudo-alacranes) presentan covarianza en la Sierra Santa Catalina. Las colectas se hicieron usando trampas de caída en 66 sitios a

lo largo de un gradiente altitudinal en las Catalinas, cubriendo desde matorral desértico, chaparral y pastizal en las elevaciones bajas, hasta los

bosques de encino, pino, y coníferas mixtas en elevaciones altas. Los datos preliminares de los ensambles de hormigas y escarabajos indican

que los diferentes biomas vegetales albergan comunidades únicas. Además de aportar datos de referencia valiosos, este estudio servirá para

identificar las especies de artrópodos vulnerables en elevaciones altas que podrían ser “empujadas” de las cumbres cuando el clima se caliente.

MIMS, MERYL C.1, Julian D. Olden1, Martin A. Schlaepfer2,3, Ivan C. Phillipsen4, Dave A. Lytle4. 1University of Washington, School of

Aquatic and Fishery Sciences, 1122 Northeast Boat Street, Seattle, WA 98105, mmims@uw.edu, olden@uw.edu; 2State University of New

York, College of Environmental Science and Forestry, 1 Forestry Drive, Syracuse, NY 13210, mschlaepfer@esf.edu; 3INRA, Ecologie et Santé

des Ecosystèmes, 35042 Rennes, France; 4Oregon State University, Department of Zoology, 501 Life Sciences West, Corvallis, OR 97331,

philliiv@science.oregonstate.edu, lytleda@oregonstate.edu.

Population structure and the landscape genetics of a common desert anuran. Desert streams play a prominent role in shaping hydrological, biogeochemical, and ecological structure of arid and semi-arid ecosystems. Many anurans in the Madrean Archipelago rely on streamflows

generated by winter storms and summer monsoons that provide favorable breeding habitat and hydrologic connectivity with individuals or

populationsthat are otherwise isolated by harsh and dry habitat. The predictability, frequency, and magnitude of these flows will likely be altered

by climate change and increasing human requirements for freshwater. Examining population structure of desert anurans - and its relationship

to hydrology - is an important step in understanding how these species will be affected by climate change and land use. We present results of

a landscape genetic analysis of one common desert anuran of the Madrean Archipelago, Spea multiplicata (Mexican spadefoot). Individuals

were collected during the summers of 2010 and 2011. We first examine population structure throughout the Huachuca, Santa Rita, and Dragoon

Mountain ranges. We then examine the relationship of population structure to major landscape features, including hydrology. These results are

the first part of a comparative study examining the role of hydrology in population structure of multiple anuran species found throughout the

Sky Islands.

Estructura de población y genética del paisaje de un anuro desértico común. Los arroyos del desierto tienen un papel fundamental en la

estructura hidrológica, biogeoquímica y ecológica de los ecosistemas áridos y semiáridos. Muchos anuros delArchipiélago Madrense dependen

de caudales generados por las tormentas invernales y el monzón del verano que generan hábitat favorable para la reproducción y la conectividad

hidrológica con individuos o poblaciones, de otra forma aislados por un entorno hostil y seco. La predecibilidad, frecuencia y magnitud de estas

corrientes probablemente serán alteradas por el cambio climático y el aumento del requisito humano de agua dulce. Investigar la estructura de la

población de los anuros desérticos, y su relación a la hidrología, es un paso importante para entender cómo estas especies serán afectadas por el

cambio climático y el uso del suelo. Presentamos el análisis de la genética de paisaje de un anuro desértico común del Archipiélago Madrense,

Spea multiplicata. Se colectaron individuos durante los veranos de 2010 y 2011. Primero examinamos la estructura de la población en las cordilleras Huachucha, Santa Rita y Dragoon. Después examinamos la relación entre la estructura de la población y los factores importantes del

paisaje, incluyendo la hidrología. Estos resultados son la primera parte de un estudio comparativo que examina el papel de la hidrología en la

estructura de población de varias especies de anuros de las Islas Serranas.
USDA Forest Service Proceedings RMRS-P-67. 2013 557

Abstracts/ Resúmenes

MINCKLEY, CHUCK1, Doug Duncan2, Ross Timmons3, Izar Izaguirre-Pompa4, Dennis Caldwell5, Phil Rosen6, and Jaime López-Méndez7. 1

Cuenca de los Ojos. cminck@npgcable.com; 2U.S. Fish and Wildlife Service, Tucson,AZ 85745, Doug_Duncan@fws.gov; 3Arizona Game and

Fish Department, Phoenix, AZ, 85086, rtimmons@azgfd.gov; 4Reserva de la Biosfera del Pinacate y Gran Desierto de Altar, Puerto Peñasco,

Sonora, iizaguirre@conanp.gob.mx; 5Caldwell Design, Tucson, AZ 85705, dennis@caldwell-design.com; 6University of Arizona, Tucson, AZ

85720, pcrosen@u.Arizona.edu; 7Colegio de Bachilleres del Estado de Sonora, Sonoyta, Sonora. jflm_77@hotmail.com.

Conservation and management of native aquatic species of the Río Sonoyta, Sonora México.The Río Sonoyta is an important aquatic

ecosystem in northern Sonora, which is disappearing because of drought and groundwater withdrawal. Its native species are also threatened

by introduced species. The only watered reach is an intermittent segment (<1 km, Agua Dulce), found just across the Southern International

Border from Organ Pipe Cactus National Monument (OPCNM). The native fish present in the river include the endangered Sonoyta pupfish

(Cyprinodon eremus) and the indigenous longfin dace (Agosia chrysogaster). In the wild, the pupfish occurs only at Quitobaquito Springs on

OPNCM and at Agua Dulce. The longfin dace may be extirpated. A partnership formed and led by the La Reserva de la Biosfera El Pinacate y

Gran Desierto deAltar included partnersfrom the United States and México to create fish refugesin México. We summarize conservation efforts

to maintain native fishes in refuge ponds, report on their status, and offer suggestions for future management. We also present information on

the future establishment of refuges for longfin dace and augmentation of pupfish refuges in the United States and México. We briefly discuss

the impact of a new wastewater treatment plant on the Sonoyta River.

Conservación y manejo de especies acuáticas nativas del río Sonoyta, Sonora, México. El río Sonoyta es un importante ecosistema acuático

en el norte de Sonora que está desapareciendo debido a la sequía y extracciones del acuífero. Sus especies nativastambién están amenazadas por

las especies introducidas. El único tramo con agua es un segmento intermitente (<1 km, Agua Dulce), justo al sur de la frontera internacional

de Organ Pipe Cactus National Monument (OPCNM). Los peces nativos presentes en el río son: el pupo del río Sonoyta (Cyprinodon eremus)

especie en peligro de extinción, y el charalito de aleta larga (Agosia chrysogaster) una especie nativa. El pupo del río Sonoyta se encuentra

silvestre sólo en Quitobaquito Springs en OPCNM y en Agua Dulce. El charalito de aleta larga probablemente ha sido extirpado. La Reserva

de la Biosfera El Pinacate y Gran Desierto de Altar formó y está a cargo de una asociación con miembros tanto de E.U. como de México para

crear refugios para peces en México.Se presenta un resumen de los trabajos de conservación para mantener los peces nativos en las tinajas

del refugio, así como un reporte de su estatus y sugerencias para el manejo futuro. También se presenta información sobre el establecimiento

de refugios para el charalito de aleta larga y un aumento de refugios para el pupo del río Sonoyta tanto en E.U. como México. Por último, se

discute brevemente el impacto de una nueva planta tratadora de aguas residuales en el río Sonoyta.

MINCKLEY, CHUCK. Flagstaff, AZ; Cuenca de los Ojos, cminck@npgcable.

Seasonal distribution of native fishes in a 10-kilometer reach of San Bernardino Creek, Sonora, México. San Bernardino Creek is a northern tributary of the Yaqui River. Originating in the United States it crosses the Southern International Border just east of Douglas, AZ/Agua

Prieta, Sonora and immediately south of the San Benardino/Leslie Canyon National Wildlife Refuge. Six of the eight Yaqui river fish species

are found in this reach and include four minnows, a sucker, and a poeciliid. Information is presented on the annual and seasonal distribution of

these fishes during 2010 - 2011. The Yaqui chub population is discussed in more detail including a population estimate of this species. A brief

history of the area is given and the use of gabion construction to change this stream during the last 3 decades is presented.

Distribución estacional de peces nativos en un tramo de 10 km del arroyo San Bernardino, Sonora, México. El arroyo San Bernardino

es un afluente norteño del río Yaqui. Nace en Estados Unidos y cruza la frontera internacional justo al este de Douglas, Arizona y Agua Prieta,

Sonora y directamente al sur de los Refugios Nacionales de Vida Silvestre San Bernardino y Leslie Canyon. En este tramo se encuentran seis

de las ocho especies de peces del río Yaqui: cuatro carpitas, el matalote y un poeciliido. Se presenta información en la distribución anual y

estacional de estos peces durante 2010-2011. Se hace una estimación de la población de la carpa púrpura. Y se incluye una breve historia del

área y la construcción de presas de gaviones para mejorar este afluente.

MINCKLEY, ROBERT L. Department of Biology, University of Rochester, Rochester, NY, robert.minckley@rochester.edu.

Changes of riparian and desert vegetation after cattle removal in the San Bernardino Valley, Arizona/ México. The San Bernardino

Valley of Arizona, USA, and Sonora, México, is one of the principal headwaters of the Río Yaqui. Conservation efforts centered on the unique

aquatic organisms and habitats found in this river system have resulted in a landscape-scale experiment. Cattle were removed from the San

Bernardino National Wildlife Refuge, Arizona, in 1979 and from the Rancho San Bernardino, México, in 2000. These adjoining areas experience the same yearly climatic variation and have similar soils and potential flora, yet differ in time since grazing. Permanent plots established

on 2000 and 2004 show vegetation differences are greater among mesic habitats in recently grazed and long-term ungrazed plots than in xeric

habitats. Plant species richness in xeric habitats (desertscrub and grassland) is greater than it is mesic habitats (riparian and mesquite bosque),

but there is little difference in richness among xeric habitats in recently grazed and long-term ungrazed areas. It is unclear if this pattern is the

result of minimal grazing away from watersources or reduced succession rate. Speciesrichnessin riparian habitats declinesin time with grazing

increases. Given that the riparian area was desert marsh in the last century, ongoing restoration efforts should continue to result in fewer plant

species as succession proceeds in habitats with permanent water.
558 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Cambios en vegetación desértica y ribereña después del desalojo de ganado en el valle de San Bernardino en Arizona y México. El

valle de San Bernardino de Arizona, E.U. y Sonora, México, es uno de los principales afluentes del río Yaqui. Los trabajos de conservación de

hábitats y organismos acuáticos exclusivos de este río han resultado en un estudio comparativo de diversidad florística. El ganado se desalojó

de San Bernardino National Wildlife Refuge, Arizona en 1979 y del Rancho San Bernardino, México en 2000. Estas áreas colindantes tienen

la misma variación climática anual, así como suelos similares y flora potencial, pero difieren en el tiempo que tienen sin ganado. Las parcelas

permanentes establecidas en 2000 y 2004 muestran que las diferencias de vegetación son mayores entre hábitats húmedos con pastoreo reciente

y parcelas con largo tiempo sin pastorear, que en hábitats áridos. La diversidad de la flora de hábitats áridos (matorral desértico y pastizal) es

mayor que en los hábitats húmedos(ribereño y bosque de mesquite). Sin embargo, hay muy poca diferencia entre la diversidad de hábitats áridos

con pastoreo reciente y áreas áridas con mucho tiempo sin pastorear. Basado en los resultados, no está claro si este patrón es el resultado de un

pastoreo mínimo lejos del agua o de un rango de sucesión reducido. La diversidad de especies en hábitats ribereños disminuye con el tiempo

conforme aumenta el pastoreo. Dado que el área ribereña fue un humedal del desierto el siglo pasado, la restauración continuará produciendo

menos especies de plantas conforme la sucesión continua en los a hábitats con agua permanente.

MINCKLEY, ROBERT L.1 and John Ascher2. 1Department of Biology, University of Rochester, Rochester, NY, robert.minckley@rochester.

edu; 2Department of Entomology, American Museum of Natural History, New York, NY.

Regional diversity of bees in northeastern Sonora, México and southeastern Arizona, USA. Museum records have long indicated that the

xeric areas of North America are unusually rich for bees (Hymenoptera: Apiformes). Two areas of the Sky Islands where bees have been most

intensively collected are the Chiricahua Mountains,Arizona, and the San Bernardino Valley,Arizona/ Sonora. Opportunistic bee sampling from

the Chiricahua Mountains date back decades and have spanned from desert to pine forest ecosystems. Intensive standardized bee sampling in

the San Bernardino Valley began recently (2000) and has been limited to a 20 km2 area of Chihuahuan desertscrub. Species density in this area

is greater than that documented elsewhere in the world; ca. 1/10th of the total bee species in the United States occur here. Species diversity is

elevated in part because of high spatial and temporal turnover of species. Despite the proximity and near continuous distribution of desertscrub

habitat between these two areas, bee species composition changes considerably and differences within this desert ecosystem are comparable to

those thatoccur amongecosystems athigher elevations.Ourpreliminary analysisindicatesregionalprocessessuchasthe intersection of ecoregions

strongly influence bee species diversity and that topographic complexity and differences in the spring and late summer fauna are also involved.

Diversidad regional de abejas en el noreste de Sonora, México y sureste de Arizona, E.U. Los registros de museos han indicado repetidamente que las áreas secas de América del Norte tienen una diversidad de abejas (Hymenoptera: Apiformes) excepcional. La Sierra Chiricahua

en Arizona y el valle de San Bernardino en Arizona y Sonora son dos áreas de las Islas Serranas donde las abejas se han colectado de forma

intensa. El muestreo oportunístico de abejas en la Sierra Chiricahua se ha realizado durante varias décadas y abarcado desde ecosistemas de

desierto a los de bosque de pino. Un muestreo de abejas, intensivo y estándar, en el valle de San Bernardino empezó recientemente (2000) y se

ha restringido a un área de 20 km2 de matorral del desierto Chihuahuense. La densidad de especies en esta área es mayor que la documentada

en otras partes del mundo; alrededor de un 1/10 del total de especies de abejas de Estados Unidos ocurre aquí. La diversidad de especies es

elevada debido en parte a un alto movimiento de especies temporal y espacial. A pesar de la cercanía y la casi continua distribución de hábitat

de matorral desértico entre estas dos áreas, la composición de especies de abejas cambia considerablemente y las diferencias dentro de este

ecosistema de desierto son comparables a las que ocurren entre ecosistemas de elevaciones más altas. Nuestros análisis preliminaresindican que

los procesos regionales tales como la intersección de ecoregiones tienen una fuerte influencia en la diversidad de especies de abejas y también

intervienen la complejidad topográfica y las diferencias entre la fauna de primavera y de finales del verano.

MINCKLEY, THOMAS A.1, A. Brunelle2, D. Turner3. 1Department of Botany, Program in Ecology, University of Wyoming; 2Department of

Geography, University of Utah; 3The Nature Conservancy, Tucson, AZ.

Paleoenvironmental perspective for understanding the development, stability and state-changes of ciénegas in the American Deserts. As

persistent wetlands in arid regions, ciénegas represent important resources for the maintenance and preservation of regional biodiversity. The

history of ciénegas in the American Southwest over the last 8,000 years provides information on the dynamics of growth, longevity, and stability of these habitats under previous climate conditions. Proxy data such as sedimentology, pollen, charcoal, and isotopes preserved in ciénega

sediments provide information on the formation, disturbance, resilience and state changes within these systems. This long-term perspective is

compared to the recent history of degradation observed in the region. We present information on how the paleoenvironmental record of change

can be used in conservation, restoration and management of ciénegas.

Perspectiva paleo-ambiental para entender el desarrollo, estabilidad y cambios en las ciénegas de los desiertos americanos. Por ser

humedales persistentes en regiones áridas, las ciénegas representan recursos importantes para mantener y preservar la biodiversidad regional.

La historia de las ciénegas en el suroeste americano durante los últimos 8000 años aporta información sobre la dinámica de crecimiento, longevidad y estabilidad de estos hábitats bajo las condiciones de climas del pasado. Datos indirectos como sedimentología, polen, carbón vegetal

e isótopos preservados en los sedimentos de la ciénega proporcionan información sobre la formación, perturbación, resistencia y cambios en

estos sistemas. Esta perspectiva a largo plazo es comparada con la historia reciente de degradación en la región. Se presenta información de

como el registro paleo ambiental de cambios puede utilizarse en la conservación, restauración y manejo de las ciénegas.
USDA Forest Service Proceedings RMRS-P-67. 2013 559

Abstracts/ Resúmenes

MISZTAL, LOUISE1, G. Garfin2, and L. Hansen. 1Sky Island Alliance, 300 E University Blvd., Suite 270, Tucson, AZ 85705, louise@skyislandalliance.org; 2

School of Natural Resources and the Environment, University of Arizona, 845 N. Park Ave., Suite 532, Tucson, AZ 85721,

gmgarfin@email.arizona.edu; 3EcoAdapt, P.O. Box 11195, Bainbridge Island, WA 981103, lara@ecoadapt.org.

Responding to climate change impacts in the Sky Island Region: From planning to action. Addressing the growing impacts of climate

change on natural resources requires multiple organizations, agencies, and institutions working cooperatively to incorporate climate change

into resource management. In the Sky Island region of the southwestern United States and northern Mexico, Sky Island Alliance, a non-governmental organization, is leading the convening of a series of climate change adaptation workshops in cooperation with a variety of agencies

and organizations. This paper demonstrates a process and methodology for convening federal and state agencies, local governments, non-profit

organizations, tribal representatives, private landowners, and academic researchers in order to develop, on-the-ground and policy-level actions

through climate change adaptation planning. Key outcomes of the workshops include: identification of climate change threats to and vulnerabilities of Madrean Forest, Riparian, Desert and Grassland ecosystems in the Sky Island region; analysis of direct and indirect climate change

threats and interacting factors; a list of ecosystem specific adaptation options for the region, a plan for implementation of one adaptation strategy and development of a regional network of professionals working cooperatively to improve natural resource management under changing

conditions. This paper highlights one approach for addressing the management and conservation challenges posed by climate change through

collaborative engagement at a regional scale.

Respondiendo al impacto del cambio climático en la Región de las Islas Serranas: De la planificación a la acción. Enfrentar los crecientes

impactos del cambio climático requiere múltiples organizaciones, agencias, e instituciones trabajando cooperativamente para incorporar el

cambio climático al manejo de recursos. En la región de las Islas Serranas en el suroeste de Estados Unidos y el norte de México, Sky Island

Alliance, una organización no gubernamental, desarrolla una serie de talleres de adaptación al cambio climático, en colaboración con varias

agencias y organizaciones. Este trabajo explica el proceso y metodología para convocar agencias federales y estatales, gobiernos locales, organizaciones sin fines de lucro, representantes tribales, propietarios privados, e investigadores académicos a fin de desarrollar acciones prácticas

y normativas para la adaptación al cambio climático. Los resultados principales de los talleres son: identificación de las amenazas del cambio

climático y la vulnerabilidad de los ecosistemas del bosque madrense, ripario, desértico y de pastizal en la región de las Islas Serranas; análisis

de las amenazas directas e indirectas del cambio climático y los factores que interactúan; listado de opciones de adaptación especificas del ecosistema para la región; plan para la implementación de una estrategia de adaptación y desarrollo de una red regional de profesionales trabajando

cooperativamente para mejorar la gestión de los recursos naturales en condiciones cambiantes. Este documento pone de relieve un enfoque para

abordar los desafíos de gestión y conservación que plantea el cambio climático a través de un compromiso de colaboración a escala regional.

MORENO-ARZATE, EFREN1,2, Carlos A. López-Gonzalez1, and Gerardo Carreón-Arroyo2. 1Universidad Autónoma de Querétaro, Cerro de

las Campanas S/N, Col. Las Campanas, Querétaro, Querétaro, C. P. 76010, arzate2786@yahoo.com.mx, cats4mex@aol.com; 2Naturalia A. C.,

El Cajón #9, Hermosillo, Sonora.

Population status of prairie dogs (Cynomys ludocivianus) in the San Pedro River Basin, Sonora. The black tailed prairie dog (Cynomys 

ludovicianus) is a species of interest for Mexico, the United States and Canada. Populations in Mexico are considered at risk, including those

in Sonora, such populations require additional conservation efforts to maintain them long term. Our objective was to determine population size

and density of the Sonoran colonies to establish a new colony. The prairie dog colonies are 92.32 ha for La Mesa and 57.86 ha for Las Palmitas

respectively. Density was calculated using visual transect counts, and Distance sampling software for the analysis of the data. La Mesa was

surveyed between November 2010 and November 2011, and las Palmitas from May to November 2011. Our density estimates range from 1.03

to 5.29 ind/ha with an average density of 3.09 ind/ha, and an estimated population of 285.26 individuals. In the case of Las Palmitas densities

ranged from 1.50 to 6.45 ind/ha with an average density of 3.62 ind/ha and an estimated population of 209.45 individuals. Our results support

a controlled but limited translocation effort that will allow the establishment of a new colony at Los Fresnos private reserve, consequently

increasing the resilience and survivorship of black-tailed prairie dogs in Sonora.

Estado de la población de perritos de la pradera (Cynomys ludocivianus) en la cuenca del río San Pedro, Sonora. El perrito de las praderas

de cola negra (Cynomys ludovicianus) es una especie de interés para México, Estados Unidos y Canadá. Poblaciones en México se consideran

en situación de riesgo, incluidas las de Sonora, estas poblaciones requieren esfuerzos de conservación adicionales para mantenerlas a largo

plazo. Nuestro objetivo fue determinar el tamaño y la densidad de las poblaciones de las colonias de Sonora para establecer una nueva colonia.

Las colonias de perritos de la pradera son 92.32 hectáreas en La Mesa y 57.86 ha en Las Palmitas, respectivamente. La densidad fue calculada

utilizando conteos visuales en transectos, y se utilizó software de muestreo a distancia para el análisis de los datos. La Mesa se monitoreó entre

noviembre de 2010 y noviembre de 2011, y Las Palmitas de mayo a noviembre de 2011. Nuestras aproximaciones de densidad están en el rango

de 1.03 a 5.29 ind / ha con una densidad media de 3.09 ind / ha y una población estimada de 285.26 individuos. En el caso de Las Palmitas

densidades variaron desde 1.50 hasta 6.45 ind / ha con una densidad media de 3.62 ind / ha y una población estimada de 209.45 individuos.

Nuestros resultados apoyan un traslado controlado pero limitado, que permitirá la creación de una nueva colonia en la reserva privada de Los

Fresnos. Y así aumentar la resiliencia y la supervivencia del perrito de las praderas de cola negra en Sonora.
560 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

MORGAN, AVRA O.1, Leslie Meyers2, Genevieve Johnson2, and Christina Vojta3. 1U.S. Bureau of Reclamation, Denver Federal Center Building 67, Denver, CO 80225; 2 U.S. Bureau of Reclamation, 6150 W. Thunderbird Rd., Glendale, AZ 85306; 3

U.S. Fish and Wildlife Service, 323

N. Leroux Suite 201, Flagstaff, AZ 86001

The Desert Landscape Conservation Cooperative: Linking science to management.The DesertLandscapeConservationCooperative (LCC)

is a self-directed, non-regulatory partnership that covers the Mojave, Sonoran, and Chihuahuan Deserts. The Bureau of Reclamation (Reclamation) provides overall coordination of the Desert LCC, and the Fish and Wildlife Service provides a permanent Science Coordinator to provide

coordination and synthesis of science activities. The Steering Committee consists of 26 members from state agencies, tribes, non-governmental

organizations, and federal agencies in the U.S. and Mexico. The primary mission of the Desert LCC is linking science to management to address

climate change and other broad scale stressors. The Madrean Archipelago is particularly vulnerable to climate change impacts. The Desert LCC

(through Reclamation) has funded the Sky Islands Alliance to conduct a survey of springs and seeps in that area in relation to climate change.

Other projects that are currently underway are analyses of climate change on major river systems (as part of Reclamation’s West-Wide Climate

Risk Assessments), a climate change vulnerability assessment of plant communities in the Mojave and Sonoran Deserts (as part of Fish and

Wildlife Service collaboration with NatureServe), mapping of surface waters in the Sonoran Desert (U.S. Geological Survey and University of

Arizona), and 8 other projects recently funded by Reclamation’s WaterSMART program.

Cooperativa de conservación de paisajes del desierto: uniendo la ciencia con el manejo. La cooperativa de conservación de paisajes del

desierto (LCC porsussiglas en inglés) es una asociación independiente, no normativa, que cubre los desiertos Mojave, Sonorense y Chihuahuense. La Oficina de Rehabilitación ofrece la coordinación de la LLC del desierto y el Servicio de Pesca y Vida Silvestre ofrece un coordinador

científico permanente y síntesis de actividades científicas. El comité directivo consiste de 26 miembros de agencias estatales, tribus, organismos

no gubernamentales y agenciasfederales en E.U. y México. El objetivo principal de la LLC del desierto es conectar la ciencia con el manejo para

abordar el cambio climático y otros eventos con impacto a gran escala. El Archipiélago Madrense es particularmente vulnerable al impacto del

cambio climático. La LLC del desierto (mediante la Oficina de Rehabilitación) ha financiado a Sky Islands Alliance para un estudio de manantiales en el área y efectos del cambio climático. Otros proyectos en curso son el análisis del impacto del cambio climático en sistemas fluviales

principales (como parte de la evaluación del riesgo climático en el oeste por parte de la Oficina de Rehabilitación), evaluación de la vulnerabilidad de comunidades vegetales al cambio climático en los desiertos Mojave y Sonorense (como parte de una colaboración entre el Servicio de

Pesca y Vida Silvestre y NatureServe), trazado de mapas de aguas superficiales en el Desierto Sonorense (University of Arizona y el Servicio

Geológico de Estados Unidos) y otros 8 proyectos recientemente financiados por el programa WaterSMART de la Oficina de Rehabilitación.

MOORE, WENDY1,2,3 (wmoore@email.arizona.edu), Wallace M Meyer III1,2, Jeff Eble1,2, Kimberly Franklin1,3, John F. Wiens3, and Richard

C Brusca2,3. 1Department of Entomology, University of Arizona, 1140 E. South Campus Dr., Forbes 410, Tucson, AZ 85721-0036; 2Center for

Insect Science, University of Arizona, 1007 E. Lowell St., P.O. Box 210106, Tucson, AZ 85721-0106; 3Arizona-Sonora Desert Museum, 2021

N. Kinney Rd., Tucson, AZ 85743.

The Arizona Sky Island Arthropod Project (ASAP): systematics, biogeography, ecology, and population genetics of ground-dwelling 

arthropods. The Arizona Sky Island Arthropod Project (ASAP) is a new research program that combines systematics, biogeography, ecology,

and population geneticsto study ground-dwelling arthropod diversity, abundance, distribution and community structure along elevation gradients

and among mountain ranges in the desert Southwest. Ground-dwelling arthropods represent taxonomically and ecologically diverse organisms

that drive key ecosystem processes. Using data from museum specimens and material collected from long-term pitfall trap monitoring, ASAP

will document diversity patterns across the Sky Islands to identify patterns of endemism, phylogeography, and the evolutionary origins of arthropods in these communities. In addition, ASAP will utilize the natural laboratory provided by the Sky Islands to evaluate the ecological roles

of arthropods across elevational and environmental gradients on the mountains. Baseline data will be used to determine climatic boundaries for

target species, which will then be integrated with climatological models to predict future changes in arthropod communities and distributions

in the wake of rapid climate change. Here, we introduce this project, outline its overarching goals, and present historical data on Sky Island

ground-dwelling arthropods harvested from the University of Arizona Insect Collection. In talks following this one, collaborators will present

analyses of data collected during the first year of ASAP on various aspects of this multidisciplinary project.

Proyecto de Artrópodos de las Islas Serranas de Arizona (ASAP): Sistemática, biogeografía, ecología y genética de poblaciones de artrópodos terrestres. El Proyecto de Artrópodos de Las Islas Serranas de Arizona (ASAP) es un nuevo programa de investigación que combina

la sistemática, biogeografía, ecología y genética de poblaciones para estudiar la diversidad, abundancia, distribución y estructura de comunidades de artrópodos terrestres a lo largo de gradientes altitudinales y entre cadenas montañosas en el desierto del suroeste norteamericano. Los

artrópodos terrestres representan organismos taxonómica y ecológicamente diversos que impulsan procesos clave en el ecosistema. Utilizando

datos de especímenes de museo y material colectado en monitoreos de largo plazo con trampas de caída, ASAP documentará los patrones de

diversidad a través de las Islas Serranas para identificar los patrones de endemismo, filogeografía, y los orígenes evolutivos de los artrópodos

en estas comunidades. Además, ASAP utilizará el laboratorio natural proporcionado por las Islas Serranas para evaluar las funciones ecológicas

de los artrópodos a través de gradientes altitudinales y ambientales en las montañas. Los datos de referencia se utilizarán para determinar los

límites climáticos de las especies en observación, y se integrarán con modelos climáticos para predecir los cambios futuros en las comunidades

de artrópodos y su distribución después de un cambio climático rápido. Se describirá este proyecto y sus objetivos generales, y presentaremos

datos históricos sobre los artrópodos terrestres de las Islas Serranas, preservados en la Colección de Insectos de la Universidad de Arizona. En

subsecuentes charlas, los colaboradores mostrarán análisis de los datos obtenidos durante el primer año de ASAP en diversos aspectos de este

proyecto multidisciplinario.
USDA Forest Service Proceedings RMRS-P-67. 2013 561

Abstracts/ Resúmenes

NAROG, MARCIA G.1, Bonni M. Corcoran1, and Ruth C. Wilson2. 1USDA, Forest Service, 4955 Canyon Crest Dr, Riverside, CA 92507,

mnarog@fs.fed.us, bcorcoran@fs.fed.us; 2Department of Biology, California State University, San Bernardino, San Bernardino, CA.

Burned saguaro: Will they live or die? Thousands of acres in giant saguaro (Carnegiea gigantea) habitat of the Sonoran Desert continue to be

scorched by fire. Resource managers struggle to maintain scenic landscapes featuring majestic saguaro with the challenges of recreation, fire,

non-native species invasion and millions of annual visitors. Successfully managing this iconic saguaro community requires understanding fire

resilience or intolerance. Our study compares survival of 164 saguaro 10 years after two wildfires and illustrates fire’s negative consequences.

Six 350 m point quarter transects were randomly placed in burned and unburned areas of The Rolls, Mesa District, Tonto National Forest, Arizona. Four saguaros were measured at each of 8 points spaced 50 m apart along each transect. Ten years post-fire, high saguaro mortality and

stunted growth were observed in both burns. Saguaro mortality was 32% with apical height growth of 0.9 m after the Vista View Fire compared

to 51% and 0.6 m after the nearby River Fire. Unburned areas had 4% saguaro mortality and apical growth of 1.13 m. High compared to low

fire severity nearly doubled saguaro mortality and stunted growth of surviving saguaro by 30%. Protracted saguaro regeneration mandates

proactive fire management to sustain this cactus-shrub community. Twenty year follow-up surveys can substantiate longer-term fire effects on

this keystone species.

Sahuaros quemados, ¿vivirán o morirán? Miles de hectáreas de hábitat del sahuaro (Carnegiea gigantea) en el Desierto Sonorense siguen

siendoarrasadas porincendios.Elpersonaldemanejode recursosluchapormantenerlospaisajes pintorescos demajestuosossahuaros enfrentando

los desafíos de recreación, fuego, invasión de especies introducidas y millones de visitantes anuales. El manejo exitoso de esta comunidad

emblemática de sahuaros requiere comprender la resistencia o intolerancia a los incendios. Nuestro estudio compara la supervivencia de 164

sahuaros 10 años después de dos incendios forestales, e ilustra las consecuencias negativas de los incendios. Seis transectos de punto de 350m

se ubicaron al azar en zonas quemadas y sin quemar del área The Rolls, Distrito de Mesa, Bosque Nacional de Tonto, Arizona. Cuatro sahuaros

se midieron en 8 puntos separados por 50 m de distancia en cada transecto. Diez años después del incendio se observó alta mortalidad y retraso

de crecimiento de sahuaros en ambas zonas quemadas. La mortalidad de sahuaros fue de 32% con un crecimiento apical de 0.9 m después del

incendio de Vista View en comparación con el 51% y 0.6 m después del incendio cercano de River. Las zonassin quemar tuvieron una mortalidad

de 4% y crecimiento apical de 1.13 m.Alta severidad comparada con baja severidad del incendio casi duplicó la mortandad de sahuaros y retrasó

el crecimiento de sahuarossobrevivientes en un 30%. La regeneración prolongada de sahuaro requiere la prevención de incendios para mantener

esta comunidad de cactus-arbusto. Monitoreos cada veinte años pueden corroborar los efectos a largo plazo por incendios en esta especie clave.

NICHOLAS, HILLARY D. and Thomas Meixner. University ofArizona, Department of Hydrology and Water Resources, 1133 E James Rogers

Way, Tucson, AZ 85721, Hillarydianne@gmail.com.

Estimating flow frequency and infiltration volumes in intermittent streams in southern Arizona. Directional climate change may lead to

increased aridity and fewer precipitation events across the American southwest. Determining infiltration fluxes during monsoonal rainstorms

may be the key to predicting how changing precipitation frequency will affect groundwater percolation and potential recharge. Our study focuses on the following research question: How much water flows and infiltrates during storm events in ephemeral streams? In order to address

these topics, twelve intermittent stream sections in southern Arizona have been chosen for infiltration flux analysis. These streams have been

instrumented with temperature and electrical resistivity loggers which can detect flow and be used to estimate percolation ratesin the subsurface.

Data collection has yielded several months of background diurnal cycles and clear changes in variable saturation which preliminarily suggest

infiltration rates on the scale of 10–5 m/s at peak flow. Each of the twelve stream reaches in this study fall across different flowpaths, geology,

vegetation, and precipitation frequency, which dictate the dynamics between the hydrology and ecosystem processes. The hydrological aspect of

this study is partnered with a vegetation analysis and an entomological study in order to develop a comprehensive investigation of the complex

characterization of variably saturated environments.

Estimación de la frecuencia de caudales y volumen de infiltración en arroyos intermitentes en el sur de Arizona. El cambio climático

direccional puede ocasionar un aumento de aridez y menor precipitación en el Suroeste Norteamericano. Determinar los flujos de infiltración

durante las tormentas del monzón puede ser la clave para pronosticar como el cambio de frecuencia de la precipitación afectará la filtración y la

potencia de recargar el acuifero. Nuestra investigación se enfoca en la siguiente pregunta: ¿Cuánta agua fluye y se infiltra durante las tormentas

en los arroyos efímeros? Con el fin de abordar este tema se escogieron doce arroyos intermitentes del sur de Arizona para un análisis del flujo

de infiltración. Se colocaron instrumentos electrónicos que registran temperatura y resistencia eléctrica para detectar el caudal, y así calcular

las tasas de filtración en el subsuelo. La toma de datos arrojó un historial de ciclos diurnos de varios meses y cambios evidentes de la saturación

variable donde los resultados preliminares muestran tasas de infiltración en una escala de 10–5 m/s durante el caudal máximo. Cada uno de los

doce tramos de arroyos en esta investigación tiene diferente cauce, geología, vegetación y frecuencia de lluvias, que dictan la dinámica entre

la hidrología y los procesos del ecosistema. El aspecto hidrológico de esta investigación está ligado con un análisis de vegetación y un estudio

entomológico con el fin de desarrollar una investigación exhaustiva de las características complejas de ambientes de saturación variable.

O’CONNOR, CHRISTOPHER D.1,3, Donald A. Falk1, Ann M. Lynch2,3, and Thomas W. Swetnam3. 1School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721, oconnorc@email.arizona.edu, dafalk@email.arizona.edu ; 2

USDA Forest Service Rocky

Mountain Research Station, Fort Collins, CO 80526, alynch@fs.fed.us; 3Laboratory of Tree-Ring Research, University of Arizona, Tucson,

AZ 85721, tswetnam@ltrr.arizona.edu.

Fire and forest dynamics along an elevational gradient of the Pinaleño Mountains of Arizona. Fire and climate are important drivers of

forest dynamicsin NorthAmerica. Studies of fire activity along elevational gradients provide an opportunity to examine how fire spreads among
562 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

stands of different species assemblages, and how climate, topography, and stand conditions influence fire behavior. We are examining spatial

and temporal distributions of fire and species dynamics in ponderosa-oak, mixed conifer, and spruce-fir forests of the Pinaleño Mountains.

Dendroecological reconstructions are based on sampling trees from a grid of 78 plots covering the area above 7,000 ft. elevation (2,134 meters).

Within and around plots we systematically sampled trees with fire scars. Preliminary results from 1,211 plot trees and 110 fire-scarred samples,

suggest distinct patterns of seedling recruitment and differences in fire frequency and severity in adjoining forest types prior to 1900. Post

1900 seedling recruitment shows synchrony across forest assemblages, suggesting a shift from stand-level to landscape level controls. Pulses

of seedling recruitment and exclusion of fire from lower elevation dry forests has resulted in forest structure that facilitates fire spread among

forest types with very different historical fire regimes. These structural and compositional changes have implications for tree recruitment and

longevity, retention of fire-adapted species, and resilience following future disturbances.

Dinámica de incendios y bosques en un gradiente altitudinal de la cordillera Pinaleño de Arizona. Fuego y clima son factores importantes

de la dinámica de bosques en América del Norte. Los estudios de actividad del fuego en gradientes de elevación proporcionan una oportunidad

para examinar cómo el fuego se propaga entre lugares con asociaciones de especies diferentes, y cómo el clima, topografía, y condiciones del

área influyen en el comportamiento del fuego. Estamos examinando la distribución espacial y temporal de la dinámica de incendios y especies

en bosques de ponderosa-encino, coníferas mixtas y picea-abies de la Sierra Pinaleño. Reconstrucciones dendroecológicas están basadas en el

muestreo de árboles de una cuadrícula de 78 parcelas que cubren el área arriba de 2,134 m de elevación. Dentro y alrededor de las parcelas se

hizo un muestreo sistemáticamente de árboles con cicatrices de fuego. Losresultados preliminares de 1,211 árboles y 110 muestras con cicatrices

de fuego, sugieren distintos patrones de reclutamiento de plántulas y diferencias en frecuencia y severidad de incendios en los bosques adyacentes antes de 1900. Después de 1900 el reclutamiento de plántulas muestra sincronía en los ensambles del bosque, lo que sugiere un cambio

del control a nivel localizado a influencia a nivel de paisaje. Los pulsos de reclutamiento de plántulas y la exclusión de incendio de bosques

secos de baja elevación ha ocasionado una estructura del bosque que facilita la propagación del fuego entre tipos de bosques con regímenes

históricos de incendios muy diferentes. Estos cambios estructurales y de composición tienen implicaciones para el reclutamiento y longevidad

de árboles, la retención de especies adaptadas al fuego, y la capacidad de recuperación de futuros disturbios.

OLDEN, JULIAN D.1, Kristin L. Jaeger1, and David A. Lytle2. 1School of Aquatic and Fishery Sciences, University of Washington, Seattle, WA

98105, olden@uw.edu; 2 Department of Zoology, Oregon State University, Corvallis, OR 97331.

Streamflow variability of dryland rivers in a changing climate. Ephemeral and intermittent streams in the southwestern United States exist

in a precarious balance between drying and flooding. Although dryland streams are already naturally fragmented ecosystems, there is concern

of further habitat loss and decreased longitudinal connectivity associated with more frequent and severe droughts caused by climate change

and greater human appropriation of water resources. Characterizing the considerable spatiotemporal variability in streamflow, both today

and into the future, remains a paramount challenge. Using a spatial array of electrical resistance sensors, we quantified streamflow continuity

(continuous through time) and longitudinal connectivity (continuous through space) across watersheds in the Huachuca Mountains, Arizona.

Continuous monitoring revealed that stream reaches are characterized by a range of hydrologic flow permanence, and that continuous streamflow in intermittent and ephemeral reaches accounted for <30% and < 5% of the year, respectively. Canyon-wide longitudinal connectivity and

reach-level flow continuity was rare. Comparison of streamflow patterns between 2010 (“normal/wet”) and 2011 (“dry”) water year showed

that decreases in winter and spring precipitation can result in earlier stream drying, with shifts from perennial to intermittent or intermittent to

ephemeral flow regimes. Rain-runoff modeling demonstrated significant potential for losses in habitat availability and depressed longitudinal

connectivity according to climate change.

Variabilidad de caudal de ríos de zonas áridas en un clima cambiante . Los arroyos efímeros e intermitentes del suroeste de Estados Unidos

subsisten en un equilibrio precario entre sequía e inundaciones.Aunque los arroyos de zonas áridasson ecosistemasfragmentados por naturaleza,

existe la preocupación de una mayor pérdida de hábitat y la disminución de la conectividad longitudinal, asociada con sequías más frecuentes y

severas ocasionadas por el cambio climático y un mayor uso humano de recursos hídricos. Caracterizar la variabilidad espacio-temporal de estos

caudales, tanto hoy como en el futuro, es un desafío primordial. Con sensores de resistencia eléctrica se cuantificó la continuidad de caudales

(continua en el tiempo) y la conectividad longitudinal (continua en el espacio) en las cuencas hidrográficas de la Sierra Huachuca, Arizona.

El monitoreo continuo reveló que los tramos de arroyos se caracterizan por una oscilación de permanencia del flujo hidrológico, y el caudal

continuo en tramos intermitentes y efímeros representan <30% y <5% del año, respectivamente. La conectividad longitudinal a lo ancho de

cañones y continuidad de corrientes en tramos fue escasa. Comparación de los patrones de caudales entre los años 2010 (“normal / húmedo”)

y 2011 (“seco”) demostró que la disminución de precipitación de invierno y primavera puede causar que los arroyos se sequen más pronto,

con cambios en los regímenes de corrientes perennes a intermitente o de intermitente a efímeras. Los modelos de lluvia-escorrentía indican un

potencial significativo de pérdidas en la disponibilidad de hábitat y una conectividad longitudinal baja de acuerdo al cambio climático.

OLGUÍN-VILLA, MÓNICA CECILIA and Melinda Cárdenas-García. Universidad de Sonora, Blvd. Luis Encinas y Rosales s/n, Col. Centro,

Hermosillo, Sonora, México, moce89@gmail.com, melindacardenas@hotmail.com.

Avifauna of the San Bernardino Ranch, Agua Prieta, Sonora, Mexico. The interest and research on grassland birds have increased in recent

decades due to the decline of many of their populations and the fragmentation or loss of their habitats. The San Bernardino Ranch is located in

desert grassland in northeastern Sonora. For 10 years restoration activities have been conducted because previous economic activities deteriorated the habitat. Therefore, it is important to develop annual lists of flora and fauna that can hel evaulate the degree of restoration. As part of
USDA Forest Service Proceedings RMRS-P-67. 2013 563

Abstracts/ Resúmenes

the Professional Practices program at the Universidad de Sonora, we conducted a taxonomic list of birds observed on the ranch in the summer

of 2011. During that season, we found a total of 53 species and 42 genera in 26 families. The sampling will be repeated during the winter to

estimate the number of migratory birds that find refuge on the ranch. With the complete record of the birds, we will learn about the effect of

habitat restoration in the area on the diversity of birds and species of concern for conservation.

Avifauna del rancho San Bernardino, Agua Prieta, Sonora, México. El interés y las investigaciones sobre las aves de pastizal se han incrementado en las últimas décadas, debido a la declinación de muchas de sus poblaciones y a la fragmentación o pérdida de sus hábitats. El rancho

San Bernardino se encuentra en la región de pastizal desértico en el noreste de Sonora; desde hace 10 años se han llevado a cabo acciones de

restauración, antiguamente se realizaban actividades económicas que deterioraban el hábitat. Por esto, es importante elaborar listados anuales

de fauna y flora que nos puedan proporcionar el grado de restauración del lugar. Como parte de las prácticas profesionales de biología, durante

el verano de 2011 se elaboró un listado taxonómico de la avifauna que alberga el rancho, durante esa estación del año se encontraron un total de

53 especies y 42 géneros distribuidos en 26 familias. Este mismo muestreo se llevará a cabo durante el invierno y de esta manera será posible

estimar la cantidad de aves migratorias que encuentran refugio en el rancho. Con el registro completo de las aves, se obtendrá información sobre

el impacto del grado de restauración del hábitat en la zona en la diversidad de aves y las especies de interés para la conservación.

PALTING, JOHN. Sky Island Alliance, 7339 N. Yucca Via, Tucson, Arizona, 85704, john.palting@ventana.roche.com.

Moth inventory of Rincón de Guadalupe, Sierra de Bacadéhuachi, Sonora, México. Moths were sampled in 2011 on two Madrean Archipelago Biodiversity Assessment (MABA) trips to the historic Rincón de Guadalupe Catholic Diocese camp in the Sierra de Bacadéhuachi in

the westernmost Serra Madre Occidental (SMO) in east-central Sonora México. The study area is in Arroyo Campo los Padres, 16.5 km (by

air) east-northeast of Bacadéhuachi. The habitat is pine-oak forest (POF) at 1680 m elevation. A trip in August during the peak of the rainy

season documented many sphingid, saturniid, and arctiid moth species typical of POF, including many previously known no farther north than

the Yécora area ca. 160 km to the south. Of special interest were a number of species associated with tropical deciduous forest, which normally

live at lower elevations. A trip in early September yielded a great diversity of smaller moths, especially those in the Noctuidae, which feed on

flowers in the larval stage. Both sampling periods resulted in more than 200 species documented on each trip, with a total of more than 400

species. The moth fauna of the area reflects a dramatic convergence of species from the Rocky Mountains, the montane SMO forests, and the

tropical lowlands of northwestern México. At least 12 species have been determined to be new to science and await description. The remarkable species richness of moths, the unspoiled natural beauty of the area, and the rich cultural history make this area a standout for preservation.

Inventario de las palomillas del Rincón de Guadalupe, Sierra de Bacadéhuachi, Sonora, México. Las palomillasfueronmuestreadas en 2011

durante dos expediciones del proyecto Evaluación de la Biodiversidad del Archipiélago Madrense (MABA) al campamento histórico Rincón de

Guadalupe, de la Diócesis Católica, en la Sierra de Bacadéhuachi al oriente centro de Sonora, México. La ubicación del área de estudio es en el

Arroyo Campo los Padres, a 16.5 km (por aire) este-noreste de Bacadéhuachi. El hábitat es bosque de pino-encino (BPE) a 1680 m de elevación.

El primer viaje en agosto durante la temporada alta de la estación lluviosa produjo varias especies de esfíngidos, satúrnidos y arctiidos típicas

de BPE, entre ellas varias especies conocidas previamente no más al norte del área de Yécora en la Sierra Madre Occidental (SMO) ca. de 160

km al sur. De especial interés son varias especies asociadas con la selva baja caducifolia, que normalmente viven en elevaciones más bajas. En

el segundo viaje a principios de septiembre se documentó una gran diversidad de palomillas pequeñas, en especial especies del género noctuid,

las que se alimentan de flores en su fase de larva. Los dos períodos de muestreo dieron como resultado más de 200 especies documentadas en

cada excursión, para un total de más de 400 especies. La fauna de palomillas del área refleja una convergencia dramática de especies de las

Montañas Rocosas, los bosques de montaña de la SMO y las regiones cálidas tropicales del noroeste de México. Por lo menos 12 especies se

han identificado como nuevas para la ciencia y se espera su descripción. Por su extraordinaria diversidad de especies de palomillas, belleza

natural y rica historia cultural este lugar debe protegerse.

PAVLIK, STEVE. Northwest Indian College, 2825 Kwina Road, Bellingham, WA 98226, spavlik@nwic.edu.

Ursus in a Sky Island range: The impact of drought and human activity on black bears in the Huachuca Mountains since 2004. At the

Madrean Archipelago II Conference, the author presented a twenty-year analysis (1983-2004) of black bear mortality in the Huachuca Mountains. This presentation will provide updated information on black bear mortality numbers and causes of mortality from 2005 through 2012,

linking this information to the drought conditions that have continued to plague the Sky Islands. This presentation will include an analysis of

the events of the fall and summer of 2006 when no less than 21 bears – over 40% of the total population – were killed or removed from the

Huachuca Mountains. Also included will be additional observations and comments regarding the role humans play in black bear mortality in

the Huachuca Mountains. The data presented in this presentation was collected through an analysis of Arizona Game and Fish records, rainfall

data provided by Fort Huachuca, newspaper articles, and personal interviews and communication with wildlife managers, bear biologists, local

landowners, and others. Among the results of this presentation will be data suggesting that far too many bears – especially females – are being

removed from the Huachuca Mountains, and that in light of recent drought, wildlife managers need to explore alternate means of managing

bears in this unique Madrean Archipelago population.

Ursus en una cordillera de Islas Serranas: impacto de la sequía y la actividad humana en osos negros en la Sierra Huachuca desde 

2004. En el II Congreso del Archipiélago Madrense, el autor presentó un análisis de veinte años (1983-2004) de la mortalidad del oso negro

en la Sierra de Huachuca. Esta presentación aportará información actualizada sobre registros y causas de mortalidad de osos negros a partir de

2005 hasta el 2012, vinculando esta información a las condiciones de sequía que han seguido afectando las Islas Serranas. Esta presentación
564 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

incluirá un análisis de los acontecimientos del otoño y verano de 2006, cuando al menos 21 osos, más del 40% de la población total, murieron

o se llevaron fuera de la Sierra Huachuca. También se incluirán observaciones y comentarios adicionales sobre el papel de los humanos en la

mortalidad del oso negro en las montañas Huachuca. Los datos de esta investigación provienen de un análisis de registros de Arizona Game and

Fish, datos de precipitación de Fort Huachuca, artículos periodísticos y entrevistas y comunicación con personal de manejo de fauna, biólogos

del oso, propietarios privados, entre otros. Los resultados sugieren que demasiados osos, especialmente las hembras, se han sacado de la Sierra

Huachuca, y en vista de la reciente sequía, el personal de manejo de fauna necesita explorar otras opciones de manejo para esta población de

osos única del Archipiélago Madrense.

PEARTHREE, PHILIPA. and Ann Youberg. Arizona Geological Survey, 416 W. Congress St., Tucson, AZ 85701, phil.pearthree@azgs.az.gov,

ann.youberg.azgs.az.gov.

Fire and debris flows in the Huachuca Mountains of southeastern Arizona. The occurrence of debris flows in the 2011 monsoon season

following the Monument Fire in the Huachuca Mountains demonstrated again the causal linkage between wildfire and debris flows in southern

Arizona. Debris flows are distinguished from water floods by their higher densities and lower viscosities, which permits them to transport large

quantities of coarse sediment. Because debris flows perform more work shaping channels and moving coarse material than the combined effects of hundreds of years of water floods, they are critical agents in shaping the watersheds of the mountains of southern Arizona. Previous

studies had documented debris flows in monsoon seasons following fires in 1977, 1988, and 2006 in the central and southern Huachucas. Vastly

increased runoff from steep, moderately to highly burned watersheds led to erosion of sediment from gullies and channels and the generation

of debris flows. Debris flows in the Huachucas have typically excavated nearly all sediment from steep channel reaches, suggesting that return

times between debris flows in individual watersheds might be several centuries. The re-occurrence of debris flows in Miller Canyon in 2011

implies that in the modern period with fairly frequent large wildfires, the return time between debris flows may be much shorter.

Incendios forestales y flujo de detritos en la Sierra Huachuca del sureste de Arizona. El flujo de detritos durante la temporada del monzón

de 2011 después del incendio forestal conocido como ‘Monument Fire’ en la Sierra Huachuca mostró la conexión entre incendios y flujos de

detritos en el sur de Arizona. Los flujos de detritos se distinguen de las inundaciones por su alta densidad y su baja viscosidad lo que permite el

transporte de grandes cantidades de sedimento grueso. Los flujos de detritos son procesos críticos en la formación de cuencas hidrográficas en

las montañas del sur de Arizona ya que su efecto es mayor formando canales y moviendo material grueso que los efectos de cientos de años de

inundaciones. Estudios previos, han documentado flujos de detritos en temporadas del monzón después de incendios forestales en 1977, 1988

y 2006 en las partes centrales y sureñas de la Sierra Huachuca. El gran aumento de escorrentía en cuencas empinadas con niveles de quemazón

de moderados a altos causó la erosión de sedimentos de barrancos y canales y la generación de flujos de detritos. En la Sierra Huachuca los

flujos de detritos típicamente han excavado casi todo el sedimento de los tramos mas empinados, sugiriendo que el flujo de detritos en cuencas

individuales puede ser de varios siglos. La reaparición de flujos de detritos en Miller Canyon en 2011 sugiere que en esta era moderna con

grandes incendios forestales frecuentes, el tiempo entre flujos de detritos es mas corto.

PERÉZ-WEIL, LUCIA. Naturalia, Hermosillo, Sonora, reservajaguar@gmail.com.

Buying land for conservation purposes in Sonora, Mexico. The Northern Jaguar Reserve is approximately 50,000 acres, comprising the

largest privately owned wildlife preserve in Sonora. Buying land in remote parts of Sonora takes special talents as ownership rules may not be

clear, boundaries may not be defined in the records, there are complex legal procedures to guarantee ownership in which Letters of Intent hold

special powers and special complexities for ejido lands. In addition, only a Presidential Decree can prevent mining, and ownership of riparian

land—whether by governmental or private parties—is controversial. The legal framework for conservation easementsisin itsinfancy in Sonora.

The lessonslearned at the NJRfor conservation land purchases will be described. NJPhasinvited participation by additional purchasers/managers

of private reserve lands in Sonora to describe their lessons learned during the Question and Answer period following the presentations. These

participants will include the Cuenca los Ojos Foundation (Valer Austin) and Rancho Los Fresnos (Juan Carlos Bravo).

Compra de tierras para conservación en Sonora, México. La Reserva del Jaguar del Norte (RJN) tiene un área aproximada de 20,000

hectáreas y representa la reserva natural privada más grande de Sonora. La compra de tierras en zonas remotas de Sonora requiere talentos

especiales, ya que los títulos de propiedad pueden ser confusos, los linderos muchas veces no están bien marcados y trámites legales engorrosos para garantizar el cambio de propietario, donde las cartas de intención conceden poderes especiales a terrenos ejidales. Además, sólo por

decreto presidencial se pueden negar actividades mineras y propiedad de terrenos ribereños, ya sea por agencias del gobierno o privados, es

objeto de controversia. El marco legal de las servidumbres ecológicas apenas empiezan en Sonora. Se presentará lo aprendido por RJN en la

compra de tierras para conservación. RJN ha extendido una invitación a otros encargados de compra de terrenos y manejo de reservas privadas

en Sonora para compartir experiencias durante la sesión de preguntas y respuestas. Se contará con la presencia de Fundación Cuenca los Ojos

(Valer Austin) y Rancho los Fresnos (Juan Carlos Bravo).
USDA Forest Service Proceedings RMRS-P-67. 2013 565

Abstracts/ Resúmenes

PONCE, EDUARDO1, Ana Davidson2, Ed Fredrickson3, and Gerardo Ceballos1. 1Instituto de Ecología, Universidad Nacional Autónoma de

México, México, DF 04510 México; 2Department of Biology, University of New Mexico, Albuquerque, New Mexico 87131; 3Department of

Agriculture, Eastern Kentucky University, Richmond, KY 40475.

Managing prairie dogs, cattle and honey mesquite: Ecological implications for grassland conservation. On last decades honey mesquite

(Prosopis glandulosa) has increased its abundance in arid grasslands of northern Mexico and southern US. This process has been attributed

mainly to overgrazing by livestock and eradication of native fauna. Although herbivory by rodents has been considered as an important element in desert grasslands, it has not been given much importance as an element capable to control the shrub establishment. Our objective was

to determine the independent and interactive effects of black-tailed prairie dog (Cynomys ludovicianus) and cattle on mesquite abundance and

structure in a desert grassland ecosystem of northern Chihuahua, Mexico. Cattle and prairie dogs were simultaneously manipulated using a

controlled, long-term, replicated experiment with a 2x2 factorial design. In five years, mesquite abundance increased in sites were prairie dog

were removed, no mater if cattle were present or not. Prairie dogs controlled the mesquite growing and cover, but the effect increased when

cattle also was present. The study of these interactions is critical to understand the role of both species on the shrub expansion and grassland

conservation in order to develop better management practices in sites were both species are still present.

El manejo de perritos de la pradera, ganado y mezquite: Implicaciones ecológicas para conservación de pastizales. En las últimas décadas

el mezquite (Prosopis glandulosa) ha incrementado su abundancia en los pastizales áridos del norte de México y sur de Estados Unidos. Este

proceso se ha atribuido principalmente al sobrepastoreo por ganado doméstico y la erradicación de la fauna nativa. A pesar que la herbivoría de

pequeños roedores ha sido considerada como un elemento importante en los pastizales desérticos, no se le ha dado mucha importancia como

un elemento capaz de controlar el establecimiento de arbustos. Nuestro objetivo fue determinar los efectos independientes e interactivos del

perrito de la pradera de cola negra (Cynomys ludovicianus) y ganado en la abundancia y estructura del mezquite en un ecosistema de pastizal

desértico del norte de Chihuahua, México. Los perros de las praderas y el ganado doméstico fueron manipulados de forma simultánea en un

experimento a largo plazo con un diseño factorial 2x2. En cinco años, la abundancia de mezquite aumentó en sitios donde los perros de la pradera

se removieron, sin importar si el ganado estaba presente o no. Los perros de la pradera controlaron el crecimiento de mezquite y la cobertura,

pero el efecto fue mayor cuando el ganado también estaba presente. El estudio de estas interacciones es fundamental para entender el papel que

las dos especies tienen en la expansión de arbustos y la conservación de los pastizales, con el fin de desarrollar mejores prácticas de manejo en

pastizales donde ambas especies todavía están presentes.

POWELL, BRIAN1,2 and RobertJ. Steidl1. 1Pima County Office of Sustainability and Conservation, 201 N. StoneAve., Tucson,AZ 85701, brian.

powell@pima.gov; 2School of Natural Resources and the Environment, University of Arizona, Tucson, Arizona 85721, steidl@ag.arizona.edu..

Habitat characteristics of montane riparian bird communities in southern Arizona. In the Sky Island region of southern Arizona, mesic

montane riparian areas comprise a small percentage of the landscape, yet provide habitat for a disproportionately large proportion of the region’s

biodiversity. This is particularly true for birds, which have higher densities and species richness in riparian as compared to upland areas. Despite the importance of riparian areas, little research has focused on riparian areas where Sonoran desert-scrub grades into Madrean evergreen

woodland. In 1997 and 1998, we surveyed birds and vegetation at 46 point-count stations in two mountain ranges, Baboquivari and Atascosa,

to describe the bird community and to understand how vegetation characteristics influence bird abundance and diversity. We recorded >3,900

individuals representing 79 species and found that both vegetation structure and floristics were important in structuring the avian community.

In general, areas of dense vegetation in the understory, which are common in riparian areas, were important predictors of bird diversity. In

addition, strong gradients that were determined largely by the composition of dominant plant species affected the distribution and abundance

of bird species in the community. Understanding the habitat associations of these species will provide additional information to make sciencebased management decisions.

Características del hábitat de las comunidades de aves ribereñas de montaña en el sur de Arizona. En la región de las Islas Serranas del

sur de Arizona, las áreas ribereñas mésicas de montaña representan un pequeño porcentaje del paisaje, sin embargo, proporcionan el hábitat

para una porción bastante grande de la biodiversidad de la región. Esto en especial para las aves, que tienen una mayor densidad y riqueza de

especies ribereñas en comparación con zonas más altas. A pesar de la importancia de las áreas ribereñas, poca investigación se ha centrado en

estas zonas, donde el matorral desértico sonorense cambia a bosque perenne madrense. En 1997 y 1998, monitoreamos pájaros y vegetación en

46 estaciones de monitoreo en dos cadenas montañosas, Baboquivari y Atascosa, para describir la comunidad de aves y para entender cómo las

características de la vegetación influyen en la abundancia de aves y su diversidad. Registramos > 3,900 individuos de 79 especies y se encontró

que tanto la estructura de la vegetación y la flora fueron importantes en la composición de la comunidad de aves. En general las áreas de densa

vegetación en el sotobosque, comunes en las zonas ribereñas, fueron indicadores importantes de la diversidad de aves. Además, los fuertes

gradientes que se determinaron principalmente por la composición de especies de plantas dominantes afectó la distribución y abundancia de

especies de aves en la comunidad. El conocimiento de las asociaciones de hábitat de estas especies, aportará información adicional para tomar

decisiones de manejo con base científica.

PRIVAL, DAVE1 and Mike Schroff2. 1Southwestern Ecological Research Company, 530 S. Herbert Ave., Tucson, AZ 85701, deprival@yahoo.

com; 23547 E. Flower St., Tucson, AZ 85716.

Ecology and conservation of a Madrean rattlesnake, Crotalus pricei. Twin-spotted rattlesnakes (Crotalus pricei) are small-bodied snakes

found in six ranges within the Madrean Archipelago as well as the Sierra Madre Occidental and Sierra Madre Oriental. Within the Madrean
566 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Archipelago they are most common in Madrean montane conifer forest above 2400 m, making them the region’s highest elevation snake. A

warming, drying climatic trend is an obvious conservation concern. We monitored populations of C. pricei in Arizona’s Chiricahua Mountains

from 1997-2009 to learn about diet, reproduction, growth rates, and survival trends. We captured 306 individuals and recaptured snakes on 155

occasions. Spiny lizards (Sceloporus) were the most frequently taken prey for both juveniles and adults. Larger females were more likely to be

gravid than smaller adult females. Our best model indicated thatsurvival rate remained constant at our most-studied site over the 13 years; detectability varied by search effort and weather.Although there was no evidence of a population decline during ourstudy, we identified waysin which

climate change and continued illegal collection for the pet trade could pose a threat to the conservation of isolated, high elevation populations.

Ecología y conservación de una víbora de cascabel madrense, Crotalus pricei. Las chachamuris (Crotalus pricei) son víboras pequeñas

que se encuentran en seis cordilleras del Archipiélago Madrense, así como en la Sierra Madre Occidental y la Sierra Madre Oriental. En el

Archipiélago Madrense son más comunes en el bosque de coníferas de montaña madrense arriba de 2400 m, por lo que son la serpiente de

más alta elevación en la región. Una tendencia climática de calentamiento y sequía es claramente una preocupación para su conservación. Se

hizo un monitoreo de las poblaciones de C. pricei en la Sierra Chiricahua en Arizona de 1997-2009 para estudiar su dieta, reproducción, tasas

de crecimiento y supervivencia. Se capturaron 306 individuos y se recapturaron en 155 ocasiones. El cachorón (Sceloporus) fue la presa más

común atrapada tanto por jóvenes y adultos. Las hembras más grandes tuvieron más probabilidades de estar grávidas que las hembras adultas

más pequeñas. Nuestro mejor modelo indicó que la tasa de supervivencia se mantuvo constante en nuestro sitio más estudiado en los 13 años;

la detectabilidad varió basada en el esfuerzo de búsqueda y el clima. Aunque no detectamos disminución de la población en nuestro estudio,

identificamos formas en las que el cambio climático y la colecta ilegal para el comercio podrían representar una amenaza para la conservación

de las poblaciones aisladas de elevaciones altas.

QUIJADA-MASCAREÑAS, ADRIÁN1,3, Donald Falk1,4, Jeremy Weiss2, Mitch McClaran1, John Koprowski1, Melanie Culver1, Sam Drake1,

Stuart Marsh1, Willem van Leeuwen1, and Matt Skroch1. 1 School of Natural Resources and the Environment, University of Arizona, Biological

Sciences East , Tucson, AZ 85721; 2 Department of Geosciences, University of Arizona, Gould-Simpson Room 208, Tucson, AZ 85721; 3 Sky

Island Alliance, 300 E University Blvd., Suite 270, Tucson, AZ 85705; 4 Laboratory of Tree-Ring Research, University of Arizona, Tucson AZ

85721.

From genes to ecosystems and human well being: Ecological effects of climate change in Sky Islands. Climate change is causing rapid

changes in mountain ecosystems. These changes may have ecological consequences at many levels of biological organization, from genetic

viability of populationsto landscape-level ecosystem processes. We present modeled projections of climate change in the Sky Islands bioregion,

and review potential effects of these changes on ecosystems of the Madrean Archipelago Sky Islands of the US-Mexico border region as a

model system. Some low elevation species may experience an expansion in suitable area, while habitat for high elevation species may contract.

As a consequence, at the population level, high elevation species will be vulnerable to genetic isolation and extinction, while population size

may increase for low-elevation species. At the community level, plant-animal interactions may be disrupted as species respond differentially

to climate change, affecting phenology patterns, insect outbreaks, and other interactions. Ecosystem processes such as fire will be altered as

changing plant distributions modify fuel loads, and as changes in temperature, wind and humidity alter the physical fire environment. Sky Island ecosystems represent important “indicator ecosystems” for monitoring ecological responses to climate change across elevation gradients.

Desde genes a ecosistemas y bienestar humano: Efectos ecológicos del cambio climático en las Islas Serranas. El cambio climático

está provocando rápidos cambios en los ecosistemas de montaña. Estos cambios pueden tener consecuencias ecológicas en muchos niveles

de organización biológica, desde la viabilidad genética de las poblaciones hasta los procesos del ecosistema a nivel de paisaje. Presentamos

modelos del cambio climático en la bioregión de las Islas Serranas, con una síntesis de los posibles efectos de estos cambios en los ecosistemas

del Archipiélago Madrense de Islas Serranas de la región fronteriza México-Estados Unidos. Algunas especies de elevaciones bajas podrían

aumentar su rango de distribución, mientras que el hábitat de especies de elevaciones altas podría reducirse. Como consecuencia, a nivel de población, las especies de elevaciones altas serán vulnerables al aislamiento genético y la extinción, mientras que el tamaño de la población podría

aumentar para las especies de elevaciones bajas. A nivel comunidad, las interacciones planta-animal podrían interrumpirse ya que las especies

responden de forma diferente al cambio climático, afectando patrones de fenología, plagas de insectos y otras interacciones. Los procesos del

ecosistema como el fuego se alterarán ya que el cambio de distribución de plantas modificará la vegetación combustible, así como los cambios

en temperatura, viento y humedad alteran el ambiente físico para incendios. Los ecosistemas de las Islas Serranas son importantes “ecosistemas

indicadores” para monitorear las respuestas ecológicas al cambio climático en gradientes de elevación.

RADKE, WILLIAM R. U.S. Fish and Wildlife Service; San Bernardino/Leslie Canyon National Wildlife Refuges, P.O. Box 3509, Douglas,

AZ 85608. Bill_Radke@fws.gov.

Land management on the United States/Mexico border. Many conservation strategies have been developed by the Service in cooperation with

others to protect habitat and enhance the recovery of fish and wildlife populations in the San Bernardino Valley, straddling Arizona and Sonora.

Damage to habitats along this international border by various illegal activities is detrimental to species recovery. In addition, potential threats

to national security have prompted the United States to improve control over the country’s boundaries, creating additional challenges for land

managers mandated with protecting the nation’s landscapes, natural resources, and associated values. Such challenges are not insurmountable,

and with minimal effort both resource management and border security can often complement one another. With or without the influence of

changes along the international border, effective species recovery methods must include a coordinated approach which involves: determining

and assessing the biological requirements of selected species through combinations of inventory, monitoring, and research activities; manag-
USDA Forest Service Proceedings RMRS-P-67. 2013 567

Abstracts/ Resúmenes

ing and protecting existing and historic habitats and populations; assessing potential reintroductions of key species into appropriate habitats;

managing exotic plants and animals that threaten the recovery of desired conditions; and developing and providing information and education

relative to the species, their habitats, and the ecosystems upon which all fish, wildlife, and humans depend.

Manejo de recursos naturales en la frontera de México y Estados Unidos. Muchas estrategias de conservación han sido desarrolladas por

U.S. Fish and Wildlife Service y colaboradores para proteger el hábitat y mejorar la recuperación de las poblaciones de peces y fauna en el

valle de San Bernardino, que se extiende a ambos lados de la frontera entre Arizona y Sonora. El daño a los hábitats causado por las diversas

actividades ilegales, en esta frontera internacional, es perjudicial para la recuperación de especies. Además, las amenazas potenciales a la

seguridad nacional han llevado a Estados Unidos a mejorar el control de las fronteras del país, creando retos adicionales para el personal de

manejo encargados de proteger los paisajes de la nación, recursos naturales y bienes asociados. Estosretos no son insuperables, y con un mínimo

esfuerzo tanto el manejo de recursos y la seguridad de la frontera a menudo se complementan entre sí. Con o sin la influencia de los cambios en

la frontera internacional, métodos eficaces de recuperación de especies deben incluir un enfoque coordinado que implica identificar y evaluar

los requisitos biológicos de especies seleccionadas al vincular inventarios, monitoreo y actividades de investigación; manejo y protección de

hábitats y poblaciones actuales e históricas; evaluar la reintroducción potencial de especies clave en hábitats adecuados; manejo de plantas y

animales exóticos que amenazan la recuperación; y el desarrollo de materiales educativos e información sobre las especies, sus hábitats y los

ecosistemas de los que tanto dependen los peces, fauna y flora y los seres humanos.

RATZLAFF, KRISTINA M.1, Cecil Schwalbe1, Phil Rosen1, and Don E. Swann2. 1University of Arizona, 1110 East South Campus Dr. Rm.

123, Bldg 33, Tucson, AZ 85721, kris.ratzlaff@gmail.com; 2Saguaro National Park, 3693 South Old Spanish Trail, Tucson, AZ 85730-5601,

Don_Swann@nps.gov.

Amphibian disease and Lowland Leopard Frogs: Trials in identification, treatment, and death. Population declines of amphibians have

been recorded worldwide. When two populations of lowland leopard frog became locally extirpated from the Rincon Mountains of Saguaro

National Park, eggs and tadpoles of this frog were taken from the park and placed in backyard ponds nearby for potential future reintroduction.

Through an unknown vector, frogs in these ponds were infected with the amphibian chytrid fungus, Batrachochytrium dendrobatidis (“Bd”),

and in some cases had near 100% mortality each winter following initial metamorphosis due to this disease. While these backyard ponds were

initially intended as ideal breeding facilities, they now provide a unique opportunity to learn more about how lowland leopard frogs interact

and respond to Bd on a smaller scale, without affecting frog populations in the wild. We present findings of what we have learned about

lowland leopard frogs and Bd from two backyard ponds, including identification of dead frogs, the possibility of freezing frogs found dead and

later obtaining usable Bd assays from them, and the feasibility of treating wild lowland leopard frogs for Bd to help them survive the winter.

La enfermedad anfibia en ranas yavapai: pruebas de identificación, tratamiento y muerte. Se han registrado descensos de poblaciones de

anfibios en todo el mundo. Cuando dos poblaciones de rana yavapai desaparecieron de las montañas Rincón del Parque Nacional Saguaro, los

huevos y los renacuajos de esta rana se llevaron del parque y se pusieron en estanques de patio cercanos para su futura reintroducción. Mediante

un vector desconocido, las ranas de estos estanques se infectaron con el hongo quítrido anfibio, Batrachochytrium dendrobatidis (“Bd”), y cada

invierno después de la metamorfosis inicial se registró una mortalidad casi del 100%, en algunos casos, por causa de esta enfermedad. Aunque

estos estanques originalmente pretendían ser instalaciones para la reproducción, ahora proveen una oportunidad única para aprender más sobre

las interacciones de las ranas yavapai y cómo responden al Bd a menor escala, sin afectar las poblaciones silvestres. Presentamos los resultados

de lo aprendido sobre las ranas yavapai y el Bd de dos estanques de patio, incluyendo la identificación de ranas muertas, la posibilidad de congelarlas y obtener de ellas muestras utilizables de Bd y la viabilidad de tratar las ranas silvestres con Bd para ayudarlas a sobrevivir el invierno.

RAYMONDI, ANN MARIE1, Cameron Childs2. School of Sustainability, Arizona State University, Tempe, AZ 85281, 1araymon3@asu.edu, 2

cchilds@asu.edu.

Institutional challenges in landscape-level fire restoration in the Sky Islands. Landscape-level management of fire transcends jurisdictional

boundaries and requires participation by multiple stakeholders. This presents a broad array of ecological and social challengesthat collaborative

projects are emerging to address. Collaborative fire management requires scientific understanding of complex system dynamics, as well as an

understanding of how to design effective partnerships, decision-making processes, and procedures to incorporate public input in management

goals. Thus, there is an inherent need to consider institutions, or formal rules and social norms, in collaborative fire management. In response

to the threat of increasingly severe wildfires, the Coronado National Forest recently unveiled FireScape, a collaborative program to safely manage and reintroduce fire into the Sky Islands of southeastern Arizona. This paper will discuss the unique social and institutional context facing

FireScape managers and partners. Other collaborative programs have struggled because of conflict with the public, lawsuits, or breakdown in

partnerships. As this collaborative arrangement develops, we will identify the unique institutional barriers that may need to be addressed to

ensure desirable social and ecological outcomes. FireScape will be used as a case study to analyze the institutional barriers and opportunities

inherent in achieving safe, ecologically sound, and sustainable fire management in the Sky Islands.

Retos institucionales en restauración del paisaje después de incendios en las Islas Serranas. El manejo del fuego a nivel paisaje trasciende

los límites jurisdiccionales y requiere la participación de los sectores interesados. Esto representa grandes retos ecológicos y sociales por lo

que están surgiendo proyectos de colaboración para enfrentarlos. El manejo colaborativo de incendios requiere la comprensión científica de la

dinámica de sistemas complejos, así como la capacidad de diseñar alianzas eficaces, procesos de toma de decisiones y formas de incorporar la

aportación del público en las metas de manejo. Por lo tanto es esencial considerar instituciones y leyes y normas sociales en el manejo colabo-
568 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

rativo de incendios. En respuesta a la amenaza de los incendios forestales cada vez más graves, el Bosque Nacional Coronado dio a conocer

recientemente FireScape, un programa de colaboración en manejo y reintroducción sin riesgo del fuego en las Islas Serranas del sureste de

Arizona. Este trabajo abordará el contexto social e institucional que enfrentan los coordinadores de FireScape y socios. Otros programas de

colaboración han tenido problemas debido a conflicto con el público, demandas, o ruptura de las alianzas. Conforme este acuerdo de colaboración se desarrolla, identificaremos las barreras institucionales que es necesario abordar para garantizar los resultados ecológicos y sociales

deseados. Se utilizará FireScape como caso de estudio para analizar las barreras institucionales y las oportunidades para lograr un manejo del

fuego seguro, ecológicamente responsable y sostenible de las Islas Serranas.

REINA-GUERRERO, ANA LILIA and Thomas R.. Van Devender. Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717, yecora4@

comcast.net .

Flora of the Limestone Sierra Anibácachi, Municipio de Agua Prieta, Sonora. A total of 590 plants collections were made in the La Calera

area in northeastern Sonora on 20 trips in 2002-2008. The 25 km2 (2.5 km2 extensively inventoried) area in the Sierra Anibácachi, Municipio

de Agua Prieta, is 11 km south of the Arizona border (31°13’59”N 109°37’53”W, ca. 1287 m elevation). Chihuahuan desertscrub on limestone

substrates is dominated by creosotebush (Larrrea divaricata), Chihuahuan whitethorn (Acacia neovernicosa), mariola (Parthenium incanum),

and tarbush (Flourensia cernua). Riparian vegetation along a rocky bedrock/gravel wash includes desert willow (Chilopsis linearis), netleaf

hackberry (Celtis reticulata), desert hackberry (C. pallida), woolly buckthorn (Sideroxylon lanuginosa), soaptree yucca (Yucca elata), Arizona

walnut (Juglans, major), and Coahuila juniper (Juniperus coahuilensis). The flora is diverse with 334 taxa in 62 families. The most important

plants are in the Poaceae (58 taxa),Asteraceae (48), Fabaceae (24), and Malvaceae (23) in the genera Euphorbia (11), Abutilon (7), Bouteloua (7),

Eragrostis (6), Ipomoea (6), and Muhlenbergia (6). Ten species are the first records for the state of Sonora, including Chamaesaracha sordida,

Cyphomeris gypsophiloides, Hybanthus verticillatus, Physaria fendleri, Quercus pungens, Ruellia parryi, Sphaeralcea polychroma, and Vicia 

ludoviciana. Other noteworthy Sonoran records include Bernardia myricaefolia, Dalea formosa, Phyllanthus polygonoides, and Vauquelinia 

californica ssp. pauciflora, among others.

Flora de las calizas de la Sierra Anibácachi, municipio de Agua Prieta, Sonora. Se colectaron un total de 590 ejemplares de herbario (con

varios duplicados) en el área de La Calera en el noreste de Sonora, en 20 salidas de campo de 2002 a 2008. Los 25 km2 del área (2.5 km2 colectados de forma intensa) de la Sierra Anibácachi, municipio de Agua Prieta, se localizan 11 km al sur de la frontera con Arizona (31°13’59”N

109°37’53”W, ca. 1287 m elevación ). El matorral del Desierto Chihuahuense en substratos de caliza está dominado por hediondía (Larrrea 

divaricata), vinorama (Acacia neovernicosa), hierba ceniza (Parthenium incanum) y Flourensia cernua. La vegetación ribereña en el arroyo de

lecho rocoso y grava incluye: jánota (Chilopsis linearis), cúmero (Celtisreticulata), garambullo (C. pallida), huajuco (Sideroxylon lanuginosa), 

nogal (Juglans, major), huata (Juniperus coahuilensis) y Yucca elata. La flora es diversa con 334 taxones en 62 familias. Las especies más

numerosas pertenecen a las familias Poaceae (58 taxones), Asteraceae (48), Fabaceae (24) y Malvaceae (23) y a los géneros Euphorbia (11),

Abutilon (7), Bouteloua (7), Eragrostis (6), Ipomoea (6) y Muhlenbergia (6). Diez especies son los primeros registros para Sonora, entre ellos

Chamaesaracha sordida, Cyphomeris gypsophiloides, Hybanthus verticillatus, Physaria fendleri, Quercus pungens, Ruellia parryi, Sphaeralcea policroma y Vicia ludoviciana. Otros registros importantes para Sonora incluyen Bernardia myricaefolia, Dalea formosa, Phyllanthus 

polygonoides y Vauquelinia californica ssp. pauciflora, entre otros.

DAN ROBINETT1 and Linda Kennedy2. 1Robinett Rangeland Resources LLC, HC 1 Box 208, Elgin,AZ 85611, dgrobinett@gmail.com; 2Linda

Kennedy, Appleton- Whittell Research Ranch of the National Audubon Society, HC 1 Box 208, Elgin, AZ 85611, lkennedy@audubon.org.

Babacomari River Riparian Protection Project. The Babacomari River is a major tributary of the San Pedro River in Santa Cruz and Cochise

Counties, Arizona. This 140,000 acre catchment includes rolling grasslands on the Sonoita plain, oak woodlands in the Canelo Hills and the

pine-oak forests of the northwestern Huachuca Mountains. The Babacomari River runs for 22 miles from its headwaters near Sonoita at 5000

feet elevation, eastward to join the San Pedro at Fairbanks at an elevation of 3850 feet. The US Geological Survey estimates that this important

tributary contributes about 6000 acre feet of water annually to the San Pedro River system. The Arizona Dept. of Water Resources funded this 5

year study with a grant (09-164WPF) in 2009. Monitoring transects were installed in 2009 and 2010 and will be re-read each year through 2013.

Objectives: Construct 2 miles of riparian boundary fence to restrict access by livestock from the Babacomari River. Install six stream riparian

vegetation and geomorphic monitoring transects and six vegetation and geomorphic transects on riparian grasslands (sacaton) tributary to the

Babacomari River. Analyze and summarize data annually and present that information to the participating ranch properties for use in making

management decisions.Riparian monitoring stations were established in May of 2010 at three locations along the Babacomari River below the

Babacomari Ranch headquarters. Riparian monitoring stations were established at three locations in June of 2010 on the Appleton-Whittell

Research Ranch of the National Audubon Society. They have been re-read in June of 2011. Riparian grasslands include large bottomlands of

giant sacaton (Sporobolus Wrightii Monro ex Scribn) on both the Babacomari Ranch (BR) and the Audubon Research Ranch (ARR). Riparian

grassland monitoring stations were established in the fall of 2009, re-read in 2010 and 2011. In addition to this monitoring effort the Babacomari

Ranch has begun to protect private lands along the Babacomari River by selling development rights and placing conservation easements on the

land. The money largely comes from the US Army (Fort Huachuca) buffering military lands used for training and testing. The majority of lands

on the Research Ranch are already protected from development.
USDA Forest Service Proceedings RMRS-P-67. 2013 569

Abstracts/ Resúmenes

Proyecto de protección ribereña en el río Babacomari. El río Babacomari es un afluente importante del río San Pedro en los condados de

Santa Cruz y Cochise, Arizona. Esta cuenca de 56, 656 has incluye pastizales de pendiente suave en la llanura de Sonoita, encinales del área de

Canelo Hills y los bosques de pino-encino del noroeste de la cordillera Huachuca. El río Babacomari recorre 35 km desde su nacimiento cerca

de Sonoita a 1524 m, hacia el este donde se une al San Pedro en Fairbanks, a una altura de 1173 m. El Servicio Geológico de E.U. estima que

este importante afluente aporta anualmente unos 6000 acres-pies de agua alsistema del río San Pedro. El Departamento de Recursos Hidráulicos

de Arizona financió este estudio de 5 años con el proyecto (09-164WPF) en 2009. Los transectos de monitoreo se ubicaron en 2009 y 2010

y continuarán hasta el 2013. Objetivos: Cercar 3.2 km para evitar el acceso del ganado desde el río Babacomari. Seleccionar seis transectos

de monitoreo de vegetación ribereña y geomorfología en el arroyo y seis transectos de vegetación y geomorfología en pastizales de zacatón

ribereños tributarios del río Babacomari. Presentar un reporte con análisis de datos a los propietarios ganaderos para el manejo del área. Estaciones de monitoreo ribereñas se establecieron en mayo de 2010 en tres lugares del río Babacomari antes de la casa del rancho Babacomari.

También se establecieron en tres localidades de la estación biológica Appleton-Whittell de la National Audubon Society, en junio de 2010. El

monitoreo se repitió en junio de 2011. Los pastizales ribereños incluyen grandes áreas de zacatón (Sporobolus wrightii Monro ex Scribn), tanto

en el rancho Babacomari y la propiedad Audubon. Se realizaron monitoreos de pastizales ribereños en el otoño de 2009 y se repitieron en 2010

y 2011. Además de este monitoreo, el rancho Babacomari ha empezado a proteger las tierras privadas del río Babacomari mediante la venta de

derechos y la creación de servidumbres ecológicas. Los fondos provienen principalmente del ejército de E.U. (Fort Huachuca) de conservación

de terrenos militares para entrenamiento y pruebas. La mayor parte del terreno de la estación biológica está protegido de la urbanización.

RORABAUGH, JAMES C.1, Jeffrey M. Servoss2, Valerie L. Boyarski3, Erin C. Fernandez2, Doug Duncan2, Carlos Robles-Elías4, and Kevin E.

Bonine5. 1U.S. Fish and Wildlife Service, Tucson, AZ, retired, jrorabaugh@earthlink.net; 2U.S. Fish and Wildlife Service, Tucson, AZ jeff_servoss@fws.gov, erin_fernandez@fws.gov, doug_duncan@fws.gov; 3

Arizona Game and Fish Department, Phoenix, AZ vboyarski@azgfd.gov; 4

Rancho El Aribabi, Carretera Imuris-Cananea Km 131+900, Imuris, Sonora, MX elaribabi@telcel.blackberry.net.mx; 5Department of Ecology

and Evolutionary Biology and School of Natural Resources, University of Arizona, Tucson, AZ, kebonine@u.arizona.edu.

A comparison of the herpetofaunas of Ranchos Los Fresnos and El Aribabi in Northern Sonora, México. To compare and contrast herpetofaunas at Ranchos Los Fresnos and ElAribabi in northern Sonora, México, we conducted herpetological surveys during 2006-2011, contacted

others working in these two areas, and queried 25 museums for specimens collected at or near these ranchos. Based on this work, nine and seven

amphibian, and 27 and 23 reptile species are known to occur at Ranchos El Aribabi and Los Fresnos, respectively. Significant findings include

at Los Fresnos, presence of presumed Sonora tiger salamanders (Ambystoma mavortium stebbinsi), of which there are only three localities in

México; Arizona treefrog (Hyla wrightorum) at several sites; and Mexican gartersnake (Thamnophis eques); and at El Aribabi, neotropical

whipsnake (Coluber mentovarius), a range extension of 212 km, Tarahumara salamander (Ambystoma rosaceum) near the northern edge of its

range, and Mexican gartersnake. Both ranchos support non-native American bullfrogs (Lithobates catesbeianus) and non-native fishes. Nonnative crayfish (Orconectes virilis) are also present at Rancho Los Fresnos. In total, 46 species of amphibians and reptiles were found, 22 of

which occur on both ranchos. Rancho Los Fresnos supports high grassland species not found at El Aribabi, while a number of typically Sonoran

Desert, foothills thornscrub, and montane species found at El Aribabi were not encountered at Los Fresnos.

Comparación de la herpetofauna de los ranchos Los Fresnos y El Aribabi en el Norte de Sonora, México. Para comparar y contrastar

las herpetofaunas de los ranchos Los Fresnos y El Aribabi en el norte de Sonora, México, realizamos un censo herpetológico de 2006 a 2011,

contactamos a otros que han trabajado en estas dos áreas y consultamos 25 museos sobre especímenes recolectados en o cerca de estos ranchos.

A partir de esta investigación, 9 y 7 especies de anfibios y 27 y 23 de reptiles se reportan de los ranchos El Aribabi y Los Fresnos, respectivamente. Registros importantes incluyen para Los Fresnos, la presencia de las presuntas salamandras tigre de Sonora (Ambystoma mavortium 

stebbinsi), de las cuales sólo hay tres localidades en México; Rana arborícola de Arizona (Hyla wrightorum) en varios sitios, y culebra del agua

(Thamnophis eques); y en El Aribabi, culebra chirrionera neotropical (Coluber mentovarius), una extensión del rango de 212 km, salamandra

tarahumara (Ambystoma rosaceum) cerca del límite norte de su distribución, y culebra del agua. Los dos ranchos tienen la rana toro introducida

(Lithobates catesbeianus) y peces no nativos. El langostino introducido (Orconectes virilis) también está presentes en el rancho Los Fresnos.

En total, se encontraron 46 especies de anfibios y reptiles, de las cuales 22 ocurren en ambos ranchos. El rancho Los Fresnos tiene especies de

pastizales altos que no se encuentran en El Aribabi, mientras una serie de especies típicas del desierto Sonorense, matorral espinoso de piedemonte y especies de montaña que se encuentran en El Aribabi no se encontraron en Los Fresnos.

SALYWON, ANDREW1 and Thomas R. Van Devender2. 1Desert Botanical Garden, 1201 N. Galvin Parkway, Phoenix, Arizona 85008, USA,

asalywon@dbg.org; 2Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717, USA.

The mustard family (Brassicaceae) of the Madrean Archipelago in Sonora, México. The Madrean Archipelago in northeastern Sonora

harbors 37 species and one hybrid of mustards in 25 genera. Lepidium, represented by six species, is the most diverse genus. Thirteen species

are non-native weeds, with several of them considered to be noxious, especially Brassica tournefortii. Recent fieldwork, including expeditions

for the Madrean Archipelago Biodiversity Assessment (MABA) Project, has discovered several new records for Sonora and greatly expanded

our knowledge of the group in this region. Draba viridis, a regional endemic species known only from the Sierra San José, Sonora, and the

Huachuca and Santa Catalina mountains in Arizona, is of conservation concern. Three species known from the Sky Island Region of southern

Arizona that should be looked for in adjacent Sonora are Draba bifurcata, Draba petrophylla, and Pennellia tricornuta. An additional seven

species (5 native) of Brassicaceae occur in the Sierra Madre Occidental (SMO) in eastern Sonora. The center of diversity for these Brassicaceae
570 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

is in the western United States. Unlike many plant groups with tropical affinities, diversity in general declines southward from the northern Sky

Islands Region in Sonora (37 taxa), although the Yécora area in the SMO is moderately diverse (23 taxa).

La familia de las mostazas (Brassicaceae) del Archipiélago Madrense en Sonora, México. El Archipiélago Madrense en el noreste de

Sonora alberga 37 especies y un híbrido de mostazas en 25 géneros. Lepidium, representado por seis especies es el género más diverso. Trece

especies son malezas introducidas y varias se consideran nocivas, especialmente Brassica tournefortii. En trabajo de campo reciente, entre éste

las expediciones del proyecto de Evaluación de la Biodiversidad del Archipiélago Madrense (MABA), se descubrieron varios registros nuevos

para Sonora y aumentó bastante nuestro conocimiento del grupo en esta región. Draba viridis, una especie endémica regional conocida sólo de la

Sierra San José en Sonora y las Sierras Huachuca y Santa Catalina en Arizona se considera como especie en riesgo. Tres especies conocidas de

la Región de las Islas Serranas del sur de Arizona que deben buscarse en el área adyacente de Sonora son: Draba bifurcata, Draba petrophylla y

Pennellia tricornuta. Otras siete especies (5 nativas) de Brassicaceae ocurren en la Sierra Madre Occidental (SMO) del este de Sonora. El centro

de diversidad de estas especies de Brassicaceae es el occidente de Estados Unidos. A diferencia de muchos grupos de plantas con afinidades

tropicales, la diversidad en general disminuye hacia el sur desde el norte de la Región de las Islas Serranas en Sonora (37 taxones); aunque el

área de Yécora en la SMO tiene una diversidad moderada (23 taxones).

SALYWON, ANDREW1, Ronald Tiller2 and Matthew King1. 1Desert Botanical Garden, 1201 N. Galvin Parkway, Phoenix, AZ 85008, asalywon@dbg.org; 2

The Nature Conservancy, 1510 E. Ft. Lowell Road, Tucson, AZ 85719, rtiller@tnc.org.

Analyzing change in ciénega area with aerial photography. Ciénegas are one of the rarest and most endangered habitats in southern Arizona. Historic alteration and destruction has eliminated/reduced the extent of approximately 95 percent of ciénegas in the region. However,

ciénegas are rather poorly studied and quantitative data regarding the area and location of historic and even current ciénegas is lacking. In order

to quantify the current area and perimeters of ciénegas at Las Ciénegas National Conservation Area we are creating ground-truthed shapefiles

with sub-meter accurate GPS units. These shapefiles will provide baseline data for informed monitoring and study of the ciénegas and will

benefit many diverse parties, including hydrologists and ecologists, working from the species-level to the landscape level. The data is necessary to evaluate any future changes in the area resulting from natural processes, climate change or anthropogenic dewatering. Additionally,

the shapefiles are being used to compare to 1930’s Fairchild aerial photography to detect historic changes in both ciénega area and vegetation.

Results and challenges are discussed.

Uso de fotografía aérea para el análisis del cambio en el área de ciénegas. Las ciénegasson uno de los hábitats más escasos y amenazados del

sur de Arizona. Cambios y destrucciones en el pasado han eliminado o reducido aproximadamente 95% de ciénegas en la región. Sin embargo,

las ciénegas son poco estudiadas y se carece de datos cuantitativos sobre el área y ubicación de ciénegas históricas e incluso actuales. Con el fin

de medir el área actual y perímetro de ciénegas en el Área de Conservación Nacional Las Ciénegas, estamos creando shapefiles de verificación

en tierra con unidades de GPS de precisión submétrica. Estos shapefiles proporcionarán información de referencia para monitoreos y estudios

de mejor calidad de ciénegas y será útil a muchos investigadores, incluyendo hidrólogos y ecólogos, y estudios desde nivel de especie hasta el

paisaje. Los datos son necesarios para evaluar cualquier cambio futuro en el área como resultado de procesos naturales, el cambio climático

o sequía antropogénica. Los shapefiles también se están utilizados para hacer una comparación de la fotografía aérea de Fairchild de 1930 y

detectar cambios históricos, tanto en área como en vegetación de la ciénega. Se presentarán resultados y dificultades.

SANDERLIN, JAMIE S.1,2, William M. Block1,3, Joseph L. Ganey1,4, and Jose M. Iniguez1,5. 1Rocky Mountain Research Station, U. S. Forest Service, 2500 S. Pine Knoll Dr., Flagstaff, AZ 86001; 2

jlsanderlin@fs.fed.us; 3wblock@fs.fed.us; 4jganey@fs.fed.us; 5jiniguez@fs.fed.us.

Species richness and avian community dynamics in the Arizona Sky Islands. The Sky Islands in southeastern Arizona contain a unique

and rich avifaunal community located within diverse habitats. The avifauna within these mountains includes many Neotropical migratory

species whose northern breeding range extends to these mountains along with many species typical of similar habitats throughout western

North America and the continent. Understanding ecological factors that influence species richness and biological diversity of both resident and

migratory species is important for conservation of this unique bird assemblage. We use a 5-year data set to evaluate avian species distribution

across a diverse set of montane habitat types within the Santa Rita, Santa Catalina, Huachuca, Chiricahua, and Pinaleño Mountains. Using

point-count data from spring-summer breeding seasons, we describe how avian diversity and community dynamics change through time. We

use Bayesian hierarchical models to describe occupancy as a function of variables like vegetation zone and year, while accounting for variables

that influence detection probability (e.g., species heterogeneity). By identifying important habitat correlates for avian species, these results help

guide management decisions to minimize loss of key habitats and guide rehabilitation and restoration efforts in response to disturbance events,

such as fire, in the Madrean Archipelago.

Riqueza de especies y dinámica de la comunidad de aves en las Islas Serranas de Arizona. Las Islas Serranas, del sureste de Arizona

contienen una comunidad de avifauna diversa y única, localizada en diversos hábitats. La avifauna en estas montañas incluye muchas especies

migratorias neotropicales cuyo límite norte de reproducción se extiende hasta estas montañas, junto con muchas especies típicas de hábitats

similares del oeste norteamericano y el continente. El estudio de los factores ecológicos que influyen en la riqueza de especies y diversidad

biológica tanto de especies residentes como migratorias es importante para la conservación de este ensamble único de pájaros. Utilizamos una

base de datos de 5 años para evaluar la distribución de especies de aves en una asociación de diversos de tipos de hábitat en las montañas Santa

Rita, Santa Catalina, Huachuca, Chiricahua, y Pinaleño. Usando datos de conteo de puntos de las temporadas de reproducción de primaveraverano, describimos cómo la diversidad y las dinámicas de la comunidad aviaria cambian con el tiempo. Usamos modelos jerárquicos Bayes-
USDA Forest Service Proceedings RMRS-P-67. 2013 571

Abstracts/ Resúmenes

ianos para describir la ocupación en función de variables como la zona de vegetación y año, teniendo en cuenta las variables que influyen en la

probabilidad de detección (por ejemplo, la heterogeneidad de las especies). Al identificar importantes correlaciones de hábitat para especies de

aves, estos resultados ayudan a tomar decisiones de manejo para minimizar la pérdida de hábitats clave y guiar los esfuerzos de rehabilitación

y restauración en respuesta a eventos de perturbación, como incendios, en el Archipiélago Madrense.

SARTOR, KARLA, Dave Gori. The Nature Conservancy in New Mexico, 212 E. Marcy St., Suite 200, Santa Fe, NM 87501, ksartor@tnc.

org, dgori@tnc.org.

Arid-grassland mapping as a decision support tool for restoration planning and rare plant survey.We used fine-scalemapping of ecological

sites and states (condition) at two locations in southern New Mexico to identify restoration priorities and effective treatment options, including

grade controlstructures, herbicide application and prescribed burning. The Pitchfork Ranch near Silver City, NM,supports grassland-dependent

species like Aplomado falcon and pronghorn and, embedded within the grassland, a rare cienega wetland. Mapping products at this site are

linked to hydrological models of the watershed to better inform restoration strategies. We have installed rock structures along almost 2km of

severely eroded floodplain grassland in addition to other treatments to prevent further down-cutting and soil loss, and plan to use the site as a

demonstration to increase landowner engagement in restoration elsewhere in the watershed. At a second site near Hachita, NM, ecological sitestate maps willsupport the BLM’s effortsto restore grassland habitat while protecting the rare Chihuahua scurfpea (Pediomelum pentaphyllum),

known from only three populations in southern New Mexico and Arizona. Mapping products and plot-level habitat characterization at known

plant locations inform a species distribution model which will improve the efficiency of locating this ephemeral plant prior to ground-disturbing

activities or herbicide treatments.

Cartografía de pastizales áridos como recurso de apoyo en los programas de restauración y estudio de plantas raras. Utilizamos cartografía a escala fina de sitios ecológicos y sus estados (condiciones) en dos localidades del sur de Nuevo México para identificar prioridades de

restauración y opciones de tratamiento eficaces, incluyendo estructuras para estabilizar pendientes, aplicación de herbicidas y quemas prescritas.

El rancho Pitchfork cerca de Silver City, NM, mantiene especies de pastizales como el halcón aplomado y el berrendo, y dentro de la pradera,

un humedal de ciénega. Los mapas del área están ligados a modelos hidrológicos de la cuenca para ilustrar mejor las estrategias de restauración.

Hemos instalado gaviones en casi 2 km del pastizal de la planicie aluvial severamente erosionado, además de otros tratamientos para prevenir

más pérdida de suelo, y planeamos usar el lugar como sitio demostrativo para aumentar la participación de propietarios en la restauración en

otras partes de la cuenca. En otro sitio cerca de Hachita, NM, mapas ecológicos del área le serán útiles al BLM para restaurar el hábitat de

pastizales, al mismo tiempo que se protege la planta escasa “contrahierba de Chihuahua” (Pediomelum pentaphyllum), conocida sólo de tres

poblaciones en el sur de Nuevo México y Arizona. Los mapas y la caracterización del hábitat a nivel de parcela de las localidades conocidas

actualizan la distribución de la especie y mejoran la eficiencia de la localización de esta planta efímera antes de actividades perturbadoras del

suelo o tratamientos herbicidas.

SÁNCHEZ-ESCALANTE, JOSÉ JESÚS1, Denise Zulema Ávila-Jiménez1, David Alfredo Delgado-Zamora1, Liliana Armenta-Cota1. 1Departamento de Investigaciones Científicas y Tecnológicas (DICTUS), Universidad de Sonora, Blvd. Luis Encinas y Rosales CP 83000 Hermosillo,

Sonora; jsanchez@guayacan.uson.mx.

Vascular plants of El Aribabi Conservation Ranch: plant diversity of a private natural protected area in northern Sonora, México. In

northeastern Sonora, isolated mountains or sky islands are recognized for their high biodiversity, sustaining vegetal communities like desert

scrubs, natural grasslands, oak woodlands and pine-oak forests; within this region (30°51’14.59”N, 110°41’11.91”W and 30°48’29.82”N,

110°32’5.59”W) is located El Aribabi Conservation Ranch. The flora of the ranch is based in more than 1000 herbarium specimens collected

by the Universidad de Sonora herbarium (USON) and with observations records from the MABA (Madrean Archipelago Biodiversity Assessment) database. The flora accounts 454 vascular plants belonging to 87 families and 279 genera. Families with the greatest number of species

are Asteraceae (65), Poaceae (41), Fabaceae (37), Euphorbiaceae (18), Malvaceae (13), Cactaceae (11) and 9 ferns. Only two species of the

flora are nominated with risk category by Norma Oficial Mexicana NOM-059. Non-native plants (27) represent 6 % of the flora and only 6 are

invasive. This work was carried out with financial support from the CONABIO and is important for its contribution to knowledge of the flora

of the northern border, in addition, the ranch was recently declared by the CONANP as a private protected natural area.

Plantas vasculares del Rancho El Aribabi: diversidad vegetal de un área natural protegida privada en el norte de Sonora, México. En

la región noreste del Estado de Sonora, montañas aisladas denominadas “islas serranas” y caracterizadas por una alta biodiversidad, sostienen

comunidades de matorrales xerófilos, pastizales naturales, encinares y bosques de pino-encino; en esta región se localiza el rancho El Aribabi,

con coordenadas 30°51’14.59”N, 110°41’11.91”W y 30°48’29.82”N, 110°32’5.59”W. La flora del rancho está respaldada por más de 1000

ejemplares depositados en el herbario USON y observaciones registradas en la base de datos del proyecto MABA (Madrean Archipelago

Biodiversity Assessment). Se contabilizaron un total de 454 especies de plantas vasculares pertenecientes a 87 familias y 279 géneros. Las

familias con mayor número de especies son Asteraceae (65), Poaceae (41), Fabaceae (37), Euphorbiaceae (18), Malvaceae (13), Cactaceae (11)

y 9 helechos. Sólo dos especies de la flora están nominadas con categoría de riesgo por la Norma Oficial Mexicana NOM-059. Las especies

no nativas (27) representan el 6 % de la flora y sólo 6 son invasoras. Este trabajo se llevó a cabo con el apoyo financiero de la CONABIO y es

importante por su contribución al conocimiento de la flora de la frontera norte; además, el rancho fue declarado recientemente por la CONANP

como área natural protegida privada.
572 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

SCHLICHTING, DEAN1 Jason Mateljak2, and Bianca Gragg3. 1National Park Service, Coronado National Memorial, 4101 East Montezuma

Canyon Road, Hereford, AZ 85615, dean_schlichting@nps.gov; 2National Park Service Chiricahua National Monument, 12856 E. Rhyolite

Creek RD Willcox,AZ 85643, Jason_mateljak@nps.gov; 3National Park Service, Coronado National Memorial, 4101 East Montezuma Canyon

Road, Hereford, AZ 85615, Bianca_torres@nps.gov.

Restoration of Lesser long-nosed bat forage resource: a group effort. In 2007, the construction of the international border fence within

Coronado National Memorial (CORO) resulted in the loss of an estimated 3700 Palmer’s agave (Agave palmeri) plants, which are the primary

food source in Southeastern Arizona of the endangered Lesser long-nosed bat (Leptonycteris curasoae yerbabuenae). A comprehensive mitigation effort was necessary, thus, CORO initiated a collaborative effort between the National Park Service (NPS), US Fish and Wildlife Service,

and the Department of Homeland Security to restore lost agaves. In 2009, CORO contracted with the Natural Resources Conservation Service,

Plant Materials Center in Tucson, Arizona to grow 4500 agave over three years. Each year, beginning in 2010, CORO hosts an annual agave

planting event, involving a coordinated effort between the public and staff from CORO, other NPS units, and other land management agencies.

In 2011, more than 130 volunteers and staff planted 1100 agaves during the one day event. Thus, there are two measures of success for this

project: (1) the replacement of 3700 lost agaves; and (2) the sense of community, wholesome outdoor experience, and feeling of ownership

afforded to all attendees. CORO will host the final annual agave planting event during the summer of 2012.

Restauración de recursos forrajeros del murciélago ‘magueyero chico’: un esfuerzo grupal. En 2007, la construcción del muro fronterizo

en el Memorial Nacional de Coronado (CORO) ocasionó la pérdida de cerca de 3700 plantas de lechuguilla (Agave palmeri), que en el sureste

de Arizona son la fuente primaria de alimento del murciélago magueyero chico (Leptonycteris curasoae yerbabuenae) en peligro de extinción.

Se necesitaba un trabajo intensivo de mitigación, por lo que CORO inició un proyecto colaborativo entre el Servicio de Parques Nacionales

(NPS), Fish and Wildlife Service de EU y el Departamento de Seguridad Nacional para reemplazar las lechuguillas perdidas. En 2009, CORO

firmó un contrato con el Servicio de Conservación de Recursos Naturales, Centro de Plantas en Tucson, Arizona para propagar 4500 agaves en

tres años. Cada año, a partir de 2010, CORO organiza un evento anual de plantación de lechuguillas, que implica un esfuerzo coordinado entre

el sector público y el personal de CORO, otras unidades del servicio de parques nacionales, y otras agencias de manejo de recursos naturales. En

2011, más de 130 voluntarios y personal plantaron 1100 agaves durante el evento de un día. Así, hay dos medidas de éxito para este proyecto:

(1) el reemplazamiento de 3700 lechuguillas perdidas, y (2) el sentido de comunidad, la experiencia saludable al aire libre, y el sentimiento de

propiedad que se ofrece a los asistentes. CORO tendrá el ultimo evento anual de plantación de agaves durante el verano de 2012.

SCHMIDT, JUSTIN O. Southwestern Biological Institute, 1965 W. Brichta Dr, Tucson, AZ 85745, ponerine@dakotacom.net.

The sand dune of Ruby, an anthropogenically created biodiversity hotspot for wasps and their velvet ant parasitoids. Ruby, Arizona,

a historically important mining town in Southern Arizona that deep mined millions of tons of ore, the crushed remains of which after mineral

extraction were piled into a large sand dune. The dune became an ideal nesting habitat for a variety of insects, plus blocked the valley, thereby

generating two lakes. Today the ghost town is an important biodiversity refuge. A large number of species of sand wasps, including cicada

killers, have colonized the dune area, sometimes with world-record-breaking populations. Concomitant with the wasp colonizations were migrations of a variety of species of velvet ant (Hymenoptera: Mutillidae) parasitoids of the wasps. Environmental resource partitioning among

the wasps and their parasitoids in this newly created habitat will be presented and the importance of preservation of various types of land will

be discussed. I thank Howard Frederick of Wildlife World Wide, Cortaro, AZ, for facilitating this study at Ruby and encouraging the research.

La duna de arena de Ruby, un centro de biodiversidad de avispas y sus parásitos las comadritas, creado antropogénicamente. Ruby,

Arizona, es un pueblo minero de importancia histórica en el sur de Arizona, que minó millones de toneladas de mineral subterráneo, donde los

jales, después de la extracción de minerales, se apilaban en una gran duna de arena. La duna se convirtió en un hábitat ideal para la anidación de

una gran variedad de insectos, además de bloquear el valle, lo que generó dos lagos. Actualmente el pueblo fantasma es un refugio importante

de biodiversidad. Un gran número de especies de avispas de arena, incluidas las asesinas de chicharras, ha colonizado la duna, a veces con poblaciones de récord mundial. Simultáneamente con las colonizaciones de avispas ocurrieron las migraciones de una gran variedad de especies de

‘comadritas’(Hymenoptera: Mutillidae) parásitos de las avispas. Se presentará la división de los recursos del medio ambiente entre las avispas

y sus parásitos en este hábitat de nueva creación y se abordará la importancia de la preservación de los distintos tipos de terreno. Agradezco a

Howard Frederick de Wildlife World Wide, Cortaro, AZ por facilitar este estudio en Ruby y fomentar la investigación.

SHALLCROSS, KEITH1 and David Hodges2. Cuenca Los Ojos A.C. Agua Prieta, Sonora Mexico and Cuenca Los Ojos Foundation, 12626 E.

Turkey Creek Road, Pearce AZ 85625, (520) 824-3566; 1yanozha2001@yahoo.com; 2dhodges73@gmail.com.

Camera trapping in the Cajon Bonito Watershed: a synthesis. The Cuenca Los Ojos Foundation, with extensive properties in the northeast

corner of the state of Sonora, has conducted a camera trapping program in Cajon Bonito for two years. The Cajon Bonito is a major interior

drainage and perennial stream in the Sierra San Luis range, flowing north then west into the Rio San Bernardino. Through long-term restoration and water conservation efforts of Cuenca Los Ojos Foundation, Cajon Bonito and surrounding areas now support a vibrant array of native

plant and animal species including the rare ocelot and the second largest population of black bear in Mexico. Our camera trapping project

has provided consistent monitoring and quantification of the wildlife found here, allowing for identification of most mammal species present

including endangered species in close proximity to the US border. In addition to providing information on presence of species, camera trapping

provides important information on local populations of prey species, with spikes or drops in animal populations as signs of possible disease or
USDA Forest Service Proceedings RMRS-P-67. 2013 573

Abstracts/ Resúmenes

poaching. Recently we have incorporated a data collection and processing system developed by Jim Sanderson and Jack Childs that has enabled

interesting comparisons to other study sites in the US and Mexico with the Cajon Bonito site.

Síntesis del trabajo con cámaras trampa en la Cuenca de Cajón Bonito. La Fundación Cuenca Los Ojos, con extensas propiedades en la

esquina noreste del estado de Sonora, ha llevado a cabo un programa de trampeo fotográfico en Cajón Bonito por dos años. El Cajón Bonito

es un drenaje interior importante de corriente perenne en la Sierra San Luis, que fluye al norte y luego al oeste hacia el arroyo San Bernardino.

Mediante trabajos a largo plazo de restauración y conservación del agua de la Fundación Cuenca Los Ojos, el Cajón Bonito y zonas adyacentes

ahora mantienen una gran diversidad de especies de plantas y animales nativos incluyendo el raro ocelote y la segunda población más grande de

oso negro en México. Nuestro proyecto con cámaras trampa ha mantenido un monitoreo constante y censo de la fauna silvestre que se encuentra

aquí, lo que permite la identificación de la mayoría de especies de mamíferos presentes, incluyendo especies amenazadas en las proximidades

de la frontera con E.U. Además de proporcionar información sobre la presencia de especies, el trampeo fotográfico proporciona información

importante sobre las poblaciones de especies presa locales, con picos o caídas en las poblaciones de animales como signos de posibles enfermedades o de caza furtiva. Recientemente Jim Sanderson y Jack Childs diseñaron una base de datos, que ha permitido hacer interesantes

comparaciones entre sitios de estudio en E.U. y México con el Cajón Bonito.

SIDNER,RONNIE1 andDebbieC.Buecher2. 1EcologicalConsulting, 1671NCliftonSt,Tucson,AZ85745,sidner@email.arizona.edu; 2Buecher

Biological Consulting, 7050 E. Katchina Court, Tucson, AZ 85715, dbuecher@comcast.net.

Recent records of Western Yellow Bats (Lasiurus xanthinus) in Arizona. Western yellow bats are insectivorous tree-roosting bats, first

recorded in Arizona in1960. Although found within or near non-native Washingtonia palm trees in Arizona, the bats are otherwise obligate

canyon or riparian dwellers at the northern end of their distribution in southeast Arizona. Hoffmeister’s (1986) ‘Mammals of Arizona’lists only

seven localities with potential information recorded from only 18 individuals. Current status of Lasiurus xanthinus is ‘priority’ or ‘sensitive’

species by interest groups, federal, state, and county government (e.g., in Pima County Sonoran Desert Conservation Plan). From 2002-2008

we captured six L. xanthinus at three new non-urban localities. Arizona Game and Fish HDMS database shows six other new localities; all new

localities are unpublished. Rarely caught in biological surveys, this species is more abundant than expected among bats sent to Arizona State

Health Department in Tucson for rabies testing. Between 2000-2007 we identified >40 western yellow bats from five sites (including three new

localities). These yellow bats were taken each month from April through November, with equal numbers of males and females, and adults and

young. Clearly this is another example of a southern species moving northward.

Registros recientes de murciélagos amarillos occidentales (Lasiurus xanthinus) en Arizona. Los murciélagos amarillos occidentales son

murciélagos insectívoros que usan los árboles como refugio; se registraron por primera vez en Arizona in 1960. Aunque se les encuentra en o

cerca de palmeras (Washingtonia introducidas en Arizona), los murciélagos son habitantes ribereños o asociados a cañones en el extremo norte

de su distribución en el sureste de Arizona. ‘Mamíferos de Arizona’ de Hoffmeister (1986) enlista tan sólo siete localidades con información

potencial registrada de sólo 18 individuos. El estatus de riesgo de Lasiurus xanthinus en E.U. es similar a ‘especie rara’ o ‘sujeta a protección

especial’ para el gobierno federal, estatal y de condado (por ejemplo, el Plan de Conservación del Desierto Sonorense del Condado de Pima).

De 2002 a 2008 se capturaron seis L. xanthinus en tres nuevas localidades no urbanas. La base de datos HDMS de Arizona Game and Fish

incluye otras seis localidades nuevas; todas las localidades nuevas no están publicadas. Rara vez atrapada en investigaciones biológicas, esta

especie es más abundante de lo esperado entre murciélagos enviados al Departamento de Salud del Estado de Arizona en Tucson para pruebas

de rabia. Entre 2000-2007 se identificaron >40 murciélagos amarillos occidentales en cinco sitios (incluyendo tres nuevas localidades). Estos

murciélagos amarillos fueron colectados cada mes desde abril hasta noviembre, con igual número de machos y hembras, y adultos y jóvenes.

Es evidente que este es otro ejemplo de una especie del sur moviéndose hacia el norte.

SIERRA-CORONA, RODRIGO1, Humberto Suzan2, Hugo Luna-Soria2, Ed L. Fredrickson3, and Gerardo Ceballos4. 1Instituto de Ecología,

Circuito Exterior s/n, anexo al Jardín Botánico, UNAM, México DF, 04510, lomogrande@hotmail.com; 2Universidad Autónoma de Querétaro,

Querétaro México, 76230, hsuzan@uaq.mx, hugo.lunas@yahoo.com.mx; 3Department ofAgriculture, Eastern Kentucky University, Richmond

KY 40475; 4Instituto de Ecología, Circuito Exterior s/n, anexo al Jardín Botánico, UNAM, México DF, 04510, gceballo@miranda.ecologia.

unam.mx.

Prairie dogs and cattle interactions: Shifting from competition to a grazing association. Since the mid nineteenth century, prairie dogs were

considered a pest by the inhabitants of rural communities through their historical range mainly because they were presumed to compete with

cattle for available forage resources; a belief reinforced by the eradication campaigns implemented from the early 1900’s to date. Nowadays,

prairie dogs occupy less than 2% of their historical distribution, and vertebrate associated species are affected, like black-footed ferret –which

is nearly extinct. Despite the intensity of the eradication campaigns, no scientific data support the competition-only interaction claims, it’s based

only in social perceptions. This research addresses the question if prairie dogs and cattle interactions could be mutualistic, similar to those observed between American bison and prairie dogs. Beef cattle (Bos taurus) habitat selection in relation with prairie dog presence was measured

on a short grass ecosystem within the Janos Biosphere Reserve. Analyses of cattle movements show a direct preference for prairie dog colonies,

with a strong preference to forage in the margins. From these observations we conclude that cattle-prairie dog interactionsresemble bison-prairie

dog grazing associations, opening a new discussion about future management strategies for the North American grasslands.
574 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Perritos de la pradera e interacciones con el ganado: De la competencia a una asociación de pastoreo. Desde mediados del siglo XIX,

los perritos de la pradera fueron considerados una plaga por los habitantes de las comunidades rurales en su área de distribución histórica,

principalmente porque se presumía que competían con el ganado por recursos forrajeros. Esta creencia fue reforzada por las campañas de

erradicación implementadas desde principios de 1900 hasta la fecha. Actualmente, los perritos de la pradera ocupan menos del 2% de su distribución histórica, y especies de vertebrados asociados se ven afectadas, como el hurón de patas negras, que está casi extinto. A pesar de la

intensidad de las campañas de erradicación, no hay datos científicos que apoyen las pretensiones de interacción únicamente competitiva, se

basan solamente en percepciones sociales. Esta investigación aborda la cuestión de si los perros de las praderas y las interacciones con ganado

pudiesen ser mutualistas, similares a las observadas entre el bisonte americano y los perros de la pradera. La selección de hábitat del ganado

vacuno (Bos taurus) en relación con la presencia de perros de la pradera se midió en un ecosistema de pastos cortos dentro de la Reserva de

la Biosfera de Janos. Los análisis del movimientos de ganado muestran una preferencia directa por colonias de perritos de la pradera, con una

fuerte preferencia por el forraje en los márgenes. A partir de estas observaciones se concluye que las interacciones entre ganado y perritos de

la pradera, se parecen a las asociaciones de pastoreo entre bisontes y perritos de la pradera, lo que abre una nueva discusión sobre las futuras

estrategias de manejo de praderas de América del Norte.

SIMMS, KAREN1, Gita Bodner2. 1 Bureau of Land Management, 12661 East Broadway, Tucson, AZ 85748, Ksimms@blm.gov; 2The Nature

Conservancy in Arizona, 1510 E Ft. Lowell Road, Tucson, AZ 85719, gbodner@tnc.org .

Moving from adaptive management to climate adaptation: experiences from agency staff and stakeholders at the Las Ciénegas National 

Conservation Area. Advocates of climate adaptation often cite adaptive management as a crucial tool for helping natural systems cope with

the impacts of climate change. Yet this tool is challenging to implement. Las Ciénegas has pursued adaptive management for some 15 years,

and is now sharing lessons on what has worked well and what has not. The site’s approach allows BLM and regional stakeholders to monitor

and evaluate ecosystem changes, and adjust management actions based on what they learn. This process builds the knowledge, flexibility and

community support needed to manage lands in a changing climate. Yet the partnership has recognized that in some cases, making adjustments

to existing activities may not be enough to buffer the watersheds from effects of rapid change. BLM and partners are examining how to further

incorporate climate adaptation by modifying monitoring protocols; implementing no regrets actions; and identifying areas where typical actions may be ineffective or even counterproductive. Current no-regrets actions include boosting the capacity of floodplains to capture sediment

and slow release of runoff water. Scenario planning activities are now helping the group identify what might need to be done quite differently

depending on how climate shifts and ecosystems respond, and on how agency resources change.

Del manejo adaptativo a la adaptación al clima: experiencias del personal de agencia y sectores interesados del Área de Conservación 

Nacional Las Ciénegas. Defensores de la adaptación al clima citan frecuentemente el manejo adaptativo como requisito para ayudar a los

sistemas naturales a superar los impactos del cambio climático. Pero este requerimiento es difícil de implementar. Las Ciénegas ha efectuado

el manejo adaptativo por unos 15 años, y ahora comparte experiencias de lo que ha funcionado y lo que no. El enfoque del sitio permite a la

Oficina de Administración de Tierras (BLM por sus siglas en inglés) y los sectores regionales interesados monitorear y evaluar los cambios del

ecosistema y ajustar el manejo según lo aprendido. Este proceso aumenta el conocimiento, la flexibilidad y el apoyo de la comunidad necesario

para el manejo de recursos naturales en un clima cambiante. Pero la asociación ha reconocido que en algunos casos, el hacer cambios en las

actividades actuales puede ser insuficiente para amortiguar los efectos del cambio rápido. BLM y socios están analizando cómo incorporar aún

más el cambio climático al modificar los protocolos del monitoreo; implementar acciones sin remordimientos; e identificar las áreas donde

las acciones típicas son inefectivas o incluso contraproducentes. Acciones sin remordimientos incluyen aumentar la capacidad de las planicies

aluviales de capturar el sedimento y disminuir el escape de escorrentía. El programar para escenarios está ayudando al grupo a identificar que

se necesita modificar dependiendo de como cambia el clima y de como responden los ecosistemas, y como cambian los recursos de la agencia.

SIVINSKI. ROBERT C. Museum of Southwestern Biology - Herbarium, MSC03 2020, University of New Mexico, Albuquerque, NM, USA

87131-0001.

Aridland spring ciénegas: Rare southwestern habitats for endangered plants. Ciénega refugia for rare plants are medium to low elevation

wet meadows created by stable springs and seeps in arid regions. A comprehensive inventory of southwestern ciénegas has not been completed;

however, these habitats are clearly rare, diminishing in extent, or already eliminated. Groundwater depletion, erosion, conversion to agriculture

or aquaculture, abusive grazing, and exotic weeds threaten most remaining ciénega habitats and the plants and animals in them. Several southwestern plant species are confined to these habitats, making aridland spring ciénegas distinct from other riverine or lentic wetlands in the region.

Some ciénega plants have widespread distributions, but since their habitat is rare these plants are also rare and, in some cases, endangered.

Ciénegas de aguajes en zonas áridas: hábitats escasos para especies en peligro en el Suroeste Norteamericano. Las ciénegas son refugio

para especies de plantas escasas y praderas húmedas de baja elevación creadas por manantiales e infiltraciones estables en regiones áridas. No

se ha hecho un inventario completo de las ciénegas del suroeste; sin embargo es evidente que estos hábitats son escasos, y su extensión está

disminuyendo o se ha eliminado. La disminución de los mantos acuíferos, erosión, cambio de uso de suelo hacia agricultura o acuacultura,

sobrepastoreo, y especies invasoras amenazan la mayoría de las ciénegas que quedan, así como su flora y fauna. Varias especies de plantas del

suroeste están restringidas a estos hábitats, distinguiendo a las ciénegas de zonas áridas de otros humedales ribereños o lenticos en esta región.

Algunas plantas de ciénegas tienen una amplia distribución, sin embargo como sus hábitats son escasos las plantas también son escasas, y en

muchos casos en peligro de extinción.
USDA Forest Service Proceedings RMRS-P-67. 2013 575

Abstracts/ Resúmenes

SMITH, STEVEN E.1, Ma. Guadalupe Mendoza,2 G. Zúñiga,2 Kandres Halbrook,1 J. L. Hayes,3 and D. N. Byrne4. 1 School of Natural Resources

and the Environment, University of Arizona, P.O. Box 210043, Tucson, AZ 85721, azalfalf@ag.arizona.edu; 2Escuela Nacional de Ciencias

Biológicas-IPN, Prol. de Carpio y Plan de Ayala s/n, Casco de Santo Tomás, México D.F., C.P. 11340, México, marnayi55@yahoo.com.mx; 3

USDA Forest Service, Pacific Southwest Research Station, P.O. Box 245, Berkeley, CA 94701; 4Department of Entomology, University of

Arizona, P.O. Box 210036, Tucson, AZ 85721.

Modeling current and future suitable habitat for a seedling-attacking bark beetle and its pine hosts in the Madrean Archipelago. Dendroctonus rhizophagus is a seedling-attacking bark beetle that occurs on pines in the Sierra Huachinera and may broadly impact pine establishment

elsewhere in the Madrean Archipelago. Little is known regarding habitat requirements of D. rhizophagus and this complicates monitoring. We

used known occurrence records to develop contemporary climate-based models to describe areas of suitable habitat for D. rhizophagus and six

of its Pinus hosts throughout the Archipelago. These models were also projected to four predicted climate scenarios for the period 2040-2069.

More than 2400 km2 of suitable habitat for both D. rhizophagus and at least one host taxon are predicted to currently occur in the Archipelago

north and west of known insect occurrences. The area predicted to be suitable for D. rhizophagus declines under two of four future climate

scenarios and increases in the other two scenarios. Suitable habitat for P. leiophylla was predicted to increase under all future climate scenarios.

Area of predicted suitable habitat generally decreases for P. engelmannii and P. strobiformis in all scenarios. Many instances were observed

of unprecedented habitats for certain Pinus taxa under future climate scenarios. This suggests opportunities for assisted migration to maintain

functioning forest ecosystems.

Modelado de hábitats actuales y futuros del descortezador de plántulas de pino y sus hospederos en el Archipiélago Madrense. El descortezador Dendroctonus rhizophagus ataca plántulas de pino en la Sierra Huachinera y puede afectar el establecimiento de pinos a lo largo del

Archipiélago Madrense. Se conoce poco sobre el requerimiento de hábitat de D. rhizophagus por lo tanto se complica su monitoreo. Empleamos

registros de incidencia para desarrollar un modelo basado en clima contemporáneo para describir los hábitats adecuados para D. rhizophagus y

seis Pinus hospederos a lo largo y ancho delArchipiélago. Estos modelosfueron proyectados para predecir 4 escenarios climáticos para el periodo

2040-2069. Más de 2400 km2 de hábitat adecuado tanto para D. rhizophagus y por lo menos un taxón hospedero se predice que actualmente se

distribuye al norte y oeste de la localidad conocida del insecto. El área pronosticada de distribución de D. rhizophagus disminuye en dos de los

cuatro escenarios de cambio climático y aumenta en los otros dos. Se predice que la distribución de P. leiophylla aumentará según los cuatro

escenarios de cambio climático. En general el hábitat de P. engelmannii y P. strobiformis disminuye en todos los escenarios. En varios casos

se observaron hábitats sin precedentes para ciertos taxones de Pinus bajo futuras condiciones de cambio climático. Esto sugiere oportunidades

para migración asistida para mantener el funcionamiento del ecosistema de bosques.

SOLES, ELLEN1 and Martha S. Cooper 2. 1PO Box 420, Cliff, NM 88028, Ellen.Soles@nau.edu; 2 The Nature Conservancy, PO Box 1603,

Silver City, NM 88062, mschumann@tnc.org.

Long-term hydrological and geomorphological monitoring, Gila River, New Mexico. Persistence of native fishes and low richness and

abundance of nonnative fishes in the Cliff-Gila Valley reach is partially attributable to maintenance of a natural flow regime and thus quantity

and quality of aquatic habitats. Mesohabitats found within and near the river are formed by the dynamic interactions among channel form, vegetation, and flooding over time. Frequent, smaller floods play a crucial role in re-wetting the floodplain, increasing alluvial groundwater levels,

and sustaining backwater wetland zones. Larger floods reconfigure extensive lengths of the active and overflow channel zones, backwaters, and

floodplains. The purpose of this study is to describe channel, floodplain, and terrace morphology; floodplain surface deposits and channel bed

material; and vegetation and mesohabitat classificationsin the Cliff-Gila Valley. Hourly alluvial groundwater elevations on selected transects and

surface flow stage provides a means of evaluating interactions between water availability and habitat creation and persistence. Long-term study

results will improve our understanding of how the flow regime of southwestern rivers—including both moderate and large flood events—create

and sustain a diversity of river-associated habitats, including in-channel fish habitat, wetland zones, and riparian grasslands and tree canopy.

Monitoreo a largo plazo de la geomorfología e hidrología del río Gila, Nuevo México. La persistencia de peces nativos y la poca abundancia

de peces exóticos en el tramo del río en el valle de Cliff-Gila se debe en parte a que mantiene el caudal natural y por consiguiente la calidad y

cantidad de hábitats acuáticos. Las interacciones dinámicas entre la vegetación, las inundaciones y las formas que toma el cauce, con el tiempo

crean los hábitats húmedos del río y sus alrededores. Las inundaciones menores y frecuentes son decisivas para regar los terrenos aluviales,

aumentar los niveles aluviales del acuífero y sostener las zonas de humedales del remanso. Las inundaciones grandes modifican largos tramos

del cauce activo y de las zonas de desbordamiento, remanso y terrenos aluviales. El propósito de este estudio es describir la morfología del

cauce, la zona de inundación y las terrazas fluviales; sedimentos de la zona de inundación y caracterización de los materiales del lecho del río; y

describir la vegetación y el hábitat húmedo del valle de Cliff-Gila. El nivel del acuífero aluvial por hora en transectosseleccionados y el volumen

de la superficie ayudan a evaluar las interacciones del agua disponible y la creación del hábitat y su persistencia. Los resultados de estudios a

largo plazo mejorarán nuestro conocimiento de como el régimen fluvial de los ríos del suroeste de E.U. (caudales grandes o moderados) crean

y sostienen una diversidad de hábitats asociados al río: hábitats para peces, humedales, pastizales ribereños y el dosel de los árboles.

SOLIZ-GARZA,GILBERTO,AgustínGómez-Alvarez, andArturoVillaba-Atondo.Departamento de InvestigacionesCientíficas yTecnológicas

de la Universidad de Sonora, Blvd. Luis Encinas y Rosales S/N, Col. Centro, Hermosillo, Sonora, México gsolis@guayacan.uson.mx.

Monitoring and conservation efforts along the Santa Cruz River in Sonora, Mexico. The Santa Cruz River basin is very important economically, ecologically, and socially as the primary source of drinking water for the border cities of Nogales, Sonora, Mexico and Nogales, Arizona,
576 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

United States, referred to as “Ambos Nogales”. In recent decades, rapid population growth and man’s desire to satisfy their basic needs, have

led to changes in the surrounding environment, such as overexploitation of the aquifers and loss of biodiversity of flora and fauna present in

these water bodies. Aware of the need for information and knowledge on the main channel of the Santa Cruz River, monitoring was conducted

on surface water quality, flora, and vegetation. Research results from various periods over the past 15 years will be presented.

Esfuerzos de monitoreo y conservación del río Santa Cruz en Sonora, México. La cuenca del río Santa Cruz (RSC) representa una gran

importancia económica, ecológica y social porser la principal fuente de abastecimiento de agua potable para las ciudadesfronterizas de Nogales

Sonora, México y de Nogales Arizona, Estados Unidos, referidas como “Ambos Nogales”. En las últimas décadas, el acelerado crecimiento

poblacional y el deseo del hombre por satisfacer sus necesidades primordiales, han provocado cambios en el ambiente que lo rodea, repercutiendo en la sobreexplotación de los mantos acuíferos y en la pérdida de la biodiversidad de la flora y fauna presente en esos cuerpos de agua.

Consientes de la necesidad de obtener información y conocimiento sobre el cauce principal del RSC se realizaron esfuerzos de monitoreo sobre

calidad del agua superficial y flora y vegetación. Se presentarán resultados de investigación obtenidos en varios períodos durante 15 años.

SOLIS-GRACIA, VERONICA1, Rurik List2, Enrique Martínez-Meyer3, and Gerardo Ceballos1

.

1

Instituto de Ecología-Universidad Nacional

AutónomadeMéxico,A.P.70-275CiudadUniversitaria,Coyoacán04510México,D.F.,México,veronicasolisg@gmail.com,gceballo@miranda.

ecologia.unam.mx; 2Departamento de Ciencias Ambientales, Universidad Autónoma Metropolitana-Lerma, Hidalgo Pte. 46, Col. La Estación,

Lerma, Estado de México 52006, México, r.list@correo.ler.uam.mx; 3Instituto de Biología-Universidad Nacional Autónoma de México, A.P.

70-275 Ciudad Universitaria, Coyoacán 04510 México, D.F., México, emm@ibiologia.unam.mx .

Identification of suitable areas for reintroduction of bison (Bison bison) in Chihuahua, Mexico. Bison (Bison bison) once ranged from

Alaska to Northern Mexico, but between 1830 and 1880 were nearly driven to extinction, and with it, their ecological role in the NorthAmerican

grasslands was lost. Efforts are underway in Canada, the United States, and more recently in Mexico, to recover the ecological role of the bison.

Using data from aerial surveys of the only free ranging bison herd in Mexico (which moves between Janos, Chihuahua and Hidalgo county,

New Mexico) and predictive models (GARP and MAXENT), we identified an area suitable for bison within the Janos Biosphere Reserve in

Chihuahua. Bison from Wind Cave National Park in South Dakota were translocated there in order to form a breeding herd for the production

of bison for reintroduction in other suitable sites within the former range of the species, as part of both grassland and bison conservation efforts.

Identificación de áreas propicias para la reintroducción del bisonte (Bison bison) en Chihuahua, México. La distribución histórica del

bisonte (Bison bison) se extendía desde Alaska hasta el norte de México, pero de 1830 a 1880 casi se extinguió y perdió su papel ecológico en

los pastizales de Norteamérica. Se está trabajando tanto en Canadá, Estados Unidos y recientemente en México para recuperar el papel ecológico

del bisonte. Utilizando datos de monitoreos aéreos de la única manada libre en México (que se mueve en la zona fronteriza entre Janos, Chihuahua y el condado de Hidalgo, Nuevo México) y modelos predictivos (GARP y MAXENT) se identificó un área propicia para bisonte dentro

de la Reserva de la Biósfera Janos, en Chihuahua. Se trasladó a esta zona un grupo de bisontes procedentes de Wind Cave National Park, en

Dakota del Sur, con el propósito de criar un grupo reproductor para posteriormente introducir más individuos a otras áreas apropiadas dentro

de su distribución histórica; en un esfuerzo de conservación tanto del bisonte como de los pastizales.

SPRINGER, ADAM C., Don E. Swann, and Chris E. Pruden. Saguaro National Park, 3693 S. Old Spanish Trail, Tucson, AZ 85730, Adam_

Springer@nps.gov, Don_Swann@nps.gov.

The role of precipitation in the distribution of the saguaro cactus at its upper elevation limit in Saguaro National Park. This presentation summarizes the results from a study of the saguaro population in its upper elevation range at Saguaro National Park from 3,500 to 5,500

feet. PRISM data indicates that the 30-year average annual precipitation for this elevation zone ranges from 400 to 600 mm, but declines of

10 percent are predicted by the end of the century. Using logistic regression, we model saguaro presence with data from 120 (0.25 ha) plots

located at these elevations. From predictive variables of elevation, slope, fire history, solar insolation, and 30-year average annual precipitation

this model predicts 70 percent of the variation in current saguaro presence at these elevations. The model indicates that when altering annual

precipitation to reflect projected declines by the end of the century, these reductions in precipitation will have detrimental effects.

Efecto de la precipitación en la distribución de sahuaros en su límite de distribución más alto en el área protegida Saguaro National Park. 

Se presentan los resultados de un estudio de la población de sahuaro en su límite de distribución más alto, 1066 a 1676 m, en Saguaro National

Park. La información de PRISM indica que la precipitación media anual en un periodo de 30 años para esta elevación varia de 400 a 600 mm,

pero se estima una disminución del 10 % a finales del siglo. Usando regresión logística, se modelo la presencia de sahuaro en 120(0.25 ha.)

cuadrantes localizados en esta elevación. Con variables predictivas de elevación, pendiente, historia de incendios, radiación solar, precipitación

media anual de 30 años, este modelo predice 70 % de variación en la presencia de sahuaro a esta elevación. El modelo indica que al alterar la

precipitación anual, para reflejar la disminución estimada para finales de siglo, esta reducción en lluvias tendrá efectos negativos.

SREDL, MICHAEL J. Arizona Game and Fish Department, 5000 W. Carefree Hwy., Phoenix, AZ 88086, msredl@azgfd.gov.

Calibrating our progress towards recovery of amphibian populations: an area-based approach and occupancy modeling. Like many

amphibian species worldwide, the Chiricahua leopard frog (Rana chiricahuensis) experienced a dramatic, range wide decline during the past

three decades and was listed under the Endangered Species Act (ESA) as threatened in 2002. A species recovery plan was finalized in 2007
USDA Forest Service Proceedings RMRS-P-67. 2013 577

Abstracts/ Resúmenes

that included four recovery criteria that, when reached, will have: 1) established sufficient populations and metapopulations, 2) managed the

necessary aquatic breeding habitats, 3) managed important dispersal corridors, and 4) reduced threats so that the Chiricahua leopard frog no

longer needs the protection of the ESA. Although great progress has been made since federal listing, progress on recovery criterion 1 has been

hampered by 1) the dearth of suitably configured landscapes that could “host” candidate metapopulations and 2) the difficulty of establishing

and monitoring stable and viable metapopulations given the limited human and financial resources available. I develop a conceptual area-based

approach to calibrate progress toward recovery that is applicable to the Chiricahua leopard frog that utilizes occupancy modeling to gauge

progress in establishing, managing, and monitoring viable metapopulations. This approach is easier to design and implement, makes fewer

assumptions, and is less biased than the current “strict metapopulation” approach and is applicable to other patchily distributed amphibians.

Medición del progreso en la recuperación de poblaciones de anfibios: enfoque basado en área y modelado de límites de ocupación. La

rana de Chiricahua (Rana chiricahuensis) como muchos anfibios del mundo, durante las últimas tres décadas, sufrió una disminución drástica

de su población, por lo que en 2002 se catalogó como especie amenazada en la Ley de Especies en Peligro de Extinción de E.U. (ESA, siglas

en inglés). Así, en 2007 se terminó un programa de acción para la conservación de la especie con cuatro metas a lograr 1) establecimiento de

poblaciones y metapoblaciones, 2) manejo de hábitats acuáticos para su reproducción, 3) manejo de corredores ecológicos para su dispersión,

4) reducción de amenazas de tal forma que la rana de Chiricahua salga de la lista de ESA. A pesar del progreso obtenido desde que se encuentra

en la lista ESA, el progreso de recuperación del criterio 1 se ha visto entorpecido por 1) la escasez de paisajes adecuados para facilitar metapoblaciones y 2) la dificultad de establecer y monitorear metapoblaciones ya que existe un limitado recurso humano y financiero. Desarrollé un

enfoque conceptual basado en área para medir el progreso de recuperación que se puede aplicar a la rana de Chiricahua y emplea un modelado

de ocupación de espacio para medir el progreso de establecimiento, manejo, y monitoreo de metapoblaciones viables. Este enfoque es más fácil

de diseñar e implementar, requiere menos suposiciones, y tiene menos sesgo que el modelo actual denominado de “metapoblaciones estrictas”,

y se puede aplicar a otros anfibios con distribución fragmentada.

STETSON, CHRISTOPHER. Coronado National Forest, 300 W. Congress, Tucson, AZ 85701, chstetson@fs.fed.us.

Factors leading to landscape-scale wildfires on the Coronado National Forest in 2011, with a summary of the affected areas. The Coronado National Forest experienced a record fire season, as far as fire size, in 2011 with over 300,000 acres impacted by wildfire. Several factors

came into alignment that ultimately led to the landscape-scale fire events that affected not only the “Sky Islands” of the Coronado but adjacent

land ownerships as well. The combination of moisture stressed vegetation, seasonal weather events, topography, and the frequent occurrence

of ignitions influenced the scale and severity of the fires across the southern half of the Coronado. The Horseshoe 2 fire covered 222, 954

acres, in the Chiricahua Ecosystem Management Area (EMA). Fires on the western portion of the Coronado impacted over 92,000 acres on the

Tumacacori, Huachuca, and Santa Rita EMA’s. A considerable amount of land was affected by the fires of 2011. However, variable levels of

soil burn severity occurred creating a “mosaic” of burned and unburned areas across the landscape. Although some areas require rehabilitation, other areas benefited from the wildfires creating favorable conditions that managers have the opportunity to maintain into the future. This

presentation will provide an overview of the factors that “lined up” to provide the recipe for one of the largest wildfire seasons in Coronado

National Forest history and the impacts of the fires on the landscape.

Factores causantes de incendios forestales a escala de paisaje en el Bosque Nacional de Coronado en 2011, con un resumen de las zonas afectadas. El Bosque Nacional de Coronado tuvo una temporada de incendios récord, en lo que a tamaño se refiere, en 2011, con más de

122.000 hectáreas afectadas por incendios forestales. Varias causas coincidieron y finalmente llevaron a incendios del paisaje que afectaron no

sólo a las “Islas Serranas” del Coronado, sino también terrenos adyacentes privados. La combinación de vegetación estresada por la sequía,

eventos climáticos estaciónales, topografía, y la ocurrencia frecuente de igniciones influyó en la magnitud y gravedad de los incendios de la

mitad sur del Coronado. El incendio La Herradura 2 cubrió 90 226 hectáreas en el Área de Manejo de Ecosistemas del Chiricahua (EMA). Los

incendios en la parte occidental del Coronado impactaron más de 37 231 hectáreas en Tumacácori, Huachuca y Santa Rita del EMA. Un área

considerable se vio afectada por los incendios de 2011. Sin embargo, se registraron variaciones en la severidad del suelo quemado, creando

un “mosaico” de áreas quemadas y sin quemar. Aunque algunas áreas necesitan restauración, otras se beneficiaron de los incendios, creando

condiciones favorables que los encargados de manejo pueden mantener. Presentamos un resumen de los factores presentes para crear la receta

de una de las mayores temporadas de incendios en la historia del Bosque Nacional de Coronado y sus impactos.

STROMBERG, J. C.1, D. Setaro1, J. Sabo1, E. Moody1, T. Meixner2, H. Nicholas2, K. Lohse3. 1Arizona State University, School of Life Sciences, Tempe AZ 85287-4501, jstrom@asu.edu; 2

University of Arizona, Dept. of Hydrology and Water Resources, Tucson AZ 85721; 3Idaho

State University, Pocatello ID 83209.

Linkages between stream hydrology, vegetation, and soil arthropods in the semi-arid highlands ecoregion of Arizona. In many streams

of the Huachuca Mountains of southern Arizona, the surface flow is ephemeral or intermittent. Despite the lack of flow permanence, drainages

there support highly diverse ecosystems. A better understanding of the processes that support such diversity might allow for more informed

decision-making and management practices. The following questions were asked to increase our knowledge about the processes controlling

species diversity: 1) Does an increase in flow permanence correlate to increased species richness for ground-dwelling arthropods and the riparian plant community? 2) Is placement within a stream network a controlling factor for species richness? Preliminary results of this multi-year

study indicate that high water availability in canyons, present as surface and sub-surface flows, sustains high richness and abundance of woody

plant species with large canopies; this, in turn, limits the number of herbaceous species due to shading and leaf litter. Ground-dwelling arthro-
578 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

pods, however, reach their greatest diversity in these dense riparian forests. The water-limited foothills streams and lower canyon sections have

more open canopies, but support a larger number of herbaceous species. Each stream type supports different (but overlapping) communities of

arthropods and plants, with the combination of different stream types thus collectively supporting high diversity.

Vínculos entre hidrología de arroyos, vegetación y artrópodos del suelo en la ecorregión del altiplano semiárido de Arizona. En muchos

arroyos de las montañas Huachuca delsur deArizona, la corriente superficial es efímera o intermitente.Apesar de la falta de corrientes continuas,

los drenajes mantienen ecosistemas altamente diversos. Una comprensión mejor de los procesos que mantienen esa diversidad podría resultar

en una toma de decisiones y de manejo con mejores bases. Se formularon las siguientes preguntas para aumentar nuestro conocimiento sobre

los procesos que controlan la diversidad de especies: 1) ¿Un aumento en la permanencia del flujo se correlaciona con aumento de la riqueza de

especies de artrópodos terrestres y la comunidad vegetal ribereña? 2) ¿Es la ubicación en una red de corrientes un factor de control de la riqueza

de especies? Los resultados preliminares de este estudio de varios años indican que la alta disponibilidad de agua en los cañones, presente en

la superficie y las corrientes del subsuelo, mantiene una alta diversidad y abundancia de especies de plantas leñosas de dosel grande, lo que, a

su vez, limita la cantidad de especies herbáceas, debido a la sombra y la hojarasca. Sin embargo, los artrópodos terrestres alcanzan su mayor

diversidad en estos densos bosques ribereños. Los arroyos del piedemonte y las áreas más bajas de cañones con agua limitada tienen un dosel

más abierto, pero mantienen un mayor número de especies herbáceas. Cada tipo de arroyo mantiene diferentes(pero superpuestas) comunidades

de artrópodos y plantas, por lo tanto la combinación de diferentes tipos de corrientes mantiene colectivamente una gran diversidad.

STROMBERG, J. C.1, M. S. White1, G. L. Katz2. 1 Arizona State University, School of Life Sciences, Tempe AZ 85287-4501, jstrom@asu.edu; 2

Appalachian State University, Dept. of Geography and Planning, Boone NC 28608

Riparian vegetation and disappearing groundwater. Over the past century in the southwestern United States, rapid economic growth, expanding urban centers and agriculture have driven steep increases in freshwater demands, which have been met through groundwater pumping,

surface flow diversions, and dams - all of which alter water availability and flow patterns in rivers. These shifting baseline conditions, combined

with episodic drought, have led both to the drying of river reaches that were once perennial as well as to the wetting of previously dry river segments. Drawing from case studies of rivers in southern Arizona including Cienega Creek, Garden Canyon, the San Pedro River, and the Santa

Cruz River, we explore several scenarios: 1) the biological changes that occur in groundwater-dependent ecosystems as water tables deepen

and no-flow periods increase, 2) the patterns and rates of vegetation change that occur when diversion and pumping pressures are reduced; and

3) the response of riparian vegetation to discharge of municipal effluent into stream channels.

Vegetación ribereña y desaparición de aguas subterráneas. Desde el siglo pasado en el suroeste de Estados Unidos, el rápido crecimiento

económico, la expansión de los centros urbanos y la agricultura han aumentado la demanda de agua dulce, que se ha resuelto a través de bombeo

del acuífero, desviaciones de corrientes en la superficie y presas, todo lo cual altera la disponibilidad de agua y los patrones de las corrientes de

ríos. El cambio de estas condiciones básicas, combinado con la sequía, han llevado tanto al secado de tramos de ríos que antes eran perennes,

y también ha causado que tramos anteriormente secos tengan agua. A partir de estudios de caso de ríos en el sur de Arizona, incluyendo los

ríos Ciénega Creek, Garden Canyon, San Pedro, y Santa Cruz, exploramos varios escenarios: 1) los cambios biológicos que ocurren en los

ecosistemas que dependen de aguas subterráneas, conforme baja el nivel freático y aumentan los períodos sin corriente, 2) los patrones e índices

de cambio de vegetación que ocurren cuando se reduce el estrés por la desviación y el bombeo, y 3) la respuesta de la vegetación ribereña a la

descarga de aguas residuales en los cauces de arroyos..

SUPPLEE, VASHTI (TICE)1 and Jennie MacFarland2. 1Arizona Important BirdArea Program,AudubonArizona, 3131 S. CentralAve., Phoenix,AZ

85040, tsupplee@audubon.org; 2Arizona Important Bird Area Program, Tucson Audubon Society, 300 E. University Blvd, #120, Tucson, AZ

85705, jmacfarland@tucsonaudubon.org.

Important bird areas of the Madrean Archipelago: A conservation strategy for avian communities. The Important Bird Areas (IBAs)

Program is a worldwide program that identifies sites considered to provide important habitats for avian species. Criteria for designation are

species abundance, diversity and range restriction. The Madrean Archipelago has a diverse range of identified IBAs that feature riparian, wetland, grassland and Madrean oak woodland habitats. Five of the 24 IBAs in the region are globally significant: the Chiricahua Mountains, San

Pedro Riparian National Conservation Area, Lower San Pedro River, Whitewater Draw State Wildlife Area, and Willcox Playa. In Mexico,

the network of IBAs continues in the Sierra Madrean Sky Islands. The purpose of designation as an IBA is highlighting the value of specific

sites. The lower elevation riparian IBAs are vital for neotropical migrants in the spring as well as provide habitat for resident species. In these

habitats, which are particularly productive in the spring, these species can refuel and rest before continuing on their journey north to breeding

grounds. The high elevation “sky island” IBAs are very productive in the fall and serve as vital stop over points for migrants headed south and

many complete their interrupted molt in these IBAs. This network serves to identify those habitats most important to conserve native birds.

Áreas importantes para las aves del Archipiélago Madrense: una estrategia de conservación para las comunidades de aves. El Programa de Áreas Importantes para la Conservación de las Aves (IBAs) es un programa mundial que identifica sitios que proporcionan hábitats

importantes para especies de aves. Criterios para la designación son la abundancia de la especie, diversidad, y restricción de localización. El

Archipiélago Madrense tiene una amplia gama de IBAs identificadas que cuentan con hábitats ribereños, humedales, pastizales y bosques de

encino Madrense. Cinco de las 24 IBAs en la región son de importancia mundial: las montañas Chiricahua, el Área Nacional de Conservación

Ribereña de San Pedro, el Área Estatal de Vida Silvestre Whitewater Draw, y Playa Willcox. En México, la red de IBAs continúa en las Islas

Serranas Madrenses. El propósito de la designación como IBA es resaltar el valor de sitios específicos. Las IBAs ribereñas de menor elevación
USDA Forest Service Proceedings RMRS-P-67. 2013 579

Abstracts/ Resúmenes

son vitales para aves migratorias neo-tropicales en la primavera, y también son el hábitat de especies residentes. En estos hábitats, que son

especialmente productivos en la primavera, estas especies pueden repostar y descansar antes de continuar su viaje hacia el norte a las zonas de

cría. Las IBAs de alta elevación en las “Islas Serranas” son muy productivas en el otoño y sirven como parada vital para aves migratorias hacia

el sur y muchos completan su muda de plumaje en estas IBAs. Esta red sirve para identificar los hábitats más importantes para la conservación

de las aves nativas.

SWANN, DON E. and Nicholas W. Perkins. Saguaro National Park, 3693 S. Old Spanish Trail, Tucson, AZ 85730, Don_Swann@nps.gov,

Nicholas_Perkins@nps.gov.

Estimating species richness and occupancy of mammals in Saguaro National Park using remote camera traps. Saguaro National Park

has numerous species of mammals, though most are hard to document and monitor due to their elusive nature. From May 2010 to April 2011

we used remote cameras to monitor mammal activity equally across stratified 1km plots based on elevation in an attempt to estimate species

richness (number of species present) and occupancy of common species. We used 48 cameras placed in 60 randomly selected plots (4 cameras

per plot) in 6-week intervals at random locations in both districts of the park (totaling 240 camera locations). We present findings from this

study, including the feasibility of using remote camera traps to estimate species richness and occupancy; other additional valuable information

that can be gained from using remote cameras; and how this information can be used to interpret mammals to public and contribute to better

resource management.

Estimación de la riqueza de especies y ocupación de mamíferos en el área protegida Saguaro National Park usando cámaras trampa 

remotas. El Parque Nacional del Sahuaro tiene numerosas especies de mamíferos, aunque la mayoría son difíciles de documentar y monitorear

debido a su naturaleza elusiva. Desde mayo de 2010 hasta abril de 2011 utilizamos cámaras remotas para monitorear la actividad de mamíferos

equitativamente en parcelas de 1 km, estratificadas por elevación, con el fin de estimar la riqueza de especies (número de especies presentes)

y la ocupación de las especies más comunes. Utilizamos 48 cámaras colocadas en 60 parcelas seleccionadas al azar (4 cámaras por parcela) en

intervalos de 6 semanas en lugares al azar en ambos distritos del parque (un total de 240 localidades de cámaras). Se presentan los resultados de

este estudio, incluyendo la posibilidad de utilizar cámaras trampas remotas para estimar la riqueza de especies y la ocupación; otra información

valiosa que puede obtenerse con el uso de cámaras remotas, y come se puede usar estos registros para informar al público sobre mamíferos y

contribuir a un mejor manejo de los recursos.

SWANN, DON E.1, Kristina M. Ratzlaff2, and Michael K. Ward3. 1Saguaro National Park, 3693 South Old Spanish Trail, Tucson, AZ 85730-

5601, Don_Swann@nps.gov; 2University of Arizona, 1110 East South Campus Dr. Rm. 123, Bldg 33, Tucson, AZ 85721, kris.ratzlaff@gmail.

com; 3Saguaro National Park, 3693 South Old Spanish Trail, Tucson, AZ 85730-5601, Michael_K_Ward@nps.gov.

Long-term dynamics of Lowland Leopard Frogs (Rana yavapaiensis). Amphibians appear to be declining throughout the globe. In the Desert Southwest, a number of amphibian species, particularly ranid frogs, have experienced dramatic decreases in population, local extinctions,

and regional extinctions. Although these declines are well documented, surprisingly little is known about the natural long-term dynamics of

amphibian populations. Lowland leopard frogs (Rana yavapaiensis) in the Rincon Mountains of Saguaro National Park are faced with challenges of drought, sedimentation of pools, potential invasion of American bullfrogs, and disease. We present findings from a 15 year monitoring

program of lowland leopard frogs in the park which provides evidence on which of these challenges pose the greatest threat to occupancy and

abundance of lowland leopard frogs.

Dinámicas a largo plazo de las ranas yavapai (Rana yavapaiensis). Los anfibios parecen estar disminuyendo alrededor el mundo. En el

desierto del suroeste norteamericano, un número de especies de anfibios, en particular ranas verdaderas, han experimentado una disminución

drástica en la población, extinciones locales y regionales. A pesar de que estas disminuciones están bien documentadas, sorprendentemente se

sabe poco acerca de las dinámicas naturales de las poblaciones de anfibios a largo plazo. Las ranas yavapai (Rana yavapaiensis) de las montañas

Rincón del Parque Nacional Saguaro se enfrentan a retos de sequía, sedimentación de las tinajas, invasión potencial de la rana toro americana, y

enfermedades. Presentamos los resultados de un programa de seguimiento de 15 años de las ranas yavapai de esta área, que ofrece datos sobre

cuál de estos desafíos plantea la mayor amenaza para la ocupación y la abundancia de ranas yavapai.

TEW, JESSICA M.1 and Shannon D. Fehlberg2. Desert Botanical Garden, 1201 N. Galvin Parkway, Phoenix, AZ 85008, 1jtew@dbg.org, 2

sfehlberg@dbg.org.

Clonal growth and genetic diversity in the endangered Huachuca Water Umbel from Southwestern Cienegas. The HuachucaWater Umbel,

Lilaeospsis schaffneriana subsp. recurva (Apiaceae)is an endangered plantfound in cienega wetlands dispersed throughoutsoutheasternArizona

and northern Sonora, Mexico. Very little is known about how much genetic variation is found within populations of this herbaceous perennial,

how different populations and watersheds are from one another, how much genetic exchange takes place among populations, and how prevalent

vegetative (rhizomes) and sexual (flowers and fruits) reproduction are within populations. We examined genetic diversity within and among ten

populations of the Huachuca Water Umbel sampled from across its range using DNA microsatellite markers. We found that patterns of genetic

diversity within populations were indicative of extensive vegetative reproduction. In general, most of the genetic diversity found in the species

was evenly distributed across populations. Genetic structure when found appeared to be influenced by geography, with populations becoming
580 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

more differentiated as proximity ofwatershedsincreased.These results provide importantinformation forimproving ongoing conservation efforts

including ex-situ collections and reintroductions and shed light on the biology and connectivity of plant populations in southwestern cienegas.

Propagación clonal y diversidad genética de la planta en peligro de extinción umbela de agua de las Huachucas, en las ciénegas del 

Suroeste Norteamericano. La umbela de agua de las Huachucas, Lilaeospsis schaffneriana subsp. recurva (Apiaceae) es una planta en peligro

de extinción que se encuentran en las ciénegas dispersas en el sureste de Arizona y el norte de Sonora, México. Se sabe muy poco sobre qué

tanta variación genética se encuentra dentro de las poblaciones de esta planta herbácea perenne, qué tan diferentes son las poblaciones y las

subcuencas hidrográficas, qué tanto intercambio genético existe entre las poblaciones, y qué tan común esla reproducción vegetativa (rizomas) y

sexual (flores y frutos). Analizamos la diversidad genética dentro y entre diez poblaciones de la umbela de agua de las Huachucas, con muestras

tomadas de toda el área de distribución utilizando marcadores microsatélites de ADN. Encontramos que los patrones de diversidad genética

dentro de las poblaciones son indicadores de extensa reproducción vegetativa. En general, la mayoría de la diversidad genética encontrada en

la especie se distribuye uniformemente a través de las poblaciones. La estructura genética, cuando se encontró, parece estar influenciada por

la geografía, con poblaciones cada vez más diferenciadas cuanto más cerca de las cuencas hidrográficas. Estos resultados aportan información

importante para mejorar los trabajos de conservación, incluyendo recolecciones y reintroducciones in situ y esclarecen la biología y la conectividad de las poblaciones de plantas en las ciénegas del Suroeste.

TILLER, RON1, Melissa Hughes2, and Gita Bodner3. The NatureConservancy, 1510 E. Fort Lowell Rd., Tucson, Arizona 85719, 1rtiller@tnc.

org, 2melissa.hughes@tnc.org, and 3gbodner@tnc.org.

Sacaton riparian grasslands of upper Ciénega Creek watershed, SE Arizona: Distribution and condition classification using state-andtransition models. Riparian grasslands dominated by Sporobolus wrightii (big sacaton) were once widely distributed in the intermountain

basins of the Madrean Archipelago. The upper Cienega Creek watershed in SE Arizona still harbors extensive sacaton stands. These alluvial

grasslands are recognized as key resources for watershed function, livestock, and wildlife. Documenting their extent and ecological state is

important for informing management in this valley and for contributing to a clearer picture of this communities’status across the region. Our

objectives were to map the distribution of sacaton; qualitatively assess stands of sacaton into condition classes; and test mapping and assessment methods for use in other valley bottoms in the region. We used a two-step approach: interpretation of aerial photography followed by

field reconnaissance. Field work consisted of qualitative, rapid assessments of ecological state using the NRCS State-and-Transition model for

the Loamy Bottom Ecological Site. The best condition “Sacaton Grassland” class occupies nearly half of alluvial habitats evaluated, with the

remainder in various states of degradation, recovery, or succession to mesquite woodland. Our observations suggest potential modifications

to the State-and-Transition model that might more accurately reflect site potential, likelihood of transitioning to other states, and management

strategies tailored to maintaining or improving condition.

Pastizales ribereños de zacatón de la cuenca alta de Ciénega Creek en el sureste de Arizona: Clasificación de la distribución y la condición 

usando modelos de estado y transición. Los pastizales ribereños dominadas por zacatón (Sporobolus wrightii) tenían una distribución amplia

en las cuencas entre montañas del Archipiélago Madrense. En la parte alta de la cuenca Ciénega Creek en el sureste de Arizona todavía existen

áreas extensas de zacatón. Se reconocen estos pastizales aluviales como un recurso vital para la función de la cuenca, el ganado y la fauna.

Es importante documentar el estado ecológico y su extensión para las actividades de manejo en este valle y tener una imagen más clara del

estado de esta comunidad en la región. Nuestro objetivo fue generar un mapa de distribución del zacatón; evaluar cualitativamente las áreas de

zacatón según su condición y probar métodos de elaboración de mapas y evaluación para su uso en otros valles ribereños de la región. Usamos

un método con dos partes: interpretación de fotografía aérea seguido por un reconocimiento de campo. El trabajo de campo consistió de una

evaluación cualitativa rápida del estado ecológico usando el modelo de estado y transición del NRCS para el sitio ecológico de Loamy Bottom. La clase con la condición mejor de “Pastizal de zacatón” ocupa casi la mitad de los hábitats de la zona aluvial evaluados, con el resto en

varios estados de degradación, recuperación o sucesión a mezquital. Nuestras observaciones sugieren una modificación al modelo de estado

y transición que podría reflejar de una manera más exacta el potencial del sitio, probabilidad de transición y estrategias de manejo adaptadas

para mantener o mejorar las condiciones.

TOLLE,CINDY1,ValerAustin1,MonicaCeciliaHolguinVilla2,JesusRamon2,ValenzuelaRosas2,MarioErandiBonillasMonge2,CarlosManuel

Valdez Colonel2, David Delgado Zamora2, and Melinda Cárdenas García2. 1Cuenca Los Ojos A.C. Agua Prieta, Sonora Mexico and Cuenca

Los Ojos Foundation, 12626 E. Turkey Creek Road, Pearce AZ 85625, (520) 824-3566; 2UNISON. Universidad de Sonora. Lic. en Biología.

Hermosillo. Blvd. Luis Encinas y Rosales s/n, Col. Centro, Hermosillo, Sonora, México. cindytolle@yahoo.com.

Monitoring restoration progress on Cuenca Los Ojos properties, Sonora Mexico. Watershed and grassland restoration began on Rancho

San Bernardino (San Bernardino Cienega) in 1999. Since this time gabions, trincheras and berms for water retention have been constructed,

and shrubs have been removed and re-seeded to native grass. In 2010, Cuenca Los Ojos initiated a restoration monitoring program. Landsat

imagery was analyzed annually in June from 1984 through 2011 using ENVI Remote Sensing software to quantify land cover changes on all

Cuenca Los Ojos properties in Sonora Mexico. Pre-restoration imagery was analyzed against post-restoration imagery to detect changes in the

San Bernardino Cienega, grasslands, shrub lands, and riparian areas. Initial results suggest an increase of 1920 acres of riparian area on Rancho

San Bernardino. Additionally, the Cienega has increased in size from 52 acres (pre-restoration) to 88 acres today. Landsat and SPOT Imagery

from 2011 was ground-truthed using resource grade GPS and permanent transects were established in several locations. Wet-dry monitoring

during the pre-monsoon month of June 2011 was done to determine perennial water at Rancho San Bernardino. Permanent Gully profile tran-
USDA Forest Service Proceedings RMRS-P-67. 2013 581

Abstracts/ Resúmenes

sects were re-measured in 2011 to determine how much sediment has been captured due to watershed restoration efforts (repeating a study

done in 2000 by Noelia de la Torre-Univ. of Sonora). Vegetation was analyzed along the gully profiles to determine changes due to restoration.

Seguimiento del progreso de restauración en las propiedades de Cuenca Los Ojos, Sonora, México. La restauración de la cuenca y el

pastizal empezaron en el rancho San Bernardino (Ciénega de San Bernardino) en 1999. Desde entonces se han construido gaviones, trincheras

y bordos para retener el agua; se removieron los arbustos y se sembraron zacates nativos. En 2010, Cuenca Los Ojos inició un programa para

darle seguimiento al progreso de restauración. Se analizaron imágenes Landsat anualmente, cada junio, desde 1984 hasta 2011 usando el software de teledetección ENVI para cuantificar los cambios de cobertura vegetal en todas las propiedades de Cuenca Los Ojos, Sonora, México.

Las imágenes tomadas antes de la restauración se compararon con imágenes tomadas después para detectar cambios en la Ciénega de San Bernardino, pastizales, matorrales y áreas ribereñas. Los resultados iniciales muestran un aumento de 780 hectáreas de áreas ribereñas en el rancho

San Bernardino. Adicionalmente, la ciénega ha aumentado en tamaño, desde 21 hectáreas (antes de la restauración) a 35 hectáreas actualmente.

Imágenes SPOT y Landsat del año 2011 fueron verificadas en tierra usando un sistema de GPS y se establecieron transectos permanentes en

varios lugares. Se hizo el seguimiento húmedo-seco durante el mes de junio de 2011, antes del monzón, para determinar las aguas perennes del

rancho San Bernardino. Se midieron de nuevo los transectos permanentes del perfil de barrancos, en 2011, para determinar cuanto sedimento se

había capturado con el trabajo de restauración (repitiendo una investigación hecha en 2000 por Noelia de la Torre- Univ. de Sonora). Se analizó

la vegetación a lo largo del perfil del barranco para determinar los cambios por la restauración.

TOWNE, GEOFFREY1, Stephen Yool1, Donald Falk2. 1School of Geography and Development, The University of Arizona, Tucson, AZ 85721; 2

School of Natural Resources and the Environment, The University of Arizona, Tucson, AZ 85721.

Using remote sensing to model fuel moisture stress for Sky Islands forests. Wildland fires have become increasingly devastating in the

Southwestern United States in the past decade, including the biodiversity rich Sky Islands of southeastern Arizona. Fire risk in this region is

increased during drought periods that lead to low fuel moisture conditions. Statistical transformations of spectral indices derived from remote

sensing provide a way to identify areas of high fuel moisture stressin vegetation.The ModerateResolution Imaging Spectroradiometer(MODIS)

provides near infrared and visible land cover data of the Sky Islands from 2000 until now. The inverse z scores of the Normalized Difference

Vegetation Index (NDVI) produce a Fuel Moisture StressIndex (FMSI) that can be used to identify when and where vegetation is under moisture

stress. The inverse z scores of these indices standardize the pixel values that may be confounded by pixels that may have the same values, but

completely different land cover types and topographies. By superimposing FMSI data from regions in years of known high fuel moisture stress,

we can establish a procedure to identify areas of high moisture stress to predict years of widespread fire occurrence. FMSI is a spatially-explicit

approach to modeling the role of interannual climate variability in modulating time-varying fire risk.

Uso de sensores remotos para el modelado de vegetación combustible en los bosques de las Islas Serranas. Desde la década pasada los

incendios forestales se han vuelto cada vez más devastadores en el Suroeste de Estados Unidos, incluyendo las Islas Serranas del sureste de

Arizona, una región de gran biodiversidad. El riesgo de incendios en esta zona aumenta durante los periodos de sequía que causa condiciones

de humedad baja en la vegetación combustible. Las transformaciones estadísticas de índices espectrales derivadas de los datos obtenidos por

sensores remotos es una manera de identificar áreas de alto riesgo de vegetación combustible por estrés de humedad. El espectroradiómetro de

imágenes de media resolución (MODIS) proporciona infrarrojo cercano y datos visibles de la cobertura del suelo de las Islas Serranas desde el

año 2000 hasta hoy. Las puntuaciones z inversas del índice de vegetación de diferencia normalizada (NDVI por sus siglas en inglés) produce un

índice de la falta de humedad del combustible (FMSI) que se puede usar para identificar el tiempo y lugar donde la vegetación está muy seca.

Las puntuaciones z inversas de estos índices estandarizan los valores de los pixeles que se pueden confundir con pixeles que tienen el mismo

valor pero que tienen tipos de cobertura de la tierra y topografías muy diferentes. Mediante la superposición de los datos acumulados por el

FMSI de lasregiones en los años conocidos de alto riesgo de vegetación combustible por estrés de humedad, podemos establecer un proceso para

identificar áreas de alto riesgo por estrés de humedad para predecir años cuando ocurrirán incendios grandes. EL FMSI es un enfoque espacialmente explícito para modelar la influencia de la variabilidad del clima interanual en la modulación del riesgo de incendios en tiempo variable.

TURNER, DALE S.1, Holly E. Richter1, Rosa Elena Jimenez Maldonado2. 1The Nature Conservancy, 1510 E. Fort Lowell Road, Tucson, AZ

85719, dturner@tnc.org, hrichter@tnc.org; 2Comisión Nacional de Áreas Naturales Protegidas, Segunda este final yAv. Sinaloa S/N. c.p. 84620,

Colonia Centro, Cananea, Sonora, elena.jimenez@conanp.gob.mx.

Mapping perennial flow in the San Pedro River: A binational monitoring project. The San Pedro River flows 279 km (173 mi) north from

its headwaters in Sonora, Mexico, to its confluence with the Gila River in Arizona, USA. It is one of the few remaining free-flowing perennial

streams in this semi-arid region, but has lost more than 50% of its perennial length. To monitor the river’s status, an annual monitoring program

was initiated in 1999 to map surface water during the driest part of the year. The length of surveyed reaches has been gradually increasing, and

in 2011 included 222 km (138 mi) of mainstem, along with 154 km (96 mi) of tributaries, in both the U.S. and Mexico. Results show a decreasing trend in some areas and increases in others, likely due to localized effects of groundwater pumping or recovery from pumping which has

recently been stopped. While some reaches were consistently wet and thus perennial, total wetted length varied widely from year to year within

most analysis segments. The results have been used for a variety of management and scientific purposes. This project also demonstrates the

power of long-term collaborative efforts across the international border, and between government agencies, non-governmental organizations,

and citizen-science volunteers.
582 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Cartografía de corrientes perennes del río San Pedro, un proyecto de monitoreo binacional. El río San Pedro recorre 279 km (173mi)

hacia el norte de su nacimiento en Sonora, Mexico hasta que se junta con el río Gila en Arizona, E.U. Es uno de los pocos ríos de corriente libre

perenne que quedan en esta región semiárida, pero ha perdido más del 50% de su longitud perenne. Para monitorear el río, se inició un programa

de monitoreo anual en 1999 para hacer mapas de las aguassuperficiales durante el periodo másseco del año. La longitud de lostramos estudiados

ha aumentado gradualmente y en 2011 fueron 222 km (138 mi) en el cauce principal y 154 km (96 mi) de corrientes tributarias tanto en E.U. y

México. Los resultados muestran una tendencia a disminuir en unas áreas y aumentar en otras, probablemente se debe a los efectos localizados

del bombeo de agua subterránea o la recuperación del bombeo que recientemente ha parado. Mientras unos tramos tienen agua constante y

por lo tanto son perennes, la longitud con agua varió ampliamente año con año en la mayoría de los segmentos analizados. Estos resultados se

han usado para diversos fines científicos y de manejo. Este proyecto también muestra el poder de los trabajos de colaboración a largo plazo a

través de la frontera internacional y entre las agencias de gobierno, organizaciones no gubernamentales y voluntarios de la ciencia ciudadana.

VALDEZ-CORONEL, CARLOS MANUEL and Reyna Amanda Castillo-Gámez. Departamento de Investigaciones Científicas y Tecnológicas

de la Universidad de Sonora, Luis Donaldo Colosio s/n, entre Sahuaripa y Reforma, Col. Centro C.P. 83000, Hermosillo, Sonora, Jim.jim.

furia@hotmail.com.

Squirrels of Sonora. Squirrels (Rodentia: Sciuridae) are usually good indicators of the conservation status of ecosystems, as they have strong

relationships with particular plant associations, plus they have a great importance in the food chain for being consumers of large amounts of

seeds and insects. Sonora is one of the states that has the biggest diversity of species of squirrels in Mexico, with 10 species in five genera,

representing 29% and 71% of the total for Mexico, respectively. These species represent three faunal influences: species of temperate forests

of the Sierra Madre Occidental, desert species with a range that extends into Arizona and grassland species that extend into New Mexico and

Chihuahua. Besides their representation in the state and its ecological and biogeographical importance, actual knowledge is currently unknown

in the state. This paper presents more information about squirrels of Sonora.

Ardillas del Estado de Sonora. Las ardillas(Rodentia: Sciuridae)suelen ser buenosindicadores del estado de conservación de los ecosistemas,

ya que presentan una fuerte relación con asociaciones vegetales particulares, además de que son de gran relevancia en las cadenas tróficas al

ser consumidores de grandes cantidades de semillas y de insectos. Sonora es uno de los estados que presenta una mayor diversidad de especies

de ardillas en México, con 10 especies en cinco géneros, que representan el 29% y 71% del total para México, respectivamente. Estas especies

representan tres influencias faunísticas: especies de bosques templados de la Sierra Madre Occidental, especies desérticas con un amplio rango

que se extiende hasta Arizona y especies de pastizales que se extienden hasta Nuevo México y Chihuahua. No obstante su representatividad en

el estado y su importancia ecológica y biogeográfica, se desconoce el estado actual de su conocimiento en el estado. Este trabajo pretende dar

a conocer más información acerca de las ardillas del estado de Sonora.

VALENZUELA-YÁNEZ, MELISSA1, Gertrudis Yanes-Arvayo1, Maria de la Paz Montañez-Armenta1, Hugo Silva-Kurumiya1, and Thomas

R. Van Devender2. 1Universidad de la Sierra, División de Ciencias Biológicas, Moctezuma, Sonora, México. 2 Sky Island Alliance, PO Box

41165, Tucson, Arizona 85717.

Floristic analysis of Ojo de Agua Tonibabi, Sierra La Madera, Sonora, Mexico. Ojo de Agua Tonibabi is an area of great historical and

biological interest located 16 km northeast of Moctezuma, Sonora, Mexico at the base of the Sierra La Madera (29o48’10”N 109o40’49”W,

624 m elevation). Plants were collected, observed, and photographed to document floristic diversity in the area on six outings between April

and November 2011. The vegetation is tropical foothills thornscrub. In the areas surrounding the spring and permanent streams, there is riparian vegetation with Goodding willow (Salix gooddingii). Collections were identified through comparisons with specimens in University of

Arizona and the Universidad de Sonora Herbaria. A total of 143 species in 114 genus and 42 families were identified. The families with the

greatest number of species are Fabaceae (26), Asteraceae (15), Euphorbiaceae (11), Convolvulaceae (8), Poaceae (8), and Cactaceae (9). Annual plants are the most abundant life form (41 taxa), followed by perennial herbs (32), trees (22), shrubs (11), vines (9), succulents (9), grasses

(8), subshrubs (7), and aquatic herbs (1).

Análisis florístico del Ojo de Agua Tonibabi, Sierra La Madera, Sonora, México. El Ojo de Agua Tonibabi es un área de gran interés biológico e histórico localizado a 16 km al noreste de Moctezuma, Sonora, México. Esta ubicado al pie de la Sierra La Madera a los 29°48’10’’

N y 109°40’49’’ O y una altitud de 624 msnm. Empezamos un inventario de la diversidad florística en esta área con seis salidas entre abril y

noviembre del 2011, para hacer observaciones y colectar ejemplares. La vegetación es de origen tropical, principalmente matorral espinoso. En

los alredores del ojo y arroyos húmedos hay vegetación riparia con presencia de sauz (Salix gooddingii). Se identificaron ejemplares mediante

visitas a los herbarios de la University of Arizona y Universidad de Sonora. Como resultado se tiene 143 especies en 114 géneros y 42 familias.

Las familias con mayor riqueza florística son Fabaceae (26), Asteraceae (15), Euphorbiaceae (11), Convolvulaceae (8), Poaceae (8) y Cactaceae

(9). Las herbáceas anuales representan la forma biológica más abundante (41 taxa), seguida por las hierbas perennes (32), árboles (22), arbustos

(11), enredaderas (9), suculentas (9), zacates (8), subarbustos (7) y hierbas acuáticas (1).
USDA Forest Service Proceedings RMRS-P-67. 2013 583

Abstracts/ Resúmenes

VALERA-AGUILAR,DANIELA1,CarlosA.López-Gonzalez2, andChristianKampichler3. 1Reservade laBiosferaMarismasNacionalesNayarit,

Comisión Nacional de Áreas Naturales Protegidas, Santiago Ixcuintla, Nayarit, daniela.valera@conanp.gob.mx; 2Laboratorio de Zoología,

Universidad Autónoma de Querétaro, Querétaro, cats4mex@aol.com; 3Dutch Centre for Avian Migration & Demography, The Netherlands,

christian.kampichler@web.de.

Identifying linkages for jaguar population connectivity in northwest Mexico. The establishment of biological corridors and linkages has

been proposed as a conservation strategy to maintain connectivity in large carnivore populations that live in highly fragmented landscapes and

require extensive home ranges. Because jaguar population connectivity in their most northern distributional limit is unknown and linkages

among populations have not been detected, we proposed to evaluate connectivity of jaguar populations and to identify potential linkages that

promote jaguar movements between populations. A virtual landscape where individuals moved was developed by SAMT software applying an

Individual Based Movement Model. The virtual landscape was developed including variables of elevation, land use types, cattle density and

human impact. Population parameters include mortality, displacement ability, and movement direction. The linkages were identified using GIS

tools. Our results suggest that most of the dispersal jaguar events are located through the Sierra Madre Occidental, showing differences between

sex and populations. Because most of the linkages important for jaguar populations are in private lands with livestock management, we propose

legal protection of them by state authorities following an archipelago reserves approach.

Identificando eslabones para la conectividad de las poblaciones de jaguar en el noroeste de México. El establecimiento de corredores

biológicos y eslabones han sido propuestos como una estrategia de conservación para mantener la conectividad en poblaciones de grandes

carnívoros que viven en paisajes altamente fragmentados. Debido a que se desconoce la conectividad de las poblaciones de jaguar en su límite

más norteño de distribución y que los eslabones entre las poblaciones no han sido identificados, nos propusimos evaluar la conectividad de las

poblaciones e identificar los eslabones potenciales que promueven los movimientos de jaguares entre las poblaciones. Se desarrolló un paisaje

virtual donde los individuos se movieron aplicando un Modelo de Movimiento Basado en Individuos en el programa SAMT. El paisaje virtual

incluye variables de altitud, tipos de uso de suelo, densidad de ganado e impacto humano. Los parámetros de poblaciones fueron mortalidad,

habilidad para desplazarse y dirección de movimientos. Los eslabones se identificaron con herramientas de SIG. Nuestros resultados sugieren

que la mayoría de los eventos de dispersión se localizan a lo largo de la Sierra Madre Occidental, mostrando diferencias entre sexos y poblaciones. Debido a que los eslabones más importantes se localizan en tierras privadas ganaderas, se propone su protección legal con autoridades

estatales, siguiendo el enfoque de las reservas archipiélagos.

VAN DEVENDER,THOMASR.1, GertrudisYanes-Arvayo2,AnaLiliaReina-Guerrero1, MelissaValenzuela-Yánez2, Maria de la Paz MontañezArmenta2

, and Hugo Silva-Kurumiya2. 1Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717 USA; 2Universidad de la Sierra, División

de Ciencias Biológicas. Moctezuma, Sonora, México.

Comparison of the tropical floras of Sierra la Madera and the Sierra Madre Occidental, Sonora, Mexico. The floras of the tropical vegetation in the Sky Island Sierra la Madera (SMA) near Moctezuma in central Sonora (30°00’N 109°18’W) and the Yécora (YEC) area in the Sierra

Madre Occidental (SMO) in eastern Sonora (28°25’N 109°15”W) were compared. The areas are 175 km apart. Tropical vegetation includes

foothills thornscrub (FTS) in both areas and tropical deciduous forest (TDF) in the Yécora area. A total of 893 vascular plant taxa are known

from these areas with 433 taxa in FTS and 793 in TDF. FTS in SMA and YEC (near Curea) had 220 and 298 taxa, with most of them also in

TDF (69.5% and 82.9%). Only 83 taxa in TDF were shared between SMA and YEC (37.7% and 27.9% of the floras). The 49 FTS species in

SMA but not YEC were not in TDF either, reflecting biotic influences from the Sonoran Desert (10), southwestern United States (8), Madrean

Archipelago (6), and a few from oak woodland and tropical western Mexico. One species (Pseudabutilon thurberi) is endemic to central Sonora

and adjacentArizona.Affinitiesto the New World tropics are very strong in both areas. The structural dominantsthat define FTS are widespread,

but composition varies greatly locally.

Comparación de la flora tropical de la Sierra la Madera con la Sierra Madre Occidental de Sonora, México. Se comparó la flora de la

vegetación tropical de la Sierra la Madera (SMA) una Isla Serrana cerca de Moctezuma en el centro de Sonora (30°00’N 109°18’W) y el área

de Yécora (YEC) en la Sierra Madre Occidental (SMO) del este de Sonora (28°25’N 109°15”W). Donde una distancia de 175 km separa las

dos regiones. La vegetación tropical incluye matorral espinoso de piedemonte (MEP) en las dos áreas y selva baja caducifolia (SBC) en el área

de Yécora. Se tienen reportados 893 taxones de plantas vasculares de estas áreas con 433 taxones en MEP y 793 en SBC. El MEP en la SMA y

YEC (cerca de Curea) tiene 220 y 298 taxones respectivamente, muchos de ellos también se encuentran en la SBC (69.5% y 82.9%). Sólo 83

taxones de la SBC se comparten entre la SMA y YEC (37.7% y 27.9% de las floras). Las 49 especies de MEP en la SMA pero no en YEC tampoco están presentes en la SBC, lo que refleja las influencias bióticas del Desierto Sonorense (10), suroeste de Estados Unidos (8), Archipiélago

Madrense (6) y unas cuantas del encinal y la región tropical del occidente de México. Una de las especies (Pseudabutilon thurberi) es endémica

del centro de Sonora y la parte adyacente de Arizona. Las afinidades con las zonas tropicales de América son muy fuertes en las dos áreas.

Las especies dominantes que determinan la estructura del MEP tienen una distribución amplia, pero la composición varía bastante localmente.
584 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

VAN DEVENDER, THOMAS R.1,,Ana Lilia Reina-Guerrero1, George M. Ferguson2, GeorgeYatskievych3, Beatriz E. Loyola-Reina4, Gertrudis

Yanes-Arvayo5, John L. Anderson6, Stephen F. Hale7, Sky Jacobs1, Maria de la Paz Montañez-Armenta5. 1Sky Island Alliance, PO Box 41165,

Tucson, Arizona 85717, vandevender@skyislandalliance.org; 2University of Arizona, Tucson, Arizona 85721; 3Missouri Botanical Garden, PO

Box 299, St, Louis, Missouri 63110; 4JRM Consultores, Ave. Sonora 89, Col. Centro, Hermosillo, Sonora, MEX; 5Universidad de la Sierra,

División de Ciencias Biológicas, Moctezuma, Sonora, MEX; 6Bureau of Land Management, 21605 North Seventh Ave., Phoenix, Arizona

85027; 7EcoPlan Associates, Inc., 701 W. Southern Ave., Suite 203, Mesa, Arizona 85210.

Preliminary flora of the Sierra Bacadéhuachi, Sonora, Mexico.TheSierra deBacadéhuachiin east-centralSonora isthewesternmostmountain

range in the Sierra Madre Occidental (SMO), located east of Bacadéhuachi, Municipio de Bacadéhuachi, 34 kilometers east of the Chihuahua

border, and 165 km south of the Arizona border. The southern portion of the range is in the Ríos Nácori Chico and Riíto drainages, both part of

the greater Río Yaqui system. The vegetation ranges from lowland foothills thornscrub (FTS) up through desert grassland to oak woodland and

pine-oak forest. The flora was sampled in December 1995 (montane forests), July 2008 (FTS), June, August, and September 2011, and March

2012 (Madrean Archipelago Biodiversity Assessment [MABA] Expeditions). The flora totals 442 taxa in 297 genera and 96 families. The most

species-rich families and genera are Asteraceae (53 taxa), Fabaceae (48 taxa each), Poaceae (46 taxa), Quercus (11 species), Cheilanthes (8 

species), Muhlenbergia (7 species), and Bouteloua (6 species) Twenty-two species are non-native (5.0%), 10 of them grasses. All observations

and collections are in the MABA database (www.madrean.org). Although tree composition and structure of the upland woodlands and forests

are similar to the Yécora area to the southeast, the preliminary Bacadéhuachi flora appears to be much less diverse.

Flora preliminar de la Sierra de Bacadéhuachi, Sonora, México. La Sierra de Bacadéhuachi en el este centro de Sonora es la región montañosa más al oeste en la Sierra Madre Occidental (SMO), ubicada al este de Bacadéhuachi, municipio del mismo nombre, 34 km al este de

la frontera con Chihuahua y 165 km al sur de la frontera con Arizona. El sur de esta sierra se localiza en la cuenca de los ríos Nácori Chico y

Riíto, afluentes de la gran cuenca del río Yaqui. La vegetación varía desde matorral espinoso de piedemonte (MEP) en la parte baja, a pastizal

del desierto y bosques de encino y de pino encino en la parte alta. La flora se colectó en diciembre de 1995 (bosques de montaña), julio de

2008 (MEP), junio, agosto y septiembre de 2011 y marzo de 2012 (expediciones del proyecto Evaluación de la Biodiversidad del Archipiélago

Madrense [MABA]). La flora actual contiene 442 taxones en 297 géneros y 96 familias. Las familias y géneros más numerosos son: Asteraceae (53 taxones), Fabaceae (48), Poaceae (43), Quercus (11 especies), Cheilanthes (8), Muhlenbergia (7 y Bouteloua (6). Veinti-dós especies

son introducidas (5.0%), 10 de ellas zacates. Todos los registros de observaciones y colectas se pueden consultar en la base de datos MABA

(www.madrean.org). Aunque la estructura y la composición de los bosques altos es similar al área de Yécora en el sureste, la flora preliminar

de Bacadéhuachi parece ser menos diversa.

VAN DEVENDER, THOMAS R.1, Erik F. Enderson2, Dale Turner3, Roberto A. Villa4, Stephen F. Hale5, George M. Ferguson6, and Charles

Hedgcock1. 1Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717, vandevender@skyislandalliance.org; 2Drylands Institute, PMB 405

2509 North Campbell Avenue, Tucson, Arizona 85719; 3The Nature Conservancy, 1510 E. Ft. Lowell Road, Tucson, Arizona 85719; 4Tucson

Herpetological Society, P.O. Box 709, Tucson, Arizona 85702; 5EcoPlan Associates, Inc., 701 W. Southern Ave., Suite 203, Mesa, Arizona

85210; 6University of Arizona, Tucson, Arizona 85721.

Comparison of preliminary herpetofaunas of the Sierras la Madera (Oposura) and Bacadéhuachi with the mainland Sierra Madre Occidental in Sonora, Mexico. Amphibians and reptiles observed in the Sierra la Madera, an isolated Sky Island mountain range in east-central

Sonora, and the Sierra de Bacadéhuachi, the westernmost Sierra Madre Occidental (SMO) range, were compared with the herpetofauna of the

Yécora area in eastern Sonora in the main Sierra Madre Occidental. Four areasin the Sierra la Madera near Moctezuma were visited in November

1981, September 2003,July 2009, and on a MadreanArchipelagoBiodiversityAssessment (MABA) Expedition inAugust 2010. MABAExpeditions went to the Sierra de Bacadéhuachi in August and September 2011, and March 2012. Observations and photos in the Sierra la Madera (59

species) and the Sierra de Bacadéhuachi (30 species) are in the online MABA database (madrean.org). About 92 species are known from along

MEX 16 in the Municipio de Yécora. Seven species in the Sky Island ranges were not in the SMO fauna. Although the Sky Island inventories

are incomplete, it is clear that the herpetofauna of the SMO is much more diverse than any Sky Island fauna. Madrean species dominate the

Sky Island faunas, which are mixtures of species with tropical, desert and northern temperate biotic affinities.

Comparasión de las herpetofaunas preliminares de la Sierra la Madera (Oposura) y Sierra de Bacadéhuachi, Sonora, México con la 

Sierra Madre Occidental. Los anfibios y reptiles observados en la Sierra la Madera, una Isla Serrana aislada en el este-centro de Sonora y la

Sierra de Bacadéhuachi, la cordillera más al oeste de la Sierra Madre Occidental (SMO) se compararon con la herpetofauna del área de Yécora

en el este de Sonora en la Sierra Madre Occidental. Cuatro áreas en la Sierra la Madera cerca de Moctezuma se visitaron en noviembre de 1981,

septiembre de 2003, julio de 2009 y en una expedición del proyecto de Evaluación de la Biodiversidad del Archipiélago Madrense (MABA) en

agosto de 2010. Las expediciones a la Sierra de Bacadéhuachi se realizaron en agosto y septiembre de 2011 y marzo de 2012. Las observaciones

e imágenes de la Sierra la Madera (59 especies) y de la Sierra de Bacadéhuachi (30 especies se pueden consultar en la base de datos en línea

MABA (madrean.org). Cerca de 92 especies son reportadas de la carretera federal MEX 16 en el municipio de Yécora. Siete especies de estas

Islas Serranas no están en la fauna de la SMO. Aunque los inventarios de Islas Serranas son incompletos, es evidente que la herpetofauna de la

SMO es mucho más diversa que cualquier fauna de Isla Serrana. Las especies madrenses dominan las faunas de las Islas Serranas, las que se

componen de especies con afinidades bióticas tropicales, desérticas y templadas del norte.
USDA Forest Service Proceedings RMRS-P-67. 2013 585

Abstracts/ Resúmenes

VAN DEVENDER, THOMAS R.1, Ana Lilia Reina-Guerrero1, and J. Jesús Sánchez-Escalante2. 1Sky Island Alliance, PO Box 41165, Tucson,

Arizona 85717, USA, yecora4@comcast.net; 2 Departmento de Investigaciones Científicas y Tecnológicas, Universidad de Sonora, Av. Rosales

y Niños Heroes s/n, Hermosillo, Sonora 83000, México.

Flora of Chihuahuan Desertscrub on Limestone in northeastern Sonora, Mexico. Transects were done in desertscrub on limestone to characterize the flora of the westernmost Chihuahuan Desert. Most of the sites (15) were in the Municipios of Agua Prieta and Naco in northeastern

Sonora. Single sites were near Ascensión in northwestern Chihuahua and east of Douglas in southeastern Arizona. A total of 250 species were

recorded on the transects. When analyzed by life form, perennial herbs (60 species) were the most numerous, followed by annual herbs (45

species), subshrubs (37 species), and woody shrubs (34 species). The most diverse limestone floras were on Rancho La Morita in the Municipio

of Naco. Two transects on a hill west of Arroyo La Bellota had 90 and 121 taxa. Ocotillo (Fouquieria splendens), sotol (Dasylirion wheeleri), 

beargrass (Nolina microcarpa), and littleleaf sumac (Rhus microphylla) were dominants. On the nearby Cerro La Bruja transect (96 taxa), Chihuahuan whitethorn acacia (Acacia neovernicosa), ocotillo, and shrubby senna (Senna wislizenii) were common on a grassy, limestone slope.

The flora was also diverse in the La Calera area on the Sierra Anibácachi in the Municipio of Agua Prieta with 93 taxa on a transect dominated

by Sonoran rosewood (Vauquelinia californica var. pauciflora), a rare Sonoran shrub.

Flora del matorral del Desierto Chihuahuense en las calizas del noreste de Sonora, México. Se utilizó el método de transectos en matorral

desértico de caliza para evaluar la flora de la región más occidental del Desierto Chihuahuense. La mayoría de los sitios de estudio (15) se ubicaron en los municipios de Agua Prieta y Naco en el noreste de Sonora. También se evaluaron sitios únicos cerca de Ascensión en el noroeste de

Chihuahua y al este de Douglas al sureste de Arizona. Se registraron un total de 250 especies en los transectos. Al hacer el análisis por forma de

vida se encontró que las hierbas perennes (60 especies) fueron las más numerosas, seguidas por las hierbas anuales (45 especies), subarbustos

(37 especies) y arbustos leñosos (34 especies). Las floras de caliza más diversas se encontraron en rancho La Morita en el municipio de Naco.

Dos transectos en este sitio, en un cerro al oeste del arroyo La Bellota registraron 90 y 121 taxones. . Ocotillo (Fouquieria splendens), sotol

(Dasylirion wheeleri), palmilla (Nolina microcarpa) y limita (Rhus microphylla) fueron las especies dominantes. En el transecto del cerro la

Bruja (96 taxones), cerca del arroyo La Bellota, vinorama (Acacia neovernicosa), ocotillo y Senna wislizenii fueron comunes en una ladera con

suelos de caliza y zacates abundantes. La flora también se encontró diversa en el área de La Calera, en la Sierra Anibácachi del municipio de

Agua Prieta con 93 taxones en un transecto dominado por Vauquelinia californica var. Pauciflora, un arbusto raro en Sonora.

VAN DEVENDER, THOMAS R.1, Sergio Avila-Villegas1, Melanie Emerson1, Dale Turner2, Aaron D. Flesch3, Nicholas S. Deyo1. 1Sky Island

Alliance, PO Box 41165, Tucson, Arizona 85717, USA, vandevender@skyislandalliance.org; 2The Nature Conservancy, 1510 E. Ft. Lowell

Road, Tucson, Arizona 85719, USA; 3Avian Science Center, Division of Biological Sciences, 32 Campus Drive, University of Montana, Missoula, Montana 59812.

Biodiversity in the Madrean Archipelago of Sonora, México. The Madrean Archipelago is the area of isolated Sky Island mountain ranges

between the Sierra Madre Occidental (SMO) in Sonora and Chihuahua and the Mogollon Rim in New Mexico andArizona. Five biotic provinces

converge in the area: tropical temperate SMO, temperate Rocky Mountains, Great Plains-Chihuahuan Desert, Sonoran Desert, and western

Mexico lowlands tropics. The transition between the New World tropics and the northern temperate zone is 28-29°N in east-central Sonora.

In 2009 Sky Island Alliance initiated the Madrean Archipelago Biodiversity Assessment (MABA) program to document distributions of all

plants and animals in the Madrean Archipelago from historical (museum collections, literature, etc.) sources, and observations on expeditions

to high diversity mountain ranges of conservation interest. Species numbers in the MABA database provide preliminary diversity estimates

for the Madrean Archipelago in Sonora. There are ca. 2880 plant taxa in northeastern Sonora. Insects are very diverse, but only 1380 taxa are

documented in the Sky Islands Region. Vertebrates are better represented: fish (39 species), amphibians and reptiles (104 species), birds (358

species), and mammals (76 species). Surprising, diversity appears to decrease southward, but probably reflects the NNW-SSE orientation of

the ranges, and decreasing area and fewer biological inventories to the south.

Biodiversidad del Archipiélago Madrense de Sonora, México. El área del Archipiélago Madrense lo forman las cadenas montañosas Islas

Serranas aisladas entre la Sierra Madre Occidental (SMO) en Sonora y Chihuahua y la región del Mogollón Rim en Nuevo México y Arizona.

Cinco provincias bióticas convergen en el área: SMO tropical templada, Montañas Rocallosas templadas, Desierto Chihuahuense – Grandes

Llanuras, Desierto Sonorense y la región tropical del occidente de México. La transición entre los trópicos de América y la zona templada del

norte se localiza en 28-29°N en la parte este centro de Sonora. Sky Island Alliance empezó el programa Evaluación de la Biodiversidad del

Archipiélago Madrense (MABA) en 2009 para documentar la distribución de las especies de plantas y animales en el Archipiélago Madrense

con registros históricos(colecciones de museos, literatura, etc.) y observaciones hechas en expediciones a sierras con alta diversidad y de interés

para su conservación. Estimaciones preliminares de la diversidad delArchipiélago Madrense en Sonora se obtienen a través de las especies en la

base de datos MABA. Se cuenta con cerca de 2880 taxones de plantas en el noreste de Sonora. Los insectos tienen una diversidad alta, pero sólo

1380 taxones están documentados en la Región de las Islas Serranas. Los vertebrados están mejor representados: peces (39 especies), anfibios

y reptiles (104 especies), aves (358 especies) y mamíferos (76 especies). Es sorprendente que la diversidad parece ser que disminuye hacia el

sur, pero probablemente refleja la orientación NNO-SSE de las cordilleras, reducción del área y muy pocos inventarios biológicos en el sur.
586 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

VAN RIPER, CHARLES, III1, J. Kellermann2, TJ Fontaine3, Susan Skagen4. 1U.S. Geological Survey, SBSC/SDRS, University AZ, Tucson

Arizona 85721; 2School of Natural Resources and the Environment, University of Arizona, Tucson, AZ 85721; 3USGS, University Nebraska,

Lincoln Nebraska 68583; 4USGS, Fort Collins, Colorado 85721.

The influence of plant phenological patterns on migrating neotropical migrant birds in western North America.In western NorthAmerica,

migration patterns of neotropical land birds evolved along landscapes at different elevations and within heterogeneous and patchy environments. Western migrant land birds appear to assess migrant routes and stop over habitats at four major scales; 1) Genetically influenced corridor

selection influenced by local weather patterns; 2) large-scale landscape features; 3) Vegetation patches; and 4) microhabitat selection within

the vegetation patch. Along the lower Colorado River in Mexico, California and Arizona, and the Santa Cruz and San Pedro rivers in Arizona,

these four scales are variously influenced by weather, vegetative species, structure, and plant phenology patterns that appear to provide a cue

to insect prey base. In migrating neotropical migrant warblers that we have examined along the Colorado, Santa Cruz and San Pedro rivers,

species arrival dates and numbers were variable among years, being largely influenced by plant phenology cycles. Therefore, stopover and bird

foraging patterns were greatly influenced by plant species and phenological patterns of the selected microhabitat. It thus appears that large scale

landscape features, along with riparian habitat structure, vegetation phenology, and insect prey base all play a role in structuring spring warbler

migration patterns along southwestern riparian corridors.

La influencia de la fenología de plantas en aves neotropicales migratorias en el oeste de América del Norte. En el oeste de América del

Norte, las rutas de migración de aves terrestres neotropicales evolucionaron en paisajes de elevaciones diferentes y entornos fragmentados y

heterogéneos. Las aves terrestres migratorias occidentales parecen evaluar rutas de migración y parar en hábitats en cuatro escalas mayores; 1)

selección de corredores por herencia genética influenciados por condiciones del clima local; 2) características mayores del paisaje; 3) manchas

de vegetación; y 4) selección de un microhábitat en la mancha de vegetación. En el río Colorado en México, California y Arizona y los ríos

Santa Cruz y San Pedro en Arizona, estas cuatro escalas son influenciadas de diversas maneras por el clima, las especies de plantas, la estructura

y las condiciones fenológicas de las plantas que parecen proporcionar una señal a los insectos presa. En los chipes neotropicales migratorios

estudiados en los ríos Colorado, Santa Cruz y San Pedro, las fechas de llegada de la especie y la cantidad variaron cada año, influenciados

principalmente por los ciclos de la fenología de plantas. Por lo tanto, las condiciones del forrajeo y paradas temporales de los pájaros fueron

influenciadas principalmente por las especies de plantas y las condiciones fenológicas del microhábitat seleccionado. En consecuencia, parece

que las características mayores del paisaje, junto con las estructuras del hábitat ribereño, la fenología de la vegetación y la base de insectos presa

todos tienen un efecto en la estructura de las rutas de migración de los chipes de primavera en los corredores ribereños del suroeste.

VARAS, CORA1, Paul R. Krausman2, Carlos A. López-Gonzalez3, Melanie Culver1, Kirby Bristow4, and Esther Rubin4. 1School of Natural

Resources and the Environment, University of Arizona, Tucson, Arizona, 85721, USA, cora@email.arizona.edu, mculver@email.arizona.edu; 2

Boone and Crockett Program in Wildlife Conservation, University of Montana, Missoula, Montana 59812, USA, paul.krausman@cfc.umt.edu; 3

Universidad Autónoma de Querétaro, Querétaro, México, C. P. 76010, cats4mex@aol.com; 4Arizona Game and Fish Department, Phoenix,

Arizona, 85086, USA, kbristow@azgfd.gov, erubin@azgfd.gov.

Are black bear moving among the mountains in the Sky Islands, Arizona and Northern Mexico? Habitat fragmentation in the sky islands

of Arizona and northern Mexico has important implications to the genetic diversity, gene flow, and ultimately the population structure of local

taxa. Black bears (Ursus americanus) inhabit the sky islands of the Madrean archipelago and are a species of public interest and management

focus in the US and of conservation concern in Mexico. We used 10 nuclear DNA (nDNA) microsatellite markers to investigate population

structure of black bears in the sky islands of Arizona and northern Mexico. We collected 396 bear samples and used spatial and non-spatial

Bayesian assignment models to evaluate nDNA genetic structure and cluster individuals into genetically distinct groups. Subtle population

structure was detected indicating high levels of gene flow in recent generations, especially throughout the sky islands region, while lower gene

flow was detected between the “mainland” Mazatzal Mountains of Arizona, and the furthest sky islands in northern Mexico, and between the

Mazatzal Mountains and the Sierra Madre Occidental, Mexico. The GENELAND, STRUCTURE AND TESS analysis indicated two populations separating the entire sky island region from the Mazatzal Mountains, with an average Fstof 0.474 yet connectivity between the two groups

was higher with an average Fst of 0.07 using these estimators (range = 0.004-0.14). These results indicate that black bears are moving among

the sky islands, with distance being the main factor that isolates populations.

¿Se trasladan los osos negros entre sierras de las Islas Serranas de Arizona y el norte de México? La fragmentación del hábitat en las Islas

Serranas de Arizona y el norte de México tiene implicaciones importantes en la diversidad genética, el flujo genético, y a lo largo la estructura

de la población de los taxones locales. Los osos negros (Ursus americanus) habitan las Islas Serranas del archipiélago Madrense y son una

especie de interés público y manejo centrado en E.U. y están protegidos en México. Usamos 10 marcadores de microsatélite de ADN del núcleo

(nDNA por sus siglas en inglés) para investigar la estructura de la población de oso negro en las Islas Serranas de Arizona y el norte de México.

Colectamos 396 muestras de osos y usamos modelos de asignación Bayesianos espaciales y no espaciales para evaluar la estructura genética

del nDNA y agrupar individuos en grupos genéticamente distintos. Detectamos estructura sutil de la población que indica niveles altos de flujo

genético en generacionesrecientes, especialmente en la región de lasIslas Serranas, aunque detectamos menos flujo genético entre las montañas

“continentales” de Mazatzal de Arizona y las Islas Serranas del norte de México más alejadas, y entre las montañas Mazatzal y la Sierra Madre

Occidental, México. Los análisis Geneland, Structure y Tess indicaron dos poblaciones que separan la región entera de las Islas Serranas de la

montañas Matzalzal, con un promedio Fst de 0.474 pero la conectividad entre los dos grupos fue más alto con un promedio Fst de 0.07 usando

estos estimadores (intervalo= 0.004-0.14). Estos resultados indican que los osos negros se desplazan entre las Islas Serranas, con la distancia

siendo el factor principal que aísla las poblaciones.
USDA Forest Service Proceedings RMRS-P-67. 2013 587

Abstracts/ Resúmenes

VILLA-ANDRADE,MARTÍNA.1,ThomasR.VanDevender2,MartínPadrés-Contreras3,FernandoPadrés3,M.Reyes-Juárez1,G.Luna-Salazar1,

Paul S. Martion4. 1Comisión de Ecología Sustentable del Estado de Sonora,Reyes yAguascalientess/n Esq.,Col. SanBenito, Hermosillo, Sonora

83190, mvilla55@yahoo.com; 2Sky Island Alliance, PO Box 41165, Tucson, Arizona 85717, vandevender@skyislandalliance.com; 3Rancho

Agua Fría,Ave. 5 de Mayo # 213 Norte, Magdalena, Sonora 84160; 4Department of Geosciences, University ofArizona, Tucson,Arizona 85721.

Biodiversity and conservation of the Ciénega de Saracachi area, Sonora, México. The Ciénega de Saracachi area, including Arroyos Santo

Domingo and Quemado and Rancho la Brisca, is in north-central Sonora (30°22’N 110°25”W; 79 km2 area; 960-1,000 m elevation) ca. 100

km south of the Arizona border in the Municipio de Cucurpe. The vegetation is desert grassland on slopes and cottonwood-willow riparian

forest in the Ciénega and rocky stream canyons. These upper tributaries of the Río San Miguel are natural corridors on the west side of the

Sierras San Antonio and Azul. The high diversity of the area has been recognized since the mid-1970s. In April 2011, a Madrean Archipelago

Biodiversity Assessment (MABA) Expedition provided additional species documentation. In the MABA database (www.madrean.org), there

are 434 species of plants in 105 families and 271 species of animals in 81 families documented in the area. Invertebrate animals with only 98

taxa in 23 families are very poorly known. Vertebrate animals with 172 species in 58 families are dominated by birds (77.9% of the species).

Additional documentation is needed for plants, amphibians, and reptiles (especially snakes) in the summer rainy season, for migratory birds,

and for invertebrates and mammals (especially bats) in general. About 10 species of animals are listed as Threatened or Of Special Concern in

the NOM-059-SEMARNAT-2010 (the Mexican endangered species law). The La Brisca talussnail (Sonorella aguafriensis) is endemic to Arroyo Santo Domingo. Several species of plants and grasshoppers are only known in Sonora from the Ciénega. In 2010, the Saracachi area was

nominated to be a Sonoran Área Natural Protegida to preserve its natural values and to develop ecotourism land use options.

Biodiversidad y conservación del área de la Ciénega de Saracachi, Sonora, México. El área de la Ciénega de Saracachi la cual incluye el

arroyo Santo Domingo, arroyo el Quemado y rancho la Brisca se localiza en la parte norte centro de Sonora (30°22’N 110°25”W; 79 km2 área;

960-1,000 m elevación) ca. de 100 km al sur de la frontera con Arizona en el municipio de Cucurpe. La vegetación es pastizal del desierto en las

laderas, y bosques ribereños de álamo y sauce en la ciénega y cañones rocosos de arroyos. Los afluentes altos del río San Miguel son corredores

naturales en el occidente de la Sierra San Antonio y de la Sierra Azul. La alta diversidad del área se ha reconocido desde mediados de 1970. En

abril de 2011 una expedición del proyecto Evaluación de la Biodiversidad delArchipiélago Madrense (MABA) agregó nuevosregistros de especies. En la base de datos de MABA (www.madrean.org), hay 434 especies de plantas en 105 familias y 271 especies de animales en 81 familias

documentadas en el área. Los invertebrados con sólo 98 taxones en 23 familias se conocen muy poco. Los vertebrados con 172 especies en 58

familias están dominados por aves (77.9% de las especies). Se necesita documentar más plantas, anfibios y reptiles (en especial serpientes) en

la temporada de lluvias del verano, así como aves migratorias e invertebrados y mamíferos (con énfasis en murciélagos) en general. Alrededor

de 10 especies de animales están en la lista de la NOM-059-SEMARNAT-2010 con categoría de amenazados o especie en riesgo. El caracol

de talud de La Brisca (Sonorella aguafriensis) es endémico del arroyo Santo Domingo. Varias especies de plantas y chapulines se conocen en

Sonora sólo de la Ciénega. El área de Saracachi se propuso como Área Natural Protegida estatal en 2010 para conservar sus atributos naturales

y desarrollar opciones de ecoturismo.

VILLARREAL, MIGUEL L.1, Leila Gass1, Jack L. Childs2, Laura M. Norman1, Joel B. Sankey1, and Cynthia S.A. Wallace1. 1U.S. Geological Survey, Western Geographic Science Center, 520 N. Park Ave., Tucson, AZ 85719, mvillarreal@usgs.gov; 2

Borderlands Jaguar Detection

Project, 1165 W. Hawk Way, Amado, AZ 85645.

Examining wildlife responses to phenology and wildfire using a landscape-scale camera trap network. Between 2001 and 2009, the

Borderlands Jaguar Detection Project deployed 174 camera traps in the Baboquivari, Sierrita, Tumacacori, Atascosa, Patagonia, and Santa Rita

mountains of southern Arizona to record jaguar activity. In addition to the jaguar, these motion-activated cameras, placed along known wildlife

travel routes, recorded occurrences of ~ 20 other animal species. We examined relationships of temporal patterns of species observations to

landscape phenology and the timing of wildfire events. Landscape phenometrics were derived from time series of monthly Normalized Difference Vegetation Index data from the Moderate Resolution Imaging Spectroradiometer using statistical and curve-fitting techniques. The remote

sensing phenometrics were stratified by vegetation type, and analyzed in conjunction with wildlife occurrences by species and camera location.

We examined temporal and spatial changes in number of sightings of select species prior to, during, and after three wildfires in the area using

fire perimeter data from the Monitoring Trends in Burn Severity database. The analyses suggest that phenology and natural disturbance are

related to local and landscape-scale dynamics of wildlife in the sky island region; information important for wildlife managers faced with uncertainty regarding changing climate and disturbance regimes.

Análisis de la respuesta de fauna a fenología e incendios forestales mediante el uso de una red de cámaras trampa en un paisaje. Entre

2011 y 2009, el proyectoBorderlandsJaguar Detection colocó 174 cámarastrampa en las cordillerasBaboquivari, Sierrita,Tumacácori,Atascosa,

Patagonia y Santa Rita delsureste deArizona para registrar la actividad de jaguares.Además del jaguar, estás cámaras activadas por movimiento,

colocadas en rutas conocidas de fauna, registraron la presencia de otras ~20 especies de animales. Analizamos la relación entre el tiempo de

ocurrencia de especies observadas, la fenología del paisaje y el tiempo de incendios forestales. La fenométrica del paisaje se derivó de series

de tiempo mensuales de los datos del índice de vegetación de diferencia normalizada tomada del espectroradiómetro de imágenes de media

resolución usando técnicas estadísticas y de ajuste de curva. Se estratificó la fenométrica de sensores remotos por tipo de vegetación y se analizó

con ocurrencias de fauna por especies y la ubicación de la cámara. Examinamos cambios temporales y espaciales en series de observaciones

de ciertas especies antes de, durante y después de tres incendios forestales del área usando datos del perímetro del incendio tomado de la base

de datos tendencia del monitoreo en la severidad del incendio. El análisis indica que la fenología y la perturbación natural están relacionadas
588 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

con la dinámica local y regional de fauna en la región de las Islas Serranas; información valiosa para los encargados del manejo de fauna que

enfrentan incertidumbre con respeto al clima cambiante y regímenes de perturbación.

WALLACE, CYNTHIA S. A.1, Miguel Villarreal1, and Charles van Riper, III2. 1U.S. Geological Survey, Western Geographic Science Center,

520 N. Park Ave., Tucson, AZ 85719, cwallace@usgs.gov. 2U.S. Geological Survey, SBSC/SDRS, University AZ, Tucson Arizona 85721

Mapping landscape phenology preferences of yellow-billed cuckoo with AVHRR data. We mapped habitat for threatened Yellow-billed

Cuckoo (Coccycus americanus occidentalis) in the State of Arizona using the temporal greenness dynamics of the landscape, or the landscape

phenology. Landscape phenometrics were derived from Advanced Very High Resolution Radiometer (AVHRR) Normalized Difference Vegetation Index (NDVI) data for 1998 and 1999 by using Fourier harmonic analysis to analyze the waveform of the annual NDVI profile at each

pixel. We modeled the spatial distribution of Yellow-billed Cuckoo habitat by coupling the field data of Cuckoo presence or absence and pointbased samples of riparian and cottonwood-willow vegetation types with satellite phenometrics for 1998. Models were validated using field

and satellite data collected in 1999. The results indicate that Yellow-billed Cuckoo occupy locations within their preferred habitat that exhibit

peak greenness after the start of the summer monsoon and are greener and more dynamic than “average” habitat. Identification of preferred

phenotypes within recognized habitat areas can be used to refine habitat models, inform predictions of habitat response to climate change, and

suggest adaptation strategies.

Cartografía del paisaje y fenología preferidos del cuco pico amarillo usando datos AVHRR. Elaboramos mapas del hábitat del cuco pico

amarillo (Coccycus americanus occidentalis) amenazado en el estado de Arizona con la dinámica temporal verdor del paisaje, o la fenología

del paisaje. Los aspectos fenometricos del paisaje se derivaron de los índices de vegetación de diferencia normalizada (NDVI) obtenidos de

las imágenes de satélite con sensor AVHRR para 1998 y 1999 mediante el análisis de Fourier para examinar el perfil anual de NDVI en cada

píxel. Hemos modelado la distribución espacial del hábitat del cuco pico amarillo al relacionar los datos de campo de presencia o ausencia de

cuco y los muestreos de puntos de los tipos de vegetación ribereña y de álamo-sauce con fenometricos del satélite para 1998. Los modelos se

corroboraron utilizando los datos de campo y de satélite tomados en 1999. Los resultados indican que el cuco pico amarillo ocupa lugares de

su hábitat preferido que exhiben un verdor pico después del comienzo del monzón de verano y son más verdes y más dinámico que el hábitat

"promedio". La identificación de fenotipos preferidos en hábitats reconocidos se puede utilizar para mejorar los modelos de hábitat, predecir la

respuesta del hábitat al cambio climático y sugerir estrategias de adaptación.

WARSHALL, PETER1 andAaronFlesch2 1 ScienceCoordinator, Northern Jaguar Project, 350 S. GrandeAve,Tucson,AZ85745, pwturtle@well.

com; 2School of Natural Resources and the Environment, University ofArizona, 325 Bioscience East, TucsonAZ 85721, flesch@ag.arizona.edu.

Lessons learned concerning management and conservation of Sky Island birds on the Northern Jaguar Reserve. The Northern Jaguar

Reserve is in the western foothills of the Sierra Madre Occidental and occupies a broad transition zone between Nearctic and Neotropical faunal

realms. We assessed the distribution and abundance of birds across all four seasons on the reserve and evaluated conservation and management issues and threats. Since 2007, we have observed 201 species of birds including 130 listed by USFWS as Neotropical migrants, 104 that

presumably breed, 20 species of bi-national conservation concern, and a surprisingly large number of species that either winter or breed north

of their previously described range limits. Observations suggest the importance of an inland migratory pathway for Neotropical migrants along

the Rio Yaqui corridor and indicate the Reserve’s value in providing habitat for short-distance migrants that breed in the Sierra Madre and Sky

Islands, and long-distance migrants that breed further north. This region faces major conservation challenges including a proposed dam and

mining and cattle impacts. Existing conservation programs include land purchase, outreach to neighboring private ranches, habitat restoration,

and environmental education. Numerous conservation and management unknowns exist in the Foothills thornscrub and will be discussed.

Lecciones aprendidas sobre manejo y conservación de aves de las Islas Serranas en la Reserva del Jaguar del Norte. La Reserva del

Jaguar del Norte se localiza en el piedemonte de la Sierra Madre Occidental y ocupa una zona ancha de transición entre dominios de fauna

neártica y neotropical. Evaluamos la distribución y abundancia de aves en las cuatro estaciones en la reserva y analizamos su conservación,

manejo y amenazas. Desde 2007, hemos registrado 201 especies de aves incluyendo 130 catalogadas como migratorias neotropicales por el

Servicio de Pesca y Vida Silvestre (USFWS por sus siglas en inglés), 104 que se presume se reproducen, 20 especies de importancia para la

conservación binacional y una cantidad sorprendente de especies que pasan el invierno o anidan al norte del límite previamente conocido. Las

observaciones sugieren la importancia de rutas migratorias del interior para migrantes neotropicales en el corredor del río Yaqui y demuestra

el valor de la reserva por ser un hábitat para migrantes de distancias cortas que se reproducen en la Sierra Madre y las Islas Serranas, así como

de migrantes de distancias largas que anidan más al norte. Esta región enfrenta enormes retos de conservación, entre ellas la propuesta de una

presa, e impactos por la minería y la ganadería. El programa de conservación actual incluye la compra de terrenos, compromisos con ranchos

privados adyacentes, restauración del hábitat, y educación ambiental. Se abordarán las numerosas lagunas de conocimiento sobre el manejo y

conservación del matorral espinoso de piedemonte.

WARSHALL, PETER. Science Coordinator, Northern Jaguar Project, 2114 W. Grant Rd. #121, Tucson, AZ 85745, pwturtle@well.com.

Can and when will female jaguars cross the border? Knowledge of the socio-demographics of the northern jaguar and implications for 

the “jaguar corridor.” A viable population of jaguars in the US cannot be achieved until females migrate across the border or are re-introduced

and then encounter an adult male. This talk will present the latest data on social life of the ordinarily considered solitary felid. This includes
USDA Forest Service Proceedings RMRS-P-67. 2013 589

Abstracts/ Resúmenes

marking, home range size, female/male differences, use of mother’s home range by young, and movements by age and sex. It will then discuss

what is known of the life-history phenomena that help push jaguars to disperse (birth and death rates, age at reproduction, sex ratios, total

young per life-time, etc.). The combined socio-demographics will be incorporated into a very simple model and applied to existing knowledge

of female geographic distribution to illustrate dispersal scenarios and times to cross into the United States. Implications for conservation and

management will be described.

¿Cruzarán la frontera los jaguares hembra? Conocimiento sobre la población del jaguar norteño e implicaciones de “corredores de 

jaguares.” Una población viable de jaguares en E.U. se logrará hasta que las hembras crucen la frontera o se reintroduzcan y encuentren un

macho adulto. Se presentarán los datos más recientes sobre la vida social del que comúnmente se considera un felino solitario. Incluye marcaje,

tamaño del territorio, diferencias entre machos y hembras, uso del territorio de la madre por las crías y movimiento por sexo y edad. Después

se abordará el fenómeno del ciclo de vida que empuja a que los jaguares se dispersen (tasas de natalidad y mortalidad, edad de reproducción,

proporción por sexo, total de crías en su vida, etc.). Los índices demográficos combinados se utilizarán para diseñar un modelo sencillo y se

aplicará a la distribución geográfica conocida de hembras para ilustrar escenarios de dispersión y plazos para cruzar a Estados Unidos. Se

abordarán las implicaciones de manejo y conservación.

WETHINGTON, SUSAN M.1, Pieter S. A. Beck2, Scott J. Goetz2, Don R. Powers3, and Catherine H. Graham4. 1Hummingbird Monitoring

Network, P.O. Box 115, Patagonia, AZ, swething@dakotacom.net; 2Woods Hole Research Center, Falmouth, Massachusetts, pbeck@whrc.org,

sgoetz@whrc.org; 3Department of Biology, George Fox University, Newberg, OR, dpowers@georgefox.edu; 4Stony Brook University, Stony

Brook, NY, cgraham@life.bio.sunysb.edu.

Understanding and predicting the impact of climate change on hummingbird diversity and migration in the Madrean Archipelago.

Perhaps the greatest threats to hummingbird survival is the effect of changing climates on flowering phenology, where even minor changes in

climate can produce large changes in nectar availability and in blooming dates that may decouple the mutualism between hummingbirds and the

plants they pollinate. The increasing drought predicted for the Madrean Archipelago could have significant impact on hummingbird diversity

in the USA by adversely affecting their nectar resources and thus their ability to migrate through and nest in this area. We are combining timeseries data for hummingbirds with climate and remote sensing data to evaluate what changes have occurred in hummingbird populations and to

predict future changes. A preliminary data analyses shows a good relationship between MODIS phenology and the abundance of Black-chinned

Hummingbirds at a site along Harshaw Creek in the Patagonia Mountains of Arizona. In this paper, we will report on the latest findings of this

effort and introduce this new research project funded by NASA through their Biological Response and Climate Change call.

Impacto del cambio climático en la diversidad y migración de colibrís en el Archipiélago Madrense, conocimiento y predicciones. Quizás

la mayor amenaza a la supervivencia de los colibrís es el efecto del cambio climático en la fenología floral, donde incluso fluctuaciones menores

del clima pueden ocasionar grandes cambios en el néctar disponible y en las fechas de floración que pueden desacoplar el mutualismo entre los

colibrís y las plantas que polinizan. Las grandes sequías pronosticadas para el Archipiélago Madrense podrían tener impactos significativos en

la diversidad de colibrís en E.U. al afectar negativamente los recursos de néctar y por consiguiente su capacidad para migrar y anidar en el área.

Se combinaron los datos de series cronológicas de colibríes con datos de clima y de sensores remotos para evaluar que cambios han ocurrido

en las poblaciones de colibrís y predecir cambios futuros. Un análisis preliminar muestra una relación buena entre la fenología MODIS y la

abundancia del colibrí barba negra en un sitio de Harshaw Creek en la Sierra Patagonia de Arizona. En esta publicación informaremos sobre

los resultados más recientes de este trabajo e introduciremos este nuevo proyecto de investigación financiado por NASA por medio del área de

“respuesta biológica y cambio climático”.

WETHINGTON, SUSAN M.1, Maria Del Rocio Meneses-Ramírez1, Andreia Malpica-Topete2, Luz Zamudio-Beltrán3, Cindy Tolle4, Valer

Austin4, and H. Ron Pulliam5. 1Hummingbird Monitoring Network, P.O. Box 115, Patagonia,AZ.swething@dakotacom.net, mr_biol@hotmail.

com; 2Department of Evolutionary Biology, Institute of Ecology, AC., Veracruz, Mexico, andreia.malpica@posgrado.inecol.edu.mx; 3Museo

de Zoología “Alfonso L. Herrera”, Departamento de Biología Evolutiva, Facultad de Ciencias Universidad Nacional Autónoma de México,

zbluze@hotmail.com; 4Cuenca Los Ojos, 12626 East Turkey Creek Road, Pearce, AZ, vaustin@elcoronadoranch.net, cindytolle@yahoo.com; 5

University of Georgia, Odum School of Ecology, Athens, Georgia, pulliam2@uga.edu.

Restoring riparian vegetation for nesting hummingbirds: Is there a conservation dilemma? Hummingbirds, as pollinators, provide important ecosystem services needed for many plants to produce viable seeds and fruit, which then become food resources for many animal species. Building upon the successful efforts by Cuenca Los Ojos to restore the San Bernardino Ciénega and associated riparian zones in northern

Sonora, we investigated if this restored habitat provided the resources needed for successful nesting by hummingbirds by comparing it with a

Sonoita Creek/Patagonia Lake site known to be used successfully by nesting Black-chinned and Broad-billed Hummingbirds. We conducted

point counts and nest searches to detect hummingbird abundance and richness and counted floral resources used by hummingbirds. During late

May, floral resources were similar at both sites but by mid-June, the only reliable nectar resource was tree tobacco (Nicotiana glauca) at Sonoita

Creek. During May, three nests were found at San Bernardino but none remained active in June.At Sonoita Creek, nesting continued throughout

the censuses with 72 nests found. Viable nectar resources likely limits the ability of hummingbirds to nest in restored riparian habitats at San

Bernardino but Tree Tobacco, considered by many to be an invasive species, may not be the best option for restoring the nectar landscape.
590 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

Restauración de la vegetación ribereña para la anidación de colibrís, ¿un dilema de conservación? Los colibrís, como polinizadores,

aportan servicios ecosistémicos necesarios para que muchas plantas produzcan semillas viables y frutos que después se convierten en recursos

alimenticios de especies animales. Partiendo del trabajo exitoso de Cuenca los Ojos de restauración de la ciénega San Bernardino y las zonas

ribereñas asociadas en el norte de Sonora, investigamos si este hábitat restaurado contaba con los recursos necesarios para la anidación exitosa

de colibrís al compararlo con un sitio de Sonoita Creek y Patagonia Lake conocido como área de anidación exitosa de dos especies de colibríes,

Archilochus alexandri y Cynanthus latirostris. Se realizaron muestreos mediante transectos de conteo y búsquedas de nidos para determinar

la abundancia y diversidad de colibríes y contamos los recursos florales usados por los colibrís. A finales de mayo, los recursos florales fueron

similares en ambos sitios, pero a mediados de junio, el único recurso fiable de néctar fue el juan loco (Nicotiana glauca) en Sonoita Creek. En

mayo, se encontraron tres nidos en San Bernardino pero ninguno estaba activo en junio. En Sonoita Creek, la anidación continuó durante el

censo, encontrándose 72 nidos. Recursos viables de néctar probablemente limitan la capacidad de anidación del colibrí en los hábitats ribereños restaurados de San Bernardino pero el juan loco considerado por muchos como una especie invasora, quizás no sea la mejor opción para

restaurar el paisaje del néctar.

WHITE , MARGARET S. and Juliet C. Stromberg. Arizona State University, School of Life Sciences, Tempe, AZ 85287-4501.

Municipal effluent: a viable source for environmental flows? Many human actions have altered the hydrological processesthatshape riparian

ecosystems. One change, the release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability

in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for environmental flows, there are numerous

unresolved ecohydrological issues. Here we summarize results of empirical research addressing several aspects of this topic. A statewide spatial

analysis of patterns of effluent generation and release was conducted, revealing that hydrogeomorphic setting influences downstream riparian

response. A field study documented differences in vegetation structure between two effluent-dominated reaches of the Santa Cruz River and a

control river (the San Pedro). Third, a greenhouse study indicated that varying concentrations of nitrogen and phosphorus, emulating levels in

effluent, influenced composition, biomass, and richness of herbaceous plant communities. We conclude that water quantity and water quality

both affect riparian ecosystem development following discharge of effluent into semiarid region streams, with these effects mediated by the

hydrogeomorphic context of the receiving stream. If effluent is to be used for environmental flows, ecological data generated by this and other

studies needs to be integrated into water resource management decision frameworks.

Aguas residuales municipales, ¿una fuente viable para corrientes ambientales? Muchas actividades humanas han alterado los procesos

hidrológicos que forman los ecosistemas ribereños. Un cambio, la liberación de aguas residuales tratadas en cauces de arroyos, ha creado corrientes perennes y ha aumentado los nutrientes disponibles en riachuelos efímeros o intermitentes. Aunque hay beneficios al utilizar el agua

residual tratada para corrientes ambientales, hay varios problemas ecohidrológicos que no se han resuelto. Aquí resumimos los resultados de

la investigación empírica abordando varios aspectos de este tema. Se llevó a cabo un análisis espacial del origen y liberación de las aguas

residuales, revelando que el marco hidrogeomórfico influye la reacción del río abajo. Un trabajo de campo documentó las diferencias de la

estructura de la vegetación entre dos tramos dominados por aguas residuales del río Santa Cruz y un río de control (el San Pedro). Tercero, un

estudio de un invernadero indicó que concentraciones variables de nitrógeno y fósforo, emulando los niveles en aguas residuales, influyeron la

composición, la biomasa y la diversidad de plantas herbáceas. Concluimos que tanto la calidad como la cantidad de agua afectan el desarrollo

del ecosistema ribereño después de la descarga de aguas residuales en arroyos de regiones semiáridas, con estos efectos influidos por el contexto

hidrogeomórfico de los arroyos que las reciben. Si el agua residual se usa para corrientes ambientales, los datos ecológicos generados por este

y otros estudios necesitan integrarse al manejo de los recursos del agua.

WILCOX, CRAIG P.1, R.Lefevre1, Ann M. Lynch2, Mellissa Merrick3, Brent Mitchell4. 1 U.S. Forest Service, Coronado N.F., 300 W Congress,

Tucson AZ 85701, cpwilcox@fs.fed.us and rlefevre@fs.fed.us; 2 U.S. Forest Service, Rocky Mountain Research Station, Tucson AZ 85721,

alynch@fs.fed.us; 3 University of Arizona, School of Natural Resources and the Environment, Tucson AZ 85721, mmerrick@email.arizona.

edu;

4 U.S. Forest Service, Remote Sensing Applications Center, Salt Lake City UT 84119, brentmitchell@fs.fed.us

Pinaleño LiDAR: Evaluating forest health and landscape factors Changes in forest conditions have led to large and severe wildfires and

devastating insect outbreaks in the Pinaleño Mountains of Southeastern Arizona. The risk of additional wildfires and insect outbreaks have

prompted restoration efforts to reduce further threats to forest resilience and provide habitat sustainability for the Mount Graham red squirrel

(MGRS) (Tamiasciurus hudsonicus grahamensis), a federally listed endangered species. Airborne Laser Scanning or LiDAR is a cost effective

means to gather forest structure and biomass information for project planning, implementation and monitoring in remote mountainous terrain.

The Coronado N.F. collected LiDAR data in September 2008 over 85,518 acres to provide current, detailed information on forest structure and

fuels. The derived information is being used to plan, implement, and monitor ecosystem restoration projects and assist in creating a MGRS

habitat model. The LiDAR data has provided continuous non-discrete forest inventory data; mapping of forest biomass, canopy fuel, and forest structure characteristics; mapping of old growth mixed-conifer (a key MGRS habitat); mapping of recent insect and fire disturbances; and

collection of forest inventory data in locations where data is difficult to obtain by traditional forest inventory methods.

Pinaleño LiDAR: Evaluación del equilibrio ecológico del bosque y factores del paisaje. Cambios en las condiciones del bosque han causado

incendios forestales severos y plagas de insectos devastadoras en las montañas Pinaleño del sureste de Arizona. El riesgo de más incendios y

plagas ha motivado esfuerzos de restauración para reducir nuevas amenazas a la resistencia del bosque y proporcionar sostenibilidad del hábitat

de la ardilla roja de Mount Graham (MGRS por sus siglas en inglés, Tamisciurus hudsonicus grahamensis), especie en peligro de extinción en
USDA Forest Service Proceedings RMRS-P-67. 2013 591

Abstracts/ Resúmenes

la norma federal. La detección de rayos láser por sensores remotos o LiDAR (por sus siglas en inglés) en una manera rentable de acumular información sobre la estructura y biomasa del bosque para usarse en la planificación de proyectos y el monitoreo de terrenos remotos montañosos.

En el Bosque Nacional de Coronado, se colectaros datos de LiDAR en septiembre de 2008 en más de 34,400 hectáreas para tener información

actual detallada sobre la estructura del bosque y vegetación combustible. La información derivada se está usando para planificar, implementar

y monitorear proyectos de restauración del ecosistema y para crear un modelo del hábitat de la MGRS. La información de LiDAR ha proporcionado datos de inventario del bosque continuos; mapas de la biomasa forestal, combustible de las copas y características de la estructura del

bosque; mapas de bosques mixto de coníferas antiguos (un hábitat clave para la MGRS); mapas de plagas e incendios recientes; y recolección

de datos de inventario forestal en lugares donde es difícil la toma de datos por métodos tradicionales.

WILSON, MICHAEL1, Joseph Scheer2, and Francisco Molina-Freaner3. 1Drylands Institute, 2509 N. Campbell Avenue, # 405, Tucson, AZ

85719, mwilson@drylands.org; 2Alfred University, 2 Pine Street, Alfred University, Alfred, New York 14843, fscheer@alfred.edu; 3Instituto de

Ecología UNAM, Avenida Colosio s/n esq. Madrid, Colonia Los Arcos, Hermosillo, Sonora C.P. 83250, freaner@unam.mx.

Promoting non-charismatic microfauna: artist and scientists join forces to document moth diversity in the Madrean region. Joseph

Scheer is an artist who for the past 15 years has been using scanning technology to create images of small biological subjects, primarily moths.

Finished prints of phenomenal clarity of up to eight feet in expanse are produced from subjects of sometimes only a few millimeters in size.

These images have generated great interest in the scientific community and it was inevitable that this work would involve the science of the

subjects themselves. Mr. Scheer has teamed with researchers and students in the USA and Mexico to document the moth diversity of the Mexican state of Sonora and has returned from a year’s stay in the state as a Fulbright fellow. This endeavor furthers Mr. Scheer’s artistic practice,

but is also being conducted in a way that is generating enormous amounts of scientifically valuable information. Furthermore, art’s ability to

reach and inspire the public furnishes influential means of raising awareness of science, biodiversity and conservation; in this case on behalf of

organisms that for one reason or another are not considered “charismatic.”

Promoviendo la microfauna no carismática: Artistas y científicos se unen para documentar la diversidad de palomillas en la región 

Madrense. Joseph Scheer es un artista que los últimos 15 años ha usado tecnología de escáners para crear imágenes de sujetos biológicos pequeños, ante todo, palomillas. Se producen imágenes claras de hasta 2.4 m de extensión de sujetos que a veces miden tan sólo milímetros. Estas

imágenes han generado mucho interés en la comunidad científica y era inevitable que esta obra involucrara la ciencia de los sujetos mismos.

Scheer se ha asociado con investigadores y estudiantes de E.U. y México para documentar la diversidad de palomillas en el estado mexicano

de Sonora y ha terminado una estancia de un año en el estado como becario Fulbright. Este esfuerzo no sólo promueve la práctica artística de

Scheer, también se conduce de una manera que genera enormes cantidades de información científica importante. Además, la capacidad del arte

de tocar e inspirar al público proporciona maneras de crear conciencia de las ciencias, la biodiversidad y la conservación; en este caso sobre

organismos, que por una razón u otra, no se consideran “carismáticos”.

WOLFSON, BARB SATINK1, Andrea E. Thode1, Molly E. Hunter1, Peter Z. Fulé1, Alexander Evans5, Jose M. Iniguez6, Donald A. Falk7. 1

Northern Arizona University School of Forestry, Flagstaff, Arizona 86011-5018; 5Forest Guild, Santa Fe, NM 87504; 6USDA Forest Service,

Rocky Mountain Research Station, Flagstaff, Arizona 86001;7University of Arizona, Laboratory of Tree Ring Research, Tucson, AZ 85721.

The Southwest Fire Science Consortium: A new opportunity in fire science and management. TheSouthwestisoneofthemostfire-dominated

regions of the US. Currently, in the Southwest, there are several localized efforts to develop fire science information and to disseminate it to

practitioners on the ground in a practical manner. However, many of these efforts are moving in parallel, without thoughtful interaction among

projects. Managers and scientists are often not aware of each other or of the external resources available. A consortium is needed to bring these

parallel efforts together to be more efficient and inclusive, allowing future fire science issues to be addressed from a broader perspective with

more information, more partners, and more resources. With support from the Joint Fire Science Program (JFSP), we have initiated the Southwest Fire Science Consortium to promote communication and meet fire knowledge needs of scientists and managers. We have organized the

Southwestern Fire Science Consortium around three key questions: (1) What do people need to know? Information needs are assessed through

workshops,surveys, and organization of a community of practice ofwildland fire professionals;(2)What information is already known? Synthesis

of existing science; and (3) What are the key information gaps between what we need to know and what is already known? This question leads

to the identification of critical areas for new research and management experiments. By focusing on these key questions we hope to provide

a mechanism for managers, scientists, and policy makers to interact and share science in ways that can effectively move new information to

management practices and facilitate new research based on management needs.

El Consorcio Suroeste de la Ciencia de Incendios: Una nueva oportunidad en la ciencia y manejo de incendios. El Suroeste de E.U.

es una da las regiones más afectadas por los incendios. Actualmente en el Suroeste, hay esfuerzos localizados para desarrollar información

sobre la ciencia de los incendios y difundirla a facultativos del terreno de una manera práctica. Sin embargo, muchos de estos esfuerzos están

moviéndose paralelamente, sin interacción atenta entre proyectos. Los administradores y científicos casi nunca son conscientes el uno del otro

o de los recursos disponibles. Se necesita un consorcio para juntar estos esfuerzos paralelos para ser más eficaz e inclusivo, y permitir que se

discutan problemas de la ciencia de incendios futuros con una perspectiva amplia, más información, más socios y más recursos. Con el apoyo

del programa conjunto de las ciencias de incendios (JFSP por sus siglas en inglés), hemos empezado el Consorcio Suroeste de las Ciencias de

Incendios para promover la comunicación y enfrentar las necesidades de los administradores y los científicos. Hemos organizado el Consorcio

Suroeste de las Ciencias de Incendios alrededor de estas preguntas claves: (1) ¿Qué es lo que debería saber la gente? Se evalúa la necesidad
592 USDA Forest Service Proceedings RMRS-P-67. 2013

Abstracts/ Resúmenes

de información a través de talleres, estudios y la organización de una comunidad de práctica de profesionales de los incendios forestales; (2)

¿Qué información ya se sabe? La síntesis de la ciencia actual; y (3) ¿Cuáles son las lagunas de información claves entre lo que se debería saber

y lo que ya se sabe? Esta pregunta lleva a la identificación de áreas críticas de nueva investigación y experimentos de manejo. Al centrarnos en

estas preguntas claves esperamos proporcionar un mecanismo para que los administradores, científicos y los que se encargan de formular leyes

interactúen y compartan la ciencia de manera que puedan convertir la información nueva en prácticas de manejo y facilitar nuevos estudios

basados en las necesidades de manejo.

YOUBERG,ANN1, Daniel G. Neary2, KarenA. Koestner2, Peter E. Koestner2. 1Arizona Geological Survey, Tucson,AZ; 2USDAForest Service,

Rocky Mountain Research Station, Flagstaff, AZ.

Post-wildfire erosion in the Chiricahua Mountains. The Horseshoe 2 Fire burned 90,226 ha (222,954 ac) of the Chiricahua Mountains in

the Coronado National Forest of southeast Arizona from May 8 to June 25, 2011. This mountain range in the Madrean Archipelago has been

burned by widespread fires prior to 1890, numerous small fires since 1890, and a more recent 11,129 ha (27,500 ac) Rattlesnake Fire in 1994.

The latter fire resulted in significant erosion during post-fire Monsoon storms that eroded deep gullies (~10 m) in Ward Canyon (upper West

Turkey Creek), and deposited large amounts of sediment in lower channels, particularly in Rucker Creek and in Rucker Lake. The Horseshoe 2

Fire burned approximately 70% of the mountain range with a mosaic of fire severities. Post-fire erosion from floods and debris flows produced

additional erosion and sedimentation that is currently being evaluated through fieldwork and repeat aerial photography. This paper discusses

the magnitude of post-fire erosion and the impacts on channel systems in the Chiricahua Mountains.

La erosión después de incendios en la sierra de Chiricahua. El incendio Horseshoe 2 quemó 90, 226 ha de la sierra de Chiricahua en el Bosque

Nacional Coronado en el sureste de Arizona desde el 8 de mayo hasta el 25 de junio de 2011. Esta cordillera del Archipiélago Madrense se ha

quemado con incendios generalizados desde antes de 1890, numerosos fuegos pequeños después de 1890 y más recientemente el incendio de

Rattlesnake de 1994 que quemó 11,129 ha..El fuego más reciente ocasionó una erosión significativa durante las tormentas del monzón después

del incendio que erosionó barrancos profundos (~10 m) en Ward Canyon (parte alta del río West Turkey Creek) y depositó grandes cantidades

de sedimento en los cauces bajos, especialmente en Rucker Creek y en Rucker Lake. El incendio de Horseshoe 2 quemó aproximadamente

70% de la sierra creando un mosaico con diferente intensidad del fuego. La erosión después del incendio, causada por inundaciones y deslaves,

produjo erosión y sedimentación adicional que actualmente se está evaluando mediante trabajo de campo y fotografía aérea repetida. Este

trabajo discute la magnitud de la erosión que ocurre después de los incendios y el impacto en el sistema de cauces de la sierra de Chiricahua.

ZAMORA-BÁRCENAS,DIANAF.1,NalleliE.Lara-Díaz1,AlejandroGonzález-Bernal2,HelíCoronel-Arellano2, andCarlosA.López-González1. 1Laboratorio de Zoología, Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Av. de las Ciencias S/N, Col. Juriquilla, Santiago de Querétaro, Querétaro, México, C.P. 76230; 2Naturalia Comité para la Conservación de Especies Silvestres A. C., Av. 32 entre Calle 3

y 4 Villa Satélite 3250, Agua Prieta, Sonora, México, C.P. 84200, daj_zb@msn.com.

Population viability of Mexican wolf (Canis lupus baileyi) in northeastern Sonora, Mexico The Mexican Wolf (Canis lupus baileyi) is

considered extinct in Mexico. Population viability analyses (PVA), was performed to evaluate the risk of extinction of a reintroduced population of five Mexican wolves. Other PVAs developed, lack demographic data of free-ranging populations, resulting in unreliable models for the

development of appropriate management strategies. We developed an AVP for wolves to be released in Mexico, with demographic parameters

of the reintroduced population in the United States to serve as a tool for management of wolves in Mexico. The demographic parameters were

obtained from reports from 1999 to 2009. Subsequently we evaluated the population dynamics under different scenarios. It was found that high

adult mortality due to social conflicts is the demographic factor that affects population growth. The scenario in establishing a viable population

of Mexican wolves in Mexico requires reducing mortality rates (20%) and supplementation of two breeding individuals in the first 10 years

after the initial release to withstand more than 50 wolves. It is necessary to improve the social perception of the wolf in the region and educate

people about livestock management practices that avoid conflict with predators, because an antagonistic attitude may limit the establishment

of a viable population.

Viabilidad de la población del lobo mexicano en el noreste de Sonora, México. El lobo mexicano (Canis lupus baileyi) se considera extinto

en México. Se hizo un análisis de viabilidad de población (PVA por sus siglas en inglés) para evaluar el riesgo de extinción de la población

reintroducida con cinco lobos. Previos PVAs no tienen datos demográficos de poblacioneslibres,resultando enmodelos poco fiables para el desarrollo de estrategias de manejo adecuadas. Realizamos un AVP de los lobos a liberarse en México, con parámetros demográficos de la población

reintroducida en Estados Unidos como referencia para el manejo de lobos en México. Los parámetros demográficos se tomaron de informes de

1999 a 2009. Posteriormente se evaluó la dinámica de población bajo diferentes escenarios. La alta mortalidad de adultos por conflictos sociales

es el factor demográfico principal que afecta el crecimiento de la población. Para establecer una población viable del lobo mexicano en México

se requiere reducir las tasas de mortalidad al 20% y agregar dos individuos en edad reproductiva los primeros 10 años después de la liberación

inicial para mantener más de 50 lobos. Es necesario mejorar la percepción social del lobo en la región y educar a la gente sobre prácticas de

manejo de ganado para evitar conflictos con depredadores, ya que una actitud antagónica no permitirá el establecimiento de una población viable.
USDA Forest Service Proceedings RMRS-P-67. 2013 593

Abstracts/ Resúmenes

All abstracts reflect the views of the authors, who are responsible for the facts and accuracy of the information presented herein. 

ZAMORA-BÁRCENAS, DIANA F.1, Helí Coronel-Arellano2, Nalleli E. Lara-Díaz1 and Carlos A. López-González1. 1Laboratorio de Zoología,

Facultad de Ciencias Naturales, UniversidadAutónoma de Querétaro,Av. de las Ciencias S/N, Col.Juriquilla, Santiago de Querétaro, Querétaro,

México, C.P. 76230; 2 Naturalia Comité para la Conservación de Especies Silvestres A. C., Av. 32 entre Calle 3 y 4 Villa Satélite 3250, Agua

Prieta, Sonora, México, C.P. 84200, daj_zb@msn.com.

The small mammal community at preserved and disturbed habitats in the Sierra San Luis, Sonora, Mexico. Species richness and community structure ofsmall mammals has proven an effective method to determine the stability or disturbance between environments. Our aim was

to determine the structure of small mammals at grasslands and scrubland, both conserved or fire affected at the Sierra San Luis during 2011. We

used Sherman traps place along sixteen transects to assess richness and diversity. Shannon diversity was higher for conserved scrubland (H’ =

1.67) and grassland (H’ = 0.70) without fires. The most abundant species were Peromyscus eremicus, P. boylii and Neotoma albigula, occurred

at the scrubland. Jaccard diversity generated two groups with a similarity of 33% and a correlation coefficient of 0.85. Although there is not a

high turnover rate between habitats, there is a trend to have abundant generalist species at sites in response to fire.

Comunidad de mamíferos pequeños en hábitats conservados y modificados en la Sierra San Luis, Sonora, México. La riqueza de especies y la estructura de comunidades de mamíferos pequeños ha demostrado ser un método efectivo para determinar la estabilidad y el disturbio

del entorno. Nuestro objetivo fue determinar la estructura de la comunidad de mamíferos pequeños del pastizal y matorral, tanto conservados

como afectados por incendios, en la sierra San Luis durante el 2011. Usamos trampas Sherman ubicadas en 16 transectos para evaluar la riqueza

y diversidad. La diversidad de Shannon fue mayor en los matorrales conservados (H’= 1.67) y pastizal sin incendios (H’= 0.70). Las especies

más abundantes fueron Peromyscus eremicus, P. boylii y Neotama albigula en el matorral. La diversidad de Jaccard generó dos grupos con

una semejanza de 33% y el coeficiente de correlación de 0.85. Aunque no se registró un volumen alto entre hábitats, hay una tendencia a tener

especies generalistas abundantes en los sitios quemados.

ZYLSTRA, ERIN R.1, Robert J. Steidl1, and Cristina A. Jones2. 1School of Natural Resources and the Environment, University of Arizona, 325

Biological Sciences East, Tucson, AZ 85721; 2Nongame Branch, Arizona Game and Fish Department, 5000 W. Carefree Highway, Phoenix,

AZ 85086-5000.

Drought decreases survival of Sonoran desert tortoises in Arizona. Decisions relevant to conservation of rare and threatened species can be

hampered by our inability to differentiate between short-term, local fluctuationsin population attributes and long-term, widespread declinesthat

can affect persistence. We used 22 years of capture-recapture data for Sonoran desert tortoises (Gopherus agassizii) collected from 15 locations

across their geographic range in Arizona to evaluate environmental factors associated with spatial and temporal variation in adult survival.

Rates of annual survival were generally high (mean = 0.92, SE = 0.01), as expected for a long-lived species, but decreased with proximity to

human habitation. Survival of adults also varied markedly in response to drought; when drought severity was high, survival of tortoises was

reduced, particularly in the most arid parts of their range. In three well-studied populations, survival of adults was markedly lower during one

survey interval (0.77-0.81) than all other intervals (0.93-0.98); these periods of reduced survival coincided with periods of extreme drought. If

drought frequency or severity increases as predicted by several climate-change models, recovery and persistence of tortoise populations in this

region could be affected adversely.

La sequía disminuye la supervivencia de la tortuga del desierto Sonorense en Arizona. Las decisiones importantes de conservación de

especies raras o amenazadas pueden dificultarse por nuestra incapacidad para diferenciar entre variaciones locales por períodos cortos de

los atributos de la población, y bajas generalizadas a largo plazo que pueden afectar su persistencia. Usamos 22 años de datos de captura y

recaptura de la tortuga del desierto Sonorense (Gopherus agassizii) colectados en 15 lugares de su rango geográfico en Arizona, para evaluar

los factores ambientales asociados con la variación espacial y temporal en la supervivencia de adultos. Las tasas anuales de supervivencia en

general fueron altas ( media = 0.92, SE= 0.01), como se espera en especies de vida larga, pero disminuyen cerca de habitaciones humanas. La

supervivencia de adultos también varió notablemente en respuesta a sequías; cuando la intensidad de la sequía fue alta, la supervivencia de la

tortuga disminuyó, particularmente en las partes más áridas de su rango. En tres poblaciones bien estudiadas, la supervivencia de adultos fue

notablemente más baja durante un intervalo del estudio (0.77-0.81) que todos los otros intervalos (0.93-0.98); estos periodos de supervivencia

reducida coincidieron con periodos de sequía extrema. Si aumenta la frecuencia o la severidad de las sequías como predicen los modelos de

cambio climático, se puede afectar negativamente la recuperación y la persistencia de la población de tortuga en esta región.

CAUTION:

PESTICIDES

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not 

contain recommendations for their use, nor does it imply that the uses 

discussed here have been registered. All uses of pesticides must be 

registered by appropriate State and/or Federal agencies before they 

can be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals, 

desirable plants, and fish or other wildlife—if they are not handled or 

applied properly. Use all pesticides selectively and carefully. Follow 

recommended practices for the disposal of surplus pesticides and 

pesticide containers.

The use of trade or firm names in this publication is for reader information and does 

not imply endorsement by the U.S. Department of Agriculture of any product or service.
The Rocky Mountain Research Station develops scientific information 

and technology to improve management, protection, and use of the 

forests and rangelands. Research is designed to meet the needs of 

the National Forest managers, Federal and State agencies, public and 

private organizations, academic institutions, industry, and individuals. 

Studies accelerate solutions to problems involving ecosystems, range, 

forests, water, recreation, fire, resource inventory, land reclamation, 

community sustainability, forest engineering technology, multiple use 

economics, wildlife and fish habitat, and forest insects and diseases. 

Studies are conducted cooperatively, and applications may be found 

worldwide. For more information, please visit the RMRS web site at: 

www.fs.fed.us/rmrs.

Station Headquarters

Rocky Mountain Research Station

240 W Prospect Road

Fort Collins, CO 80526

(970) 498-1100

Research Locations

The U.S. Department of Agriculture (USDA) prohibits discrimination against its 

customers, employees, and applicants for employment on the bases of race, 

color, national origin, age, disability, sex, gender identity, religion, reprisal, and 

where applicable, political beliefs, marital status, familial or parental status, 

sexual orientation, or all or part of an individual’s income is derived from any 

public assistance program, or protected genetic information in employment 

or in any program or activity conducted or funded by the Department. (Not all 

prohibited bases will apply to all programs and/or employment activities.) For 

more information, please visit the USDA web site at: www.usda.gov and click on 

the Non-Discrimination Statement link at the bottom of the page.

Reno, Nevada

Albuquerque, New Mexico

Rapid City, South Dakota

Logan, Utah

Ogden, Utah

Provo, Utah

Flagstaff, Arizona

Fort Collins, Colorado

Boise, Idaho

Moscow, Idaho

Bozeman, Montana

Missoula, Montana

To learn more about RMRS publications or search our online titles:

www.fs.fed.us/rm/publications

www.treesearch.fs.fed.us

Federal Recycling Program Printed on Recycled Paper