U.S. Geological Survey

Southwest Biological Science Center

2255 N. Gemini Drive

Flagstaff, AZ 86001

 

In cooperation with the University of Arizona, School of Natural Resources 

Vascular Plant and Vertebrate Inventory of 

Saguaro National Park, Rincon Mountain District 

Edited by Brian F. Powell, William L. Halvorson, and Cecilia A. Schmidt 

Open-File Report 2006-1075 

November 2006

 

U.S. Department of the Interior 

U.S. Geological Survey 

National Park Service 

USGS Southwest Biological Science Center 

Sonoran Desert Research Station 

University of Arizona 

School of Natural Resources 

125 Biological Sciences East 

Tucson, Arizona 85721
U.S. Department of the Interior 

DIRK KEMPTHORNE, Secretary 

U.S. Geological Survey

Mark Myers, Director

U.S. Geological Survey, Reston, Virginia: 2006 

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 Suggested Citation 

Powell, B. F, W. L. Halvorson, and C. A. Schmidt. Vascular Plant and Vertebrate Inventory of Saguaro 

National Park, Rincon Mountain District. OFR 2006-1075. U.S. Geological Survey, Southwest Biological 

Science Center, Sonoran Desert Research Station, University of Arizona, Tucson, AZ. 

Cover photo: Rincon Creek (left); Sonoran Desertscrub with Tanque Verde ridge in the background 

(right). Photograph by Greg Levandowski. 

Any use of trade, product, or firm names is for descriptive purposes only and does not imply 

endorsement by the U.S. Government. 

 Printed on recycled paper

ii 
Editors and Authors Authors

Brian F. Powell and Cecilia A. Schmidt Aaron D. Flesch

School of Natural Resources School of Natural Resources 

125 Biological Sciences East, Building 43 325 Biological Sciences East, Building 43 

The University of Arizona The University of Arizona 

Tucson, AZ 85721 Tucson, AZ 85721 

William L. Halvorson Don E. Swann 

USGS SBSC Sonoran Desert Research Station Saguaro National Park 

125 Biological Sciences East, Building 43 3693 South Old Spanish Trail 

The University of Arizona Tucson, AZ 85730 

Tucson, AZ 85721 

U.S. Geological SBSC Survey Sonoran Desert Research Station Personnel 

Charles van Riper III, Station Leader 

William L. Halvorson, Research Ecologist 

Cecil R. Schwalbe, Ecologist 

Michael R. Kunzmann, Ecologist (Emeritus) 

Kathryn Thomas, Ecologist 

Pamela Nagler, Physical Scientist 

Phil Rosen, Ecologist 

Program and Expertise Areas of USGS and UA Personnel

Administration & Outreach 

Jennifer Meador 

Wendy Parrish 

Emily Sherbrooke 

Charles van Riper III 

Avian Ecology 

Claire Crow 

Glenn Johnson 

Chris O’Brien 

Brian Powell 

Charles van Riper III 

Data Management 

Brent Sigafus 

Ecology of Amphibians & Reptiles 

Kevin Baker 

Cristina Jones 

Dave Prival 

Phil Rosen 

Cecil Schwalbe 

Brent Sigafus 

Fire Management 

Dennis Suhre 

Cori Dolan 

James Feldmann 

Bill Halvorson 

Invasive Species Research 

Patricia Guertin 

Jim Malusa 

Phil Rosen 

Cecil Schwalbe 

Brent Sigafus 

Dennis Suhre 

Kathryn Thomas 

Inventory & Monitoring 

Patricia Guertin 

Bill Halvorson 

Pamela Nagler 

Brian Powell 

Cecilia Schmidt

Vegetation Mapping & Ecology 

Patricia Guertin 

Bill Halvorson 

Jim Malusa 

Kathryn Thomas 

USGS Southwest Biological Science Center http://sbsc.wr.usgs.gov 

USGS Southwest Biological Science Center, Sonoran Desert Research Station http://sbsc.wr.usgs.gov/sdrs

iii 
iv 
Table of Contents 

Report Dedication ..................................................................................................................................... xi

Acknowledgements.................................................................................................................................. xiii

Executive Summary ..................................................................................................................................xv

Chapter 1: Introduction to the Inventories...............................................................................................1

Project Overview...................................................................................................................................1

Rerport Format and Data Organization .................................................................................................2

Verification and Assessment of Results................................................................................................3

Sampling Design ...................................................................................................................................4

Chapter 2: Park Overview..........................................................................................................................7

Park Area and History ...........................................................................................................................7

Natural Resources Overview.................................................................................................................7

Natural Resource Management Issues.................................................................................................12

Chapter 3: Plant Inventory .....................................................................................................................15

Previous and Ongoing Research..........................................................................................................15

Methods...............................................................................................................................................15

Results.................................................................................................................................................19

Inventory Completeness......................................................................................................................21

Discussion ...........................................................................................................................................23

Chapter 4: Amphibian and Reptile Inventory.......................................................................................27

Previous Research ...............................................................................................................................27

Methods...............................................................................................................................................27

Results.................................................................................................................................................32

Inventory Completeness......................................................................................................................40

Discussion ...........................................................................................................................................40

Chapter 5: Bird Inventory.......................................................................................................................45

Previous Research ...............................................................................................................................45

Methods...............................................................................................................................................45

Results.................................................................................................................................................52

Inventory Completeness......................................................................................................................57

Discussion ...........................................................................................................................................62

Chapter 6: Mammal Inventory ...............................................................................................................69

Previous and Ongoing Research..........................................................................................................69

Methods...............................................................................................................................................69

Results.................................................................................................................................................78

Inventory Completeness......................................................................................................................81

Discussion ...........................................................................................................................................84

Chapter 7: Literature Cited ....................................................................................................................89

v 
List of Tables

Table 1. Summary of vascular plant and vertebrate inventories at Saguaro National Park, Rincon 

Mountain District, 1999–2005. ..................................................................................................xv

Table 1.1. Museums that were queried in 1998 for vertebrate voucher specimens with “Arizona” and 

“Saguaro National Park” and “National Monument” in the collection location.........................3

Table 2.1. Average monthly climate data for Manning Camp (high elevation), Saguaro National Park, 

Rincon Mountain District, 1994–2004........................................................................................9

Table 2.2. Average monthly climate data for the University of Arizona (low elevation; the closest climate 

monitoring station to Saguaro National Park, Rincon Mountain District) 1894–2004...............9

Table 4.1. Characteristics of three major active survey methods used during surveys for herpetofauna, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002..........................................27

Table 4.2. Herpetofaunal survey effort by year, Saguaro National Park, Rincon Mountain District, 2001 

and 2002....................................................................................................................................29

Table 4.3. Environmental factors considered when modeling variation in relative abundance of species 

and species groups and species richness of herpetofauna, using stepwise multiple linear 

regression, Saguaro National Park, Rincon Mountain District, 2001 and 2002. .....................30

Table 4.4. Number of animals and species detected per hour during herpetofaunal surveys by year and 

survey method, Saguaro National Park, Rincon Mountain District, 2001 and 2002. ...............33

Table 4.5. Relative abundance (mean + SE; no./ha/hr) of herpetofauna detected during intensive surveys 

in spring (9 April – 24 May) along focal point-transects by elevation strata, Saguaro National 

Park, Rincon Mountain District, 2001. ...................................................................................34

Table 4.6. Relative abundance (mean + SE; no./ha/hr) of herpetofauna detected during intensive surveys 

along random transects (n = 7) surveyed in both spring (9 April – 8 May) and summer (18 – 

31 July), Saguaro National Park, Rincon Mountain District, 2001...........................................35

Table 4.7. Environmental factors that explained relative abundance (no./ha/hr) of species (with >15 

observations), species groups, and species richness of lizards and snakes detected during 

intensive surveys, Saguaro National Park, Rincon Mountain District, spring 2001. ................36

Table 4.8. Relative abundance (mean + SE; no./10 hrs) of herpetofauna detected during extensive 

surveys (n = 85), by elevation strata, Saguaro National Park, Rincon Mountain District, 2001 

and 2002. .................................................................................................................................37

Table 4.9. Relative abundance (no./hr) of herpetofauna detected during road surveys, Saguaro National 

Park, Rincon Mountain District, 2001 and 2002.......................................................................38

Table 5.1. Characteristics of the three major VCP survey types for birds, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. ...........................................................................................46

Table 5.2. Summary of bird survey effort, Saguaro National Park, Rincon Mountain District, 2001–

2003. ........................................................................................................................................49

Table 5.3. Bird measures by community type and compared using Tukey-Kramer multiple pairwise 

procedure, Saguaro National Park, Rincon Mountain District, 2001 and 2002. .....................54

Table 5.4. Relative abundance (mean + SD) by community type for birds recorded during repeat-visit 

VCP surveys, Saguaro National Park, Rincon Mountain District, 2001 and 2002. ................55

Table 5.5. Mean relative abundance of birds from reconnaissance VCP surveys by strata and transect, 

Saguaro National Park, Rincon Mountain District, 2002..........................................................58

Table 5.6. Relative abundance (mean + SE) of birds from line-transect surveys, Saguaro National Park, 

Rincon Mountain District, 2002 and 2003................................................................................60

Table 5.7. Mean relative abundance of birds from nocturnal surveys by elevation strata and transect, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002..........................................61

vi 
Table 5.8. Number of breeding behavior observations for birds from all survey types, Saguaro National 

Park, Rincon Mountain District, 2001 and 2002. ....................................................................61

Table 6.1. Summary of small-mammal trapping effort, Saguaro National Park, Rincon Mountain District, 

2001 and 2002. ........................................................................................................................72

Table 6.2. Summary of infrared-triggered camera effort, Saguaro National Park, Rincon Mountain 

District, 1999–2005. ................................................................................................................77

Table 6.3. Relative abundance of small mammals by strata and site type (R = random [focal-point 

transects]; NR = non-random), Saguaro National Park, Rincon Mountain District, 2001 and 

2002. ........................................................................................................................................79

Table 6.4. Results of netting for bats, by elevation strata, site, and visit, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. ...........................................................................................80

Table 6.5. Number of photographs of mammals from infrared-triggered photography by elevation strata, 

Saguaro National Park, Rincon Mountain District, 1999 – 2005..............................................81

vii 
List of Figures 

Figure 1.1. Layout of 1-km focal-point transects showing layout of amphibian and reptile plots (C), 

small-mammal trapping grids (D), and bird survey stations (E).................................................5

Figure 2.1. Location of the two districts of Saguaro National Park in southern Arizona. ...........................8

Figure 2.2. Aerial photograph showing major features of Saguaro National Park, Rincon Mountain 

District.........................................................................................................................................9

Figure 2.3. Comparison of monthly weather data during the time of the majority of the inventory effort 

(2001–2003) compared to the mean (1994–2004 for Manning Camp, 1894–2004 for 

University of Arizona; thick solid line in all figures), Saguaro National Park.. .......................10

Figure 2.4. Diagram of the major vegetation communities of the Santa Catalina Mountains, adjacent to 

the Rincon Mountains (from Whittaker and Niering 1965). ...................................................11

Figure 3.1. Locations of general botanizing collection sites, Saguaro National Park, Rincon Mountain 

District, 2001 and 2002.............................................................................................................17

Figure 3.2. Layout of a modified-Whittaker plot, Saguaro National Park, Rincon Mountain District, 

2001...........................................................................................................................................17

Figure 3.3. Locations of modified-Whittaker plots and point-intercept transects (line transect), Saguaro 

National Park, Rincon Mountain District, 2001........................................................................18

Figure 3.4. Typical layout of point-intercept transects, Saguaro National Park, Rincon Mountain District, 

2001...........................................................................................................................................18

Figure 3.5. Summary (mean + SD) of data from point-intercept transects by community type and height 

class, Saguaro National Park, Rincon Mountain District, 2001................................................21

Figure 3.6. Percent (mean + SD) ground cover from point-intercept transects by community type, 

Saguaro National Park, Rincon Mountain District, 2001..........................................................22

Figure 4.1. Layout of herpetofauna survey plots along focal-point transects, Saguaro National Park, 

Rincon Mountain District, 2001. .............................................................................................28

Figure 4.2. Locations of intensive and extensive survey sites for herpetofauna, Saguaro National Park, 

Rincon Mountain District, 2001 and 2002................................................................................29

Figure 4.3. Species accumulation curve for herpetofauna surveys, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. .........................................................................................39

Figure 5.1. Locations of VCP survey stations (random [focal-point transects], non-random, and 

reconnaissance), Saguaro National Park, Rincon Mountain District, 2001 and 2002. .............47

Figure 5.2. Location of section breaks for non-breeding season (winter) bird transects and nocturnal 

survey stations, Saguaro National Park, Rincon Mountain District, 2001 and 2002. ...............48

Figure 5.3. Dendrogram of bird community groups from Ward’s hierarchical cluster analysis, Saguaro 

National Park, Rincon Mountain District, 2001 and 2002. .....................................................53

Figure 5.4. Species accumulation curve for all survey methods for birds, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. .........................................................................................63

Figure 6.1. Layout of small-mammal trapping grids along focal-point transects, Saguaro National 

 Park, 2001. ..............................................................................................................................70

Figure 6.2. Detailed layout of small-mammal trapping grids at Saguaro National Park, 

 2001 and 2002. .......................................................................................................................70

Figure 6.3. Locations of random (focal-point transect) small-mammal trapping sites, pitfall traps for 

shrews, and bat trapping stations, Saguaro National Park, Rincon Mountain District, 

 2001 and 2002. ..........................................................................................................................71

Figure 6.4. Locations of non-random small-mammal trapping sites, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. ...........................................................................................72

viii 
Figure 6.5. Locations of non-random infrared-triggered cameras, Saguaro National Park, Rincon 

Mountain District, 2000-2005...................................................................................................75

Figure 6.6. Locations of random infrared-triggered cameras, Saguaro National Park, Rincon Mountain 

District, 2000-2005. ..................................................................................................................76

Figure 6.7. Example of three-camera placement at one of the random points, Saguaro National Park, 

Rincon Mountain District, 2001 and 2002. .............................................................................76

Figure 6.8. Typical configuration for an active infrared-triggered camera system. .................................77

Figure 6.9. Species accumulation curve for small-mammal trapping by elevation stratum, Saguaro 

National Park, Rincon Mountain District, 2001 and 2002. .....................................................82

Figure 6.10. Species accumulation curve for bat trapping, Saguaro National Park, Rincon Mountain 

District, 2001 and 2002. ..........................................................................................................82

Figure 6.11. Species accumulation curve for infrared-triggered cameras, Saguaro National Park, Rincon 

Mountain District, 1999-2005. ................................................................................................83

ix 
List of Appendices 

Appendix A. List of plant species that were observed (O) or collected (X) at Saguaro National Park, 

Rincon Mountain District. .......................................................................................................99

Appendix B. List of amphibian and reptile species observed or documented at Saguaro National Park, 

Rincon Mountain District by UA inventory personnel (total number of observations; 2001-

2002) or by other survey efforts or lists. ...............................................................................129

Appendix C. List of bird species observed at Saguaro National Park, Rincon Mountain District by UA 

inventory personnel (2001-2003) or by other survey efforts or lists. ....................................131

Appendix D. Number of observations of mammal species by University of Arizona and Saguaro National 

Park Inventory personnel by survey type, Saguaro National Park, Rincon Mountain District, 

2001 and 2002 (small mammals, bats, and observations of all taxa) and 1999-2005 (infraredtriggered photography). .........................................................................................................137

Appendix E. Vertebrate specimen and photograph vouchers collected by University of Arizona or park 

personnel, Saguaro National Park, Rincon Mountain District, 1997–2002. .........................139

Appendix F. List of existing voucher specimens collected prior to this inventory effort. .....................142

Appendix G. Mean frequency of detection of birds, by community type and transect, recorded during 

repeat-visit VCP surveys, Saguaro National Park, Rincon Mountain District, 

 2001 and 2002. ......................................................................................................................146

Appendix H. Mean density (number of stems/hectare) of large trees and potential cavity-bearing plants at 

non-random, repeat-visit VCP stations, Saguaro National Park, Rincon Mountain District, 

2001 and 2002. ...................................................................................................................... 151

Appendix I. Details of small-mammal trapping effort, Saguaro National Park, Rincon Mountain District, 

2001 and 2002.........................................................................................................................152

Appendix J. Summary of field effort for bats, Saguaro National Park, Rincon Mountain District, 2001 

and 2002. ...............................................................................................................................152

Appendix K. Details of infrared-triggered camera effort and results, Saguaro National Park, Rincon 

Mountain District, 1999-2005. ..............................................................................................153

x 
Report Dedication 

Eric Wells Albrecht 

1970-2004

This report, as others in the series, is dedicated to Eric’s life and work; he was an extraordinary ecologist, 

community member, father, partner, and friend. Eric was co-coordinator of the University of Arizona 

(UA) biological inventory and monitoring program from 2002 until his sudden and unexpected death on 

September 20, 2004. Eric was near completion of his MS degree in Wildlife Conservation from the UA, 

which was awarded posthumously in November 2004. In his last year, Eric spearheaded projects to 

investigate the efficiency of current monitoring programs; he was passionate about using the best 

available information to guide vertebrate monitoring efforts in the region. He is survived by his partner, 

Kathy Moore, and their two young children, Elizabeth and Zachary. We hope that the lives of his 

children will be enriched by Eric’s hard work on behalf of the national parks in the Sonoran Desert 

Network. 

Don Swann dedicates the mammal chapter to Lowell Sumner for his elegant study of mammals in 

the Rincon Mountains in 1950-1951 and for his life-long dedication to biological research in U.S. 

National Parks; and to Russell Davis and Ronnie Sidner for their significant and on-going contributions to 

our understanding of mammals in Saguaro National Park. 

xi 
xii 
Acknowledgements 

Thanks to Saguaro National Park Superintendent Sarah Craighead, Chief of Science and 

Resource Management Meg Weesner, and biologists Natasha Kline and Don Swann for providing 

leadership and administrative support for this project. Other park staff who assisted our project 

included Matt Daniels, Mark Holden, Bob Lineback, Todd Nelson, Kathy Schon, Mike Ward, 

and Jim Williams. This project was funded by the National Park Service (NPS) Inventory and 

Monitoring program (I&M) and resulted from the collaboration of many people at the University 

of Arizona (UA), NPS, and U.S. Geological Survey (USGS). Administration of the project was 

facilitated by the Desert Southwest and Colorado Plateau Cooperative Ecosystem Studies Units 

(CESUs). The Southern Arizona Office of the NPS assisted with the development of the original 

study plan that led directly to initiation of this project. Additional support (monetary and inkind) 

for infrared-triggered photography was provided by the Western National Parks Association, 

Friends of Saguaro National Park, and the UA Undergraduate Biology Internship Program. 

Andy Hubbard at the Sonoran Desert Network I&M program has been a great advocate 

of our program. He also provided funds for Don Swann to work on this report. Kathy Davis, 

Superintendent of Tuzigoot and Montezuma Castle national monuments played an instrumental 

role in this project by providing important early initiative. Larry Norris at the Desert Southwest 

CESU has provided strong support for our program and spent considerable time and effort 

providing clear and timely administrative assistance. Matt Goode, Don Swann, and Dale Turner 

provided much of the early planning for this project; we are indebted to their vision and work. 

Eric Albrecht, to whom this report is dedicated, was an outstanding spokesperson and leader of 

the program; he was an invaluable member of the team and his contributions are sorely missed. 

We thank a core group of dedicated field biologists who collected a wealth of data at 

Saguaro National Park: Greta Anderson, Theresa DeKoker, Sky Jacobs, Shawn Lowery, Meg 

Quinn, Rene Tanner, Dale Turner, and Emily Willard (plants); Dan Bell, Kevin Bonine, James 

Borgmeyer, Matt Goode, Dave Prival, and Mike Wall (amphibians and reptiles); Eric Albrecht, 

Gavin Beiber, Aaron Flesch, Chris Kirkpatrick, and Gabe Martinez (birds); Clare Austin, Eric 

Albrecht, Mike Chehoski, Ryan Gann, Michael Olker, Neil Perry, Jason Schmidt, Ronnie Sidner, 

Mike Sotak, Albi von Dach, Michael Ward, and Sandy Wolf (mammals). We are appreciative of 

the following people, many of whom never ventured into the field, but whose work in the office 

made the field effort successful: Debbie Angell, Jennifer Brodsky, Chuck Conrad, Louise Conrad, 

Brian Cornelius, Taylor Edwards, Carianne Funicelli, Marina Hernandez, Colleen McClain, 

Heather McClaren, Lindsay Norpel, Ryan Reese, Jill Rubio, Brent Sigafus, Taffy Sterpka, Jenny 

Treiber, Zuleika Valdez, Alesha Williams, and Erin Zylstra. Pam Anning, Kristen Beaupre, and 

Matthew Daniels assisted with database design. Pam Anning also provided the maps for this 

report. Additional administrative support was provided by Valery Catt, Jenny Ferry, Andy 

Honaman, Terri Rice, and especially Cecily Westphal of the School of Natural Resources at the 

UA. Special thanks to Pam Anning, Lisa Carder, and Kathleen Docherty for their years of hard 

work on all aspects of the project. 

Technical support was graciously provided by the following experts: Dan Austin, 

Michael Chamberland, Phil Jenkins, and Charlotte and John Reeder at the UA Herbarium; Tom 

Huels of the UA ornithology collection; George Bradley of the UA herpetology collection; and 

Yar Petryszyn and Melanie Bucci of the UA mammal collection. Thanks to Sharon Megdal and 

Peter Wierenga, the current and former directors, respectively, of the UA Water Resources 

Research Center, and all their staff. Thanks to Mau-Crimmins et al. (2005) and Sprouse et al. 

(2002) for use of their background information on the park and Aaron Flesch (Flesch 2001) for 

use of some of his discussion in the mammal chapter. We received helpful reviews of earlier 

versions of this report from Danielle Foster, Natasha Kline, Jeff Lovich, Theressa MauCrimmins, Larry Norris, Cecil Schwalbe, Don Swann, and Meg Wessner. 

xiii 
xiv 
This report summarizes the results of the 

first comprehensive inventory of plants and 

vertebrates at the Rincon Mountain District 

(RMD) of Saguaro National Park, Arizona. 

From 2001 to 2003 we surveyed for vascular 

plants and vertebrates (amphibians, reptiles, 

birds, and mammals) at the district to document 

the presence of species within its boundaries. 

Park staff also surveyed for medium and large 

mammals using infrared-triggered cameras 

from 1999 to 2005. This report summarizes the 

methods and results of these two efforts. Our 

spatial sampling design was ambitious and was 

one of the first of its kind in the region to colocate study sites for vegetation and vertebrates 

using a stratified random design. We also chose 

the location of some study sites non-randomly 

in areas that we thought would have the highest 

species richness. Because we used repeatable 

study designs and standardized field methods, 

these inventories can serve as the first step in a 

biological monitoring program for the district. 

We also provide an important overview of most 

previous survey efforts in the district. We use 

data from our inventory and other surveys to 

compile species lists and to assess inventory 

completeness. 

With the exception of plants, our 

survey effort was the most comprehensive ever 

undertaken in the district. We recorded a total 

of 801 plant and vertebrate species, including 

50 species not previously found in the district 

(Table 1) of which five (all plants) are non-native 

species. Based on a review of our inventory and 

past research at the district, there have been a 

total of 1,479 species of plants and vertebrates 

found there. We believe inventories for all 

taxonomic groups are nearly complete. In 

particular, the plant, amphibian and reptile, and 

mammal species lists are the most complete of 

any comparably large natural area of the “sky 

island” region of southern Arizona and adjacent 

Mexico. 

For each taxon-specific chapter 

we discuss patterns of species richness and 

environmental determinants of these patterns. 

For all groups except medium and large 

mammals, the low elevation stratum (<4,000 

feet) contained the highest species richness, after 

accounting for differences in survey effort among 

strata. This is consistent with known patterns 

of species richness in the sky island mountain 

ranges. Using data on relative abundance for 

plants and birds, we were able to identify a 

number of distinct ecological communities, which 

were consistent with known patterns in the sky 

islands. 

Our review of species lists and park 

records reveals that the district has lost species, 

particularly plants and mammals, in the past few 

decades. Because of the district’s close proximity 

to the rapidly growing city of Tucson, there are 

a number of development-related threats that 

could cause additional species loss or decline in 

abundance of some species. In particular, the 

increasing groundwater pumping near Rincon 

Creek, the most species-rich area in the park, is 

likely to impact the unique riparian vegetation 

and animals of that area. We discuss this and 

other demands on the ecological integrity of 

the district. We also recommend additional 

inventory, monitoring, and research studies. 

Executive Summary

xv

Table 1. Summary of vascular plant and vertebrate inventories at Saguaro National Park, Rincon Mountain District, 

1999–2005. 

UA inventory

Taxonomic group

Number of 

species recorded

Number of new species 

added to district list 

Number of 

non-native species

Total number of species 

on district list

Plants 523 39 78 1,162

Amphibians and Reptiles 46 0 2 56

Birds 173 10 3 198

Mammals 59 1 3 63

Totals 801 50 86 1,479
xvi
Project Overview

Inventory: A point-in-time effort to document the 

resources present in an area. 

In the early 1990s, responding to criticism that 

it lacked basic knowledge of natural resources 

within parks, the National Park Service (NPS) 

initiated the Inventory and Monitoring Program 

(I&M) to detect long-term changes in biological 

resources (NPS 1992a). At the time of the 

program’s inception, basic information, including 

lists of plants and animals, was absent or 

incomplete for most park units (Stohlgren et al. 

1995b).

Species inventories have both direct and 

indirect value for management of the park and are 

an important first step in long-term monitoring. 

Species lists are not only useful in resource 

interpretation and facilitating visitor appreciation 

of natural resources, but are also critical for 

making management decisions. Knowledge of 

which species are present, particularly sensitive 

species, and where they occur provides for 

informed planning and decision-making (e.g., 

locating new facilities). Thorough biological 

inventories provide a basis for choosing 

parameters to monitor and can provide baseline 

data for monitoring ecological populations and 

communities. Inventories can also test sampling 

designs, field methods, and data collection 

protocols, and provide estimates of variation that 

are essential in prospective power analysis. 

Goals

The purpose of this study was to complete basic 

inventories for vascular plants and vertebrates 

at the Rincon Mountain District (RMD) of 

Saguaro National Park. This effort was part of a 

larger biological inventory of eight NPS units in 

southern Arizona and southwestern New Mexico 

(Davis and Halvorson 2000, Powell et al. 2004, 

2005). Our goals were to: 

(1) Conduct field surveys to document at 

least 90% of all species of vascular plants 

and vertebrates expected to occur at the 

district. 

(2) Use repeatable sampling designs and 

survey methods that allow estimation 

of parameters of interest (e.g., relative 

abundance).

(3) Compile historic occurrence data for all 

species of vascular plants and vertebrates 

from three sources: museum records 

(specimen vouchers), previous studies, 

and park records. 

(4) Create resources useful to park managers, 

including detailed species lists, maps 

of study sites, and high-quality digital 

images for use in resource interpretation 

and education. 

The bulk of our effort addressed the 

first two goals. To maximize efficiency (i.e., the 

number of species recorded by effort) we used 

field methods designed to detect multiple species. 

We did not undertake single-species surveys for 

threatened or endangered species. This report 

supersedes results reported in Powell et al. (2002 

and 2003). 

Administrative History

The original study plan for this project was 

developed, and an inventory of one park unit 

(Tumacácori National Historical Park) was 

completed, through a cooperative agreement 

among NPS, University of Arizona (UA), and the 

United States Geological Survey (USGS). This 

project was funded through Task Agreements 

UAZ-03, UAZ-05, and UAZ-06 (under the 

Colorado Plateau Cooperative Ecosystems 

Studies Unit [CESU] cooperative agreement 

number 1200-99-009). NPS thereafter obligated 

additional funds through the Colorado Plateau 

CESU (UAZ-07) and the Desert Southwest 

CESU (cooperative agreement number CA1248-

00-002, reference UAZ-39, UAZ-77, UAZ-87, 

UAZ-97, and UAZ-128) for administration and 

management of the biological inventories. 

Chapter 1: Introduction to the Inventories 

Brian F. Powell, Cecilia A. Schmidt, and William L. Halvorson

1
Report Format and Data Organization

Unlike others in the series, each taxonspecific chapter in this report has separate 

authorship. As such there are some differences 

in the organization and content of each chapter. 

Appendices related to each chapter are attributed 

to the respective author(s). We organized a single 

literature cited chapter at the end of the report. 

In the text, we report both common and 

scientific names for plants, and for vertebrates 

we report only common names (listed in 

phylogenetic sequence in tables) unless we 

reference a species that is not listed later in an 

appendix; in this case, we present both common 

and scientific names. For each taxonomic 

group we include an appendix of all species 

that we recorded in the district (Appendices 

A–D). In the amphibian and reptile and mammal 

chapters we review species that were likely 

or confirmed to have been present historically 

or that we suspect are currently present and 

may be recorded with additional survey effort. 

Scientific and common names used throughout 

this document are current according to accepted 

authorities for each taxonomic group: Integrated 

Taxonomic Information System (ITIS 2005) and 

the PLANTS database (USDA 2005) for plants; 

Stebbins (2003) for amphibians and reptiles; 

American Ornithologists’ Union (AOU 1998, 

2003) for birds; and Baker et al. (2003a) for 

mammals. We recognize that the designation of 

a plant as “non-native” using the aforementioned 

lists may lead to the misclassification of some 

species, because these lists indicate only species 

status in North America as a whole, not regions 

within the continent. Therefore, our flora 

underestimates the number of non-native species, 

but because no authoritative list of non-native 

species exists for the region, we believe that use 

of these lists is justified.

Spatial Data

Most spatial data are geographically referenced 

to facilitate mapping of study plots and locations 

of plants or animals. Coordinates were stored 

in the Universal Transverse Mercator (UTM) 

projection (Zone 12), using the North American 

Datum of 1983 (NAD 83). We recorded UTM 

coordinates using hand-held Garmin E-Map®

Global Positioning System (GPS) units (Garmin 

International Incorporated, Olathe, KS; horizontal 

accuracy approximately 10–30 m). We obtained 

some plot or station locations by using more 

accurate Trimble Pathfinder® GPS units (Trimble 

Navigation Limited, Sunnyvale, CA; horizontal 

accuracy about 1 m). Although we map the 

locations of study plots, stations, or transects 

on Digital Orthophoto Quarter Quads (DOQQ; 

produced by the USGS), the locations of study 

areas will remain with the park and NPS Sonoran 

Desert Network I&M office in Tucson. We also 

produced distribution maps for all vertebrate 

species from this and other recent survey efforts 

(including wildlife observation cards at the 

park). Those maps will be archived in the same 

locations as the GPS coordinates. 

Species Conservation Designations

We indicate species conservation designations by 

the following agencies: U.S. Fish and Wildlife 

Service (responsible for administering the 

Endangered Species Act), USDA Forest Service, 

Arizona Game and Fish Department, and Partners 

in Flight (a partnership of dozens of federal, 

state and local governments, non-governmental 

organizations, and private industry). 

Databases and Data Archiving

We entered field data into taxon-specific 

databases (Microsoft Access version 97) and 

checked all data for transcription errors. From 

these databases, we reproduced copies of the 

original field datasheets using the “Report” 

function in Access. The output looks similar 

to the original datasheets but data are easier to 

read. The databases, printouts of field data, and 

other data such as digital photographs have been 

distributed to park staff and will be distributed to 

Special Collections at the University of Arizona. 

Original copies of all datasheets currently 

reside at the I&M office in Tucson and may be 

permanently archived at another location. Along 

with the archived data, we will include copies 

of the original datasheets and a guide to filling 

2
them out. This information, in conjunction 

with the text of this report, should enable future 

researchers to repeat our work. 

Verification and Assessment of Results

Photograph Vouchers

Whenever possible, we documented vertebrate 

species with analog color photographs. Many 

of these photographs show coloration or other 

characteristics of visual appearance in detail, 

and they may serve as educational tools for the 

park staff and visitors. We obtained a closeup photograph of each animal “in hand” and, 

if possible, another photograph of the animal 

in natural surroundings. Photographs will be 

archived with other data as described above.

Specimen Vouchers

Specimen vouchers are an indisputable form of 

evidence of a species occurrence. For plants, we 

searched the University of Arizona Herbarium 

for existing specimens from the district (see 

Appendix A for results), and we collected 

herbarium specimens whenever flowers or fruit 

were present on plants in the field. All specimens 

that we collected were accessioned into the 

University of Arizona Herbarium. To prioritize 

vertebrate species for voucher collection, we 

first searched the park’s specimen collection and 

that of other universities and collections (Table 

1.1; see Appendix F for results). When we did 

collect specimens, most were found dead. When 

necessary, we euthanized animals according to 

standardized and approved procedures, prepared 

the specimens using accepted methods, and 

deposited them in the appropriate collection at the 

University of Arizona. 

Assessing Inventory Completeness

We assessed inventory completeness by (1) 

examining the rate at which new species were 

recorded in successive surveys (i.e., species 

accumulation curves; Hayek and Buzas 1997) 

and (2) comparing the list of species we recorded 

with a list of species likely to be present based 

on previous research and/or expert opinion. 

We created species accumulation curves for 

all taxonomic groups except plants. For all 

accumulation curves (unless indicated otherwise), 

we randomized the order of the sampling periods 

to break up clusters of new detections that 

resulted from temporal conditions (e.g., monsoon 

initiation) independent of cumulative effort. We 

used the computer program Species Richness 

and Diversity III (Pisces Conservation Ltd., IRC 

House, Pennington, Lymington, UK) to calculate 

species accumulation curves where the order 

of samples was shuffled the maximum number 

of times and the average was plotted, thereby 

smoothing the curve.

3

Table 1.1. Museums that were queried in 1998 for vertebrate voucher specimens with “Arizona” and 

“Saguaro National Park” and “National Monument” in the collection location. 

Brigham Young University Oklahoma Museum of Natural History, Norman

Chicago Academy of Sciences Peabody Museum, Yale University

Cincinnati Museum of Natural History & Science Saguaro National Park (collection now at the Western 

Cornell Vertebrate Collections, Cornell University Archaeological and Conservation Center, Tucson

George Mason University (Fairfax, VA) Strecker Museum, Baylor University, Waco

Illinois Natural History Survey Texas Cooperative Wildlife Collection

Marjorie Barrick Museum, University of Nevada-Las Vegas Tulane Museum of Natural History

Michigan State University Museum (East Lansing) University of Arizona

Milwaukee Public Museum University of Texas, Arlington 

Museum of Natural History, University of Kansas University of Illinois, Champaign-Urbana

Museum of Texas Tech University University of Colorado Museum

Museum of Vertebrate Zoology, University of California, Berkeley United States National Museum

Museum of Life Sciences, Louisiana State University, Shreveport Walnut Canyon National Monument, Arizona

Natural History Museum of Los Angeles County Western Archaeological and Conservation Center, Tucson

North Carolina State Museum of Natural Sciences Wupatki National Monument, Flagstaff
Estimating Abundance

Estimating population size is a common goal 

of biologists who are motivated by the desire 

to reduce (pest species), increase (endangered 

species), maintain (game species) or monitor 

(indicator species) population size. Our surveys 

at the park were generally focused on detecting 

species rather than estimating population size. 

In many cases, however, we present estimates 

of “relative abundance” by species to provide 

information on areas in which species might be 

more or less common. Relative abundance is 

an index to population size; we calculate it as 

the number of individuals of a species recorded, 

scaled by survey effort. If we completed multiple 

surveys in comparable areas, we included a 

measure of precision (usually standard error) with 

the mean of those survey results. 

Indices of abundance are presumed to 

correlate with true population size but ecologists 

do not typically attempt to account for variation 

in detectability among different species or groups 

of species under different circumstances. Metrics 

(rather than indices) of abundance do consider 

variation in detection probability, and these 

include density (number of individuals per unit 

area; e.g., one Arizona black rattlesnake per km2

) 

and absolute abundance (population size; e.g., 30 

Arizona black rattlesnakes at the district). These 

estimates are beyond the scope of our research. 

While it is true that indices to abundance have 

often been criticized (and with good reason, c.f. 

Anderson 2001a), the abundance information that 

we present in this report is used to characterize 

the commonness of different species rather than 

to quantify changes in abundance over long 

periods of time (e.g., monitoring). As such, 

relative abundance estimates are more useful 

than detectability-adjusted estimates of density 

for only a few species or raw count data for all 

species without scaling counts by survey effort. 

Sampling Design

Overview

Sampling design is the process of selecting 

sample units from a population or area of interest. 

Unbiased random samples allow inference to 

the larger population from which those samples 

were drawn and enable one to estimate the true 

value of a parameter. The precision of these 

estimates, based on sample variance, increases 

with the number of samples taken; theoretically, 

random samples can be taken until all possible 

samples have been selected and precision is exact 

– a census has been taken and the true value is 

known. Non-random samples are less likely to be 

representative of the entire population, because 

the sample may (intentionally or not) be biased 

toward a particular characteristic, perhaps one of 

interest or convenience. 

In our surveys we employed both 

random and non-random spatial sampling 

designs for all taxa. For random sites, we colocated all taxonomic studies at the same sites 

(focal points and focal-point transects; see 

below for more information) because some 

characteristics, especially vegetation, could be 

used to explain differences in species richness 

or relative abundance among transects. We also 

used vegetation floristics and structure to group 

transects into community types that allowed more 

accurate data summaries. The location of nonrandom study sites was entirely at the discretion 

of each field crew (i.e., plants, birds, etc.) and we 

made no effort to co-locate them. 

Focal Points and Focal-point Transects: Random 

Sampling 

To account for differences in plant and animal 

communities at different elevation zones (e.g., 

Whittaker and Niering 1965) at the district, 

we used a stratified random design using 

elevation to delineate three strata: <4000, 4,000-

6,000, and >6,000 feet. We chose a stratified 

design over a simple random design because 

stratified sampling better captures the inherent 

environmental variability within each stratum, 

allowing for greater precision of parameter 

estimates and increased sampling efficiency 

(Levy and Lemeshow 1999). This design also 

generates a better spatial dispersion of sampling 

units. Further, we chose to delineate strata 

based on elevation because it can be a good 

predictor of changes in vegetation and animal 

4
communities and is especially useful when no 

reliable vegetation maps exist, as was the case for 

the district. 

Locating Random Study Sites 

We used the following process to assign the 

location of random study areas. First, we created 

100 random (hereafter referred to as “focal”) 

points using the Animal Movement extension 

for ArcView (developed by the USGS Alaska 

Science Center – Biological Science Office), 

using uniform distribution, allowing zero meters 

to the district boundary, and zero meters between 

points. For each focal point, we generated a 

random bearing (the numbers ranged from 0 to 

359). We then used the Bearing and Distance 

extension for ArcView (developed by Ying Ming 

Zhou, March 29, 2000; downloaded from ESRI 

ArcScripts website) to create points based on the 

distance and bearing from the original points. 

This gave us start points and end points for all 

100 focal points. We then used the “from” and 

“to” coordinates to draw the transect line using 

an Avenue script (“Draw line by coordinates,” 

developed by Rodrigo Nobrega, August 13, 1998; 

downloaded from ESRI ArcScripts website). The 

result was randomly placed, 1000-m line transects 

(hereafter referred to as “focal-point transects” 

or “transects”). Focal-point transects were not 

allowed to overlap. If this occurred, an entire new 

selection was conducted until a scenario of no 

overlapping transects was achieved.

Many focal-point transects were not used 

because (1) some part of them lay outside of the 

district boundary, (2) at least 67% of the line did 

not fall within a single stratum, or (3) they were 

in areas where the terrain was too steep to work 

safely (i.e., crossed areas with slopes exceeding 35 

degrees). These “danger” areas were derived from 

30-m Digital Elevation Models using the Spatial 

Analyst extension for ArcView. The final design 

produced four bird-survey stations spaced 250 

m apart; 10, 100 x 100 m amphibian and reptile 

plots; and 20, 50 x 50 m mammal plots along the 

focal-point transect line (Fig. 1.1). We sampled 

5

100m

 1 2 3 4 5 6 7 8 9 10

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

1 2 3 4

A

B

E

D

C

 1 2 3 4

Figure 1.1. Layout of 1-km focal-point transects showing layout of amphibian and 

reptile plots (C), small-mammal trapping grids (D), and bird survey stations (E). 
6

vegetation by point intercept along six, 50-m 

transects (see Chapter 3 for more information).

To map the location of plots, we designed 

a footprint of the sampling grids using an 

Avenue Script (“View.CreateTransectLines,” by 

Neal Banerjee, October 5, 2000; downloaded 

from ESRI ArcScripts website) to create grid 

lines every 100 m that were perpendicular (90 

degrees) to a “dummy” transect (Fig. 1.1A). 

These grid lines were converted from graphics to 

shapes using the XTools extension for ArcView 

(developed by the Oregon Department of 

Forestry). We then generated points where each 

grid line intersected the transect using the Themes 

Intersections to Points extension for ArcView 

(developed by Arun Saraf, November 11, 1999; 

downloaded from ESRI ArcScripts website) (Fig. 

1.1B). 

We created 100 x 100 m squares 

centered on each intersection point to generate 

the amphibian and reptile plots using the 

Square Buffer Wizard extension for ArcView 

(developed by Robert J. Scheitlin, May 12, 2000; 

downloaded from ESRI ArcScripts website). 

These squares were numbered 1 to 10 in the 

direction of the transect bearing (Fig. 1.1C). The 

same process was repeated to create the mammal 

plots (Fig. 1.1D). Four bird survey stations were 

created by selecting the center of mammal plots 

3, 8, 13, and 18 and buffering each of these 

points with a radius of 125 m (Fig. 1.1E). 

These circles were numbered 1 to 4 in the 

direction of the transect bearing.

Non-random Selection of Study Sites

Many areas of the district contain unique areas 

requiring special surveys for all taxa. Riparian 

areas, cliffs, rocky outcrops, and ephemeral 

pools were likely to be missed if we located 

our study sites only in random areas. Yet these 

areas are diversity “hotspots” and are therefore 

crucial to visit in order to complete the species 

inventories. We selected these study areas 

based on our knowledge of the district. The 

area deemed to be of importance differed 

by taxonomic group, but we chose to do 

surveys for all taxa in low-elevation riparian 

areas (e.g., Rincon Creek). For plants, we 

concentrated on Rincon Creek and drainages 

on the east slope of the Rincon Mountains. For 

reptiles and amphibians we searched dozens of 

canyons at low and medium elevations, and for 

mammals we concentrated on middle elevation 

semi-desert grasslands (for more complete 

descriptions of survey areas, see each taxonspecific chapters.


Park Area and History

Saguaro National Park is located in eastern Pima 

County adjacent to Tucson, Arizona (Fig. 2.1). 

Originally designated as a national monument, 

the park was created in 1933 to preserve the 

“exceptional growth” of the saguaro cactus (NPS 

1992b). In 1961, the park was expanded to 

include over 9,000 ha of the Tucson Mountains 

(know as the Tucson Mountain District). The 

Rincon Mountain District (referred to as “the 

district”) is the subject of this report. It is 27,233 

ha in size and is bounded by USDA Forest Service 

land to the east; Forest Service and private land to 

the north; Forest Service, private and state land to 

the south; and private land to the west (Fig. 2.2). 

Although created to preserve natural resources, the 

park is also home to native American campsites 

and petroglyphs and contains remnants of early 

ranching and mining (NPS 1992b). Annual 

visitation to both districts of the park averages 

approximately 700,000 (NPS 2005). 

Natural Resources Overview

Physiography, Geology, and Soils

Saguaro National Park is located within the Basin 

and Range Physiographic Province. The district 

encompasses most of the Rincon Mountains, one 

of the region’s prominent “sky island” mountain 

ranges. Topography at the district varies from 

low-elevation desert flats to steep rocky canyons 

and high-elevation meadows. Elevation ranges 

from 814 m (2,670 feet) in the northwestern 

corner of the district to 2,641 m (8,665 feet) at 

Mica Mountain. The Rincon Mountains are 

primarily metamorphic in origin, with rocks of the 

Santa Catalina Group, a mixture of Pinal Schist, 

Continental Granodiorite, and Wrong Mountain 

Quartz Monzonite (McColly 1961, Drewes 

1977). All components are of Precambrian 

rock parentage, subsequently deformed and 

recrystalized. Sedimentary rocks in the vicinity 

are largely Permian limestones of Earp and 

Horquilla formations (Drewes 1977). 

Hydrology

The Rincon Mountain District has several sources 

of perennial water: Chimenea, Madrona, Rincon, 

and Wild Horse Creeks; and Deer Head, Spud 

Rock, Italian, and Manning Camp Springs. The 

most prominent hydrologic feature is Rincon 

Creek, which drains approximately one-half of 

the district. 

Climate

Saguaro National Park experiences an annual 

bimodal pattern of precipitation which is 

characterized by heavy summer (monsoon) 

storms brought about by moisture coming from 

the Gulf of Mexico, and less intense frontal 

systems coming from the Pacific Ocean in the 

winter. On average, approximately one-half of 

the annual precipitation falls from July through 

September (Tables 2.1, 2.2; WRCC 2005, PCFCD 

2005). The area’s hot season occurs from April 

through October; daily maximum temperatures 

exceed 40o

C at lower elevations and 30oC at 

high elevations. Winter temperatures dip below 

freezing and snow is common at high elevations. 

From 2001 to 2003, during the time of 

most of our inventory effort, average annual 

precipitation totals for the high elevation areas 

were slightly below the long-term mean of 69.1 

cm (60.6 cm in 2001, 38.6 cm from May to Dec 

2002 [no data for Jan–Apr 2002] and 60.0 cm in 

2003; Fig. 2.3; PCFCD 2005). Average annual 

precipitation totals for low elevations ranged 

from slightly to substantially below the longterm mean of 28.6 cm (21.7 cm in 2001, 19.0 cm 

in 2002 and 26.5 cm in 2003; Fig. 2.3; WRCC 

2005). The percent of the total precipitation 

during the monsoon season (July through 

September) was higher in the low elevation 

(50%) than in the high elevation (40%) areas 

(Tables 2.1, 2.2). 

Chapter 2: Park Overview

Brian F. Powell, Cecilia A. Schmidt, and William L. Halvorson
Average annual temperatures for low 

elevations from 2001 to 2003 were above the 

long-term mean of 21.3o

C (21.5oC in 2001, 

21.6o

C in 2002 and 22.0oC in 2003; Fig 2.3; 

WRCC 2005). Average annual temperatures for 

high elevations ranged from slightly below to 

slightly above the long-term mean of 8.5o

C (6.7oC 

in 2001, 7.3oC in 2002 and 9.5oC in 2003; Fig 

2.3; PCFCD 2005), though these records have 

only been kept for 10 years. 

Vegetation and Biotic Communities

The Rincon Mountain District encompasses 

most of the Rincon Mountains, one of the “sky 

island” mountain ranges of southeast Arizona 

and northern Mexico. Sky islands, so called 

because the “sky” mountains are isolated by 

“seas” of desert and semi-desert grasslands, are 

areas of remarkable biological diversity as a 

result of elevational gradients and subsequent 

Figure 2.1. Location of the two districts of Saguaro National Park in southern Arizona.
Figure 2.2. Aerial photograph showing major features of Saguaro National Park, Rincon Mountain District. 

Table 2.1. Average monthly climate data for Manning Camp (high elevation), Saguaro National Park, 

Rincon Mountain District, 1994–2004. Data from PCFCD (2005).

Month

Characteristic Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Annual

Maximum temperature (o

C) 15.6 15.0 17.9 19.8 27.0 27.4 29.2 27.1 25.6 23.0 17.9 15.2 21.7

Minimum temperature (o

C) -10.6 -9.6 -9.8 -5.6 -4.1 1.6 7.3 7.0 3.8 -3.7 -8.1 -10.6 -3.5

Precipitation (cm) 6.5 6.6 8.3 3.4 0.8 0.9 12.2 11.2 4.6 3.7 3.9 7.0 5.8

Table 2.2. Average monthly climate data for the University of Arizona (low elevation; the closest climate 

monitoring station to Saguaro National Park, Rincon Mountain District) 1894–2004. Data from WRCC 

(2005).

Month

Characteristic Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Annual

Maximum temperature (o

C) 18.6 20.5 23.5 27.8 32.6 37.7 37.8 36.7 35.1 29.9 23.5 19.0 28.6

Minimum temperature (o

C) 3.1 4.5 6.7 9.9 14.2 19.3 23.3 22.4 19.3 12.7 6.6 3.4 12.1

Precipitation (cm) 2.3 2.2 1.9 1.0 0.4 0.7 5.2 5.4 3.0 1.9 2.0 2.5 2.3
10

differences in precipitation and temperature. 

These mountain ranges extend from subtropical 

to temperate latitudes, hosting species whose 

core distributions are from the Sierra Madre of 

Mexico and the Rocky Mountains of the United 

States and Canada (Warshall 1994). In southern 

Arizona, the sky island mountain ranges have 

similar and predictable vegetation communities 

across elevational gradients, from low-elevation 

Sonoran desertscrub to high-elevation conifer 

forests. Below we review the major vegetation 

and biotic communities found in the Rincon 

Mountains. 

Sonoran Desertscrub

Sonoran Desertscrub (Sonoran Desert Scrub; 

Fig. 2.4) is found in the lowest elevation and 

driest areas of the district on its west and 

southern boundaries. The dominant shrubs 

are velvet mesquite (Prosopis spp.), acacias 

(Acacia spp.), palo verdes (Cercidium spp.), and 

creosote bush (Larrea tridentata). Succulents 

are ubiquitous and include: agave (Agave spp.), 

yucca (Yucca spp.), barrel cactus (Ferrocactus

and Echinocactus spp.), pincushion cactus 

(Mammalaria spp.), and prickly pear and cholla 

(Opuntia spp.). Warm- and cool-season annuals, 

both native (e.g., woolly plantain, [Plantago 

patagonia]) and introduced (e.g., red brome, 

[Bromus rubens]) are common following rainfall.

Southwestern Deciduous Riparian Forest

These forests (Canyon Woodland; Fig 2.4) are 

found along low-elevation washes and creeks 

and are among the most biologically unique 

communities in the Sonoran Desert ecoregion. 

At the district they are found along Rincon Creek 

and to a lesser extent along its tributaries. The 

dominant tree species are Fremont cottonwood 

(Populus fremonti), Arizona sycamore (Platanus 

wrightii), velvet ash (Fraxinus velutina), willow 

(Salix spp.), and netleaf hackberry (Celtis 

reticulata). In the Rincon Mountain District 

Sonoran Desertscrub bounds these zones. 

Figure 2.3. Comparison of monthly weather data during the time of the majority of the inventory 

effort (2001–2003) compared to the mean (1994–2004 for Manning Camp, 1894–2004 for University of 

Arizona; thick solid line in all figures), Saguaro National Park. Data for Manning Camp from PCFCD 

(2005) and data for University of Arizona from WRCC (2005).

Month

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

-10

-5

0

5

10

2001

2002

2003

Precipitation

Month

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

-4

-2

0

2

4 2001

2002

2003

Temperature

Manning Camp 

Month

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Centimenters from mean -4

-2

0

2

4 2001

2002

2003

Precipitation

Month

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

-4

-2

0

2

4

6

2001

2002

2003

Temperature

University of Arizona 

Centimeters from mean Degrees Celsius difference from mean

 Centimeters from mean Degrees Celsius difference from mean
11

Figure 2.4. Diagram of the major vegetation communities of the Santa Catalina Mountains, adjacent to the 

Rincon Mountains (from Whittaker and Niering 1965). The Rincon Mountains have similar communities with the 

exception of the subalpine forest community. Reprinted with permission from the Ecological Society of America. 
12

Semi-desert Grassland

Semi-desert grasslands (Desert Grassland; Fig 

2.4) occur in some middle elevation areas of the 

district, primarily along the northern boundary of 

and in a few areas of Tanque Verde Ridge. The 

community is composed of perennial short- and 

mid-grass species, with most areas invaded by 

velvet mesquite (Prosopis velutina Woot.). 

Oak Savannah

The oak savannah community (Open Oak 

Woodland; Fig 2.4) is found at higher elevations 

than the semi-desert grassland community and 

lower elevations than the pine-oak woodland, and 

it contains elements of both communities. It is 

ecologically similar to the chaparral communities 

of central Arizona. In this community there are 

dense stands of manzanita (Arctostaphylos spp.) 

and oak (Quercus spp.), with a variety of annual 

and perennial grasses. 

Pine-oak Forest and Woodland

Pine-oak forest and woodland (sometimes 

referred to as Madrean evergreen woodland; 

Fig. 2.4) is ubiquitous at mid-elevations 

throughout the Apache Highlands (Bailey 

1998, McPherson 1993). Madrean evergreen 

woodland is characterized by evergreen oaks 

with thick sclerophyllous leaves, such as emory 

oak (Quercus emoryi Torr.), Arizona white oak 

(Quercus arizonica Sarg.), and Mexican blue oak 

(Quercus oblongifolia Torr.). Mexican pinyon 

pine (Pinus cembroides Zucc.) and alligator 

juniper (Juniperus deppeana Steud.) are the 

common gymnosperms. Understory grasses are 

usually abundant. At the higher elevations and in 

drainages, there is also ponderosa pine.

Coniferous Forest

Dominated by gymnosperms such as pines 

(Pinus spp.), and firs (Abies spp.), coniferous 

forests (Pine and Montane Fir Forests; Fig 2.4) 

represent the cold-hardiest biotic community 

in the district. In these communities in the 

district, ponderosa pine (Pinus ponderosa P. &C. 

Lawson) and Douglas fir (Pseudotsuga menziesii

[Mirbel] Franco) dominate, with some temperate 

deciduous plants intermixing, primarily on 

the north-facing slopes: Gambel oak (Quercus 

gambelii Nutt.), quaking aspen (Populus 

tremuloides Michx.), and maples (Acer spp.) and 

boxelder (Acer spp.). Conifer forests are fireadapted ecosystems, with natural low-intensity 

fires occurring every 6 to 15 years (Baisan and 

Swetnam 1990, Dimmitt 2000). 

Natural Resource Management Issues

Adjacent Land Development

Increasing housing development along the 

western and southern boundaries has become 

the most pressing natural resource issue for the 

district. Sandwiched between both districts 

of the park, the greater Tucson metropolitan 

area is one of the fastest growing in the United 

States. The area currently has an estimated 

population of 800,000, a 44% increase over the 

last two decades (PAG 2005). The increase 

in human residents brings with it a variety of 

natural resource-related problems including 

harassment and predation of native species by 

feral animals, increased traffic leading to altered 

animal movement patterns and mortality, the 

spread of non-native species, illegal collections 

of animals, vandalism, increased water demands, 

air pollution from vehicle emissions, and visual 

intrusions to the natural landscape (Briggs et al. 

1996). Throughout this document we highlight 

some of these impacts as they pertain to each 

taxonomic group. 

Of immediate concern for park 

managers is the depletion of groundwater and 

its effects on the ecologically valuable Rincon 

Creek, in particular (Baird et al. 2000). There 

are numerous single-family and large-scale 

housing units being constructed (or planned) 

directly adjacent to the district, including the 

proposed Rocking K Ranch development, 

which anticipates 9,000 residents and has been 

granted a permit by the Arizona Department of 

Water Resources to withdraw 4,400 acre feet per 

year from the underlying aquifer (Mott 1997). 

Rincon Creek has the most well-developed 

stretch of southwestern deciduous riparian forest 

in the district, which will likely be impacted 

by drawdown of the aquifer. Groundwater 

drawdown at Tanque Verde Wash has already 

affected the riparian community there (Mott 

1997). 
13

Non-native Species and Changes to Vegetation

The spread of non-native species within the 

district is an important natural resource issue. 

In particular, buffelgrass (Pennisetum ciliare), 

Lehmann lovegrass (Eragrostis lehmanniana), 

red brome (Bromus rubens) and other nonnative grasses, have increased in the last ten 

years (Funicelli et al. 2001). The spread of 

some non-native plants used for landscaping, 

such as crimson fountaingrass (Pennisetum 

setaceum) from development bordering the 

district is also a concern. The invasion of nonnative grasses has led to structural changes in 

vegetation, from areas that supported mostly 

sparse bunchgrasses to areas of uniform grass. 

This change in species composition and structure 

can alter the fire regime of the area by supporting 

higher fire frequencies, thereby leading to other 

changes in vegetation composition and structure 

(Anable et al. 1992). Nowhere are these effects 

more evident than in the Sonoran Desertscrub 

vegetation community, which rarely burned 

historically (Steenbergh and Lowe 1977). Many 

native plant species, especially succulents, are not 

adapted to short duration but high-intensity fires 

and therefore die (Schwalbe et al. 1999, Dimmitt 

2000). Fires such as the Mother’s day fire, 

which was fueled largely by non-native grasses, 

have caused a high mortality of saguaro cactus 

(Carnegiae gigantea Britt. & Rose), which is of 

great concern to park managers (Schwalbe et al. 

1999; see Chapter 3 for additional information). 

Wildland Fire

Since the park began keeping records in 1937, 

there have been 572 fires in the district, and since 

1984, park personnel have burned approximately 

1,450 ha through their active fire-management 

program. Fires play a crucial role in the middle 

and high-elevation semi-desert grasslands and 

forests by depleting dense understory vegetation 

and downed-woody debris. Even in these 

fire-adapted ecosystems, however, fire can 

be devastating, particularly after decades of 

suppression and subsequent buildup of fuel loads. 

A number of large fires in the last few decades, 

most notably the Chiva and Box Canyon fires, 

caused massive runoffs of sediment and ash. 

The Chiva fire apparently eliminated lowland 

leopard frog habitat and may have destroyed the 

district’s only population of (federally listed) 

Gila topminnow (Poeciliopsis occidentalis 

occidentalis) at Little Wildhorse Tank, though 

their status as a natural or introduced population 

was uncertain (Don Swann, pers. comm.). The 

Box Canyon fire of 1999 led to the sedimentation 

of perennial pools, where lowland leopard frogs 

once bred (Don Swann, unpubl. data). Despite 

some problems, the NPS is committed to 

returning natural fire cycles to the high elevation 

areas of the district. 
14
15

Previous and Ongoing Research

Floras and Plant Collections

We located specimens representing 883 species at 

the University of Arizona Herbarium (Appendix 

A). Many of these specimens were collected 

or reported in Bowers and McLaughlin (1987). 

Their treatise is the most comprehensive 

annotated flora for the Rincon Mountains, though 

species have been added to the list since its 

publication. Bowers and McLaughlin (1987) 

also provide an excellent overview of previous 

research and collecting from the range (as does 

Bowers [1984]), the plant communities present, 

species richness gradients, and a list of species 

extirpated from the range. The Bowers and 

McLaughlin list was compiled from work by 

Bowers (1984) above 4,500 feet elevation and by 

Carole Jenkins who collected from 1978 to 1982 

below 4,500 feet elevation. Jenkins never wrote 

up the results of her work. The list was updated 

in 1996 to include the addition of 34 species 

and the subtraction of four (due to incorrect 

identifications; Fishbein and Bowers 1996). 

There have been floras for four designated natural 

areas of the district: Wildhorse Canyon (Rondeau 

and Van Devender 1992), Chimenea Canyon 

(Fishbein et al. 1994a), Box Canyon (Fishbein 

et al. 1994b), and Madrona Canyon (Fishbein 

1995). Halvorson and Gebow (2000) compiled 

these works into a single volume. Halvorson and 

Guertin (2003) mapped locations of 27 species of 

non-native plants.

Monitoring, Research, and Single-species Studies

Park personnel established long-term monitoring 

plots in low-elevation areas of both units 

(Saguaro NP 2005). They used the pointintercept method at 25 plots in the Rincon 

Mountain District and 20 plots in the Tucson 

Mountain District and surveyed these transects 

from 1998 to 2004 (Mark Holden, pers. 

comm.). Funicelli et al. (2001) resurveyed 25, 

10 x 10 m vegetation plots (established 10 years 

prior to their surveys) and mapped each plant 

species. These plots were also used by Turner 

and Funicelli (2000) to resurvey the condition 

and population structure of the saguaro cactus. 

Swann et al. (2003a) used the same protocol as 

that used by Funicelli et al. (2001) to survey for 

plants on the east slope of the Rincon Mountains. 

Anderson (2001b) surveyed vegetation transects 

at random sites in the Rocking K and adjacent 

expansion areas. 

The saguaro cactus, the park’s namesake 

species, has been one of the most investigated 

non-agricultural plants in the world. McAuliffe 

(1993) provided an overview of saguaro research 

at the park as well as its political and scientific 

context. Schwalbe et al. (1999) surveyed 

vegetation in and adjacent to the area burned 

by the Mother’s Day fire of 1994. Baisan and 

Swetnam (1990) constructed a fire history (1657–

1893) of the conifer forest in the vicinity of Mica 

Mountain. Though there is a GIS layer of 15 

dominant vegetation communities in the district, 

there is not a current, detailed vegetation map. 

In fact, the most current vegetation map was by 

Roseberry and Dole (1939).

Current projects include a fire-effects 

monitoring program in the high elevation areas 

of the district (Saguaro NP, unpubl. data) and a 

program to map and remove non-native species 

(e.g., buffelgrass, fountaingrass, Saharan mustard, 

and Malta starthistle) from low-elevation areas of 

both districts of the park. 

Methods

We used three field methods to survey for 

vascular plants. General botanizing surveys 

involved opportunistically collecting what we 

thought might be new additions to the district’s 

flora or plants that we could not identify in the 

field. We also used modified-Whittaker plots 

and point-intercept transects to make quantitative 

comparisons among areas and provide data for 

long-term monitoring. 

Chapter 3: Plant Inventory

Brian F. Powell
16

General Botanizing

Methods

We collected species opportunistically and when 

we thought we had found a species not on the 

district list (derived principally from Bowers 

and McLaughlin [1987]). We also searched 

specifically for species that were listed as 

possibly extirpated from the district (in Bowers 

and McLaughlin 1987). Whenever possible we 

collected at least one representative specimen 

with reproductive structures for each plant 

species that we encountered. We also maintained 

a list of species observed but not collected. 

When we collected a specimen, we assigned it a 

collection number and recorded the flower color, 

associated dominant vegetation, date, collector 

name(s), and UTM coordinates. We pressed and 

processed the specimens on site. Specimens 

remained pressed for two to three weeks and 

were later frozen for 48 hours or more to prevent 

infestation by insects and pathogens. Mounted 

specimens were accessioned into the University 

of Arizona Herbarium.

Effort

We collected specimens during 38 days of 

fieldwork between 10 April and 24 September 

2001 and 4 and 5 May 2002. We collected 

specimens from 41 locations throughout the 

district (Fig. 3.1) and many of the collections 

were made in the course of traveling to and from 

focal points. 

Analysis

We present a variety of summary statistics 

including total number of species found and 

number and percent of native and non-native 

species. 

Modified-Whittaker Plots

We used modified-Whittaker plots to characterize 

the plant community at a single area associated 

with focal points. Each plot was 20 x 50 m 

(1000 m2) and contained 13 subplots of three 

different sizes (see Stohlgren et al. 1995a): 0.5 

x 2 m (10 subplots), 2 x 5 m (2 subplots), and 5 

x 20 m (1 subplot) (Fig. 3.2; Shmida 1984). We 

estimated the coverage (m2) of each plant species 

for the entire 1000 m2 plot. For all subplots we 

simply noted the presence of each species. For a 

more detailed explanation of the data collection 

method, see Shmida (1984). We deviated from 

the methods outlined in Shmida (1984) by not 

surveying against the contours in steep areas, 

because of safety reasons. 

Effort

We used modified-Whittaker plots at 13 of the 17 

focal points (Fig. 3.3). We excluded four plots 

(numbers 120, 121, 125, and 155) because of 

logistical constraints. We used a single observer 

(Patty West) to estimate percent cover in the 20 

x 50 m plot, but other observers occasionally 

assisted with noting presence of plants in 

subplots. 

Analysis

We note patterns of species richness among plots 

and community types. In this report we do not 

present a complete summary of the data, but 

instead will archive these summaries (see Chapter 

1 for archive locations). 

Point-intercept Transects

Methods

We used the point-intercept method (Bonham 

1989) to sample vegetation along 50-m transects 

located along each focal-point transect (Fig. 

3.4). Point-intercept transects began at 25, 125, 

425, 525, 825 and 925 m from the beginning of 

the transect (i.e., focal point). For example, the 

first transect started at 25 m from the focal point 

and went to the 75-m mark. We placed a 50-m 

transect tape along the length of each transect 

section. In each of four height categories (<0.5 

m, 0.5–2 m, 2-4 m, and >4 m) we recorded the 

species of the first plant intercepted by a vertical 

line every 1 m along the transect line (n = 300 

points for most transects). We created the vertical 

line using a graduated pole and extrapolated 

contacts in a fourth height category (>4 m), 

which was rarely used in the desert areas. We 

classified groundcover as rock, bare ground, 

annual forb, grass or woody debris.
17

Figure 3.1. Locations of general botanizing collection sites, Saguaro National Park, Rincon Mountain 

District, 2001 and 2002.

0.5 x 2m 

2 x 5m 

5 x 20m 

20m

50 m 

Figure 3.2. Layout of a modified-Whittaker plot, Saguaro National Park, Rincon Mountain District, 2001.
18

Effort

We surveyed along each of the 17 random 

transects (Fig. 3.3) in the spring of 2001. We 

typically worked in groups of two or three field 

personnel, but sometimes had as many as five 

field personnel. We surveyed a total of 300 

points along most transects. Additional points 

were surveyed on a subset of transects when time 

permitted; transects with difficult terrain resulted 

in fewer than 300 points being surveyed. 

Analysis

We calculated percent cover and percent 

composition for each species in each height 

category. Percent cover is the number of times a 

Figure 3.3. Locations of modified-Whittaker plots and point-intercept transects (line transect), 

Saguaro National Park, Rincon Mountain District, 2001.

Distance from focal point 

125 m 25 m 

Focal point (beginning 

of transect) Point-intercept transects 

925 m 825 m 525 m 425 m 

50 m

0 m 

Figure 3.4. Typical layout of point-intercept transects, Saguaro National Park, Rincon Mountain District, 2001. 
19

species was encountered along the entire length 

of the transect divided by effort (in most cases a 

maximum of 300 intercepts per height category) 

and multiplied by 100. We calculated percent 

composition of each species in each height 

category as the number of times a species was 

encountered divided by the number of times all 

other species were encountered. If there was at 

least a single species encountered along a transect 

(in each height category), the total percent 

composition equaled 100 percent. 

Community Types

We sought to identify plant communities within 

the district and to compare characteristics among 

them. We did not use the original stratification 

of random transects for this analysis because we 

were more interested in classifying communities 

than drawing inference to a larger area. To group 

transects, we used Ward’s hierarchical cluster 

analysis using data from point-intercept transects. 

Cluster analysis is a multivariate technique that 

groups like entities (in our case transects) that 

share similar values. We used the total number 

of point intercepts by the most common plant 

species in all four height categories for this 

analysis. A detailed summary of point-intercept 

data will be available along with other archived 

materials (see Chapter 1).

Results

We collected 741 specimens representing 523 

species from the Rincon Mountain District 

of Saguaro National Park (Appendix A). We 

found 39 species that had not previously been 

documented in the district, almost one-half of 

them (n = 19) during the course of surveying 

at point-intercept and/or modified-Whittaker 

plots. The list of new species that we found 

included five non-native species, most notably 

African sumac (Rhus lancea). Native species of 

note that we added to the flora included cleftleaf 

wildheliotrope (Phacelia crenulata), Arizona 

dewberry (Rubus arizonensis), and American 

black nightshade (Solanum americanum). 

Based on a thorough review of past 

studies, floras, and collections located at the 

University of Arizona, there have been a total of 

1,170 specific and intraspecific taxa documented 

at the district, of which 78 (6.7%) are non native. 

Excluding eight species in the UA collection that 

Bowers and McLaughlin (1987) cite as likely 

extirpated from the district, there have been 

1,120 species (1,162 including intraspecific taxa) 

documented since the early 1980s (Appendix 

A). Of these species, six were thought to be 

extirpated by Bowers and McLaughlin (1987) but 

were found by other studies: purple scalystem 

(Elytraria imbicata), Lemmon’s hawkweed 

(Hieracium lemmonii), alderleaf mountain 

mahogany (Cercocarpus montanus), Baltic rush 

(Juncus balticus), poverty rush (J. tenius), and 

common barley (Hordeum vulgare) (Appendix 

A). 

Community Types

Based on our interpretation of the cluster analysis 

using data from point-intercept transects, there 

are four communities (i.e., clusters) represented:

•	 Sonoran Desertscrub. Five lowelevation transects (112, 115, 130, 138, 

and 139) and one middle elevation 

transect (121). Mixed cacti and 

paloverde (Parkinsonia spp.), with some 

velvet mesquite (Prosopis velutina), 

especially in the dry washes.

•	 Oak Savannah. Four middle-elevation 

transects (101, 106, 189, and 111). Open 

areas dominated by perennial grasses 

with scattered trees, mostly oaks.

•	 Pine-oak Woodland. Two middle 

(125 and 120) and three high (107, 

155, and 128) elevation transects. 

Most transects had dense stands of 

manzanita (Arctostaphylos spp.) and 

oaks, interspersed with some pine trees, 

mostly pinon and ponderosa pine (Pinus 

ponderosa).

•	 Conifer Forest. Two high elevation 

random transects (113 and 191). Tall 

forests of ponderosa pine, Douglas fir 

(Pseudotsuga menziesii), and some 

oaks, especially Gambel oak (Quercus 

gambelii). 
20

Focal-points: General Patterns

We found 367 species associated with the 17 

focal points. Approximately 47% of these species 

(n = 173) we found associated with only a single 

focal point, whereas six species (spidergrass 

[Aristida ternipes], side-oats grama [Bouteloua 

curtipendula], plains lovegrass [Eragrostis 

intermedia], bullgrass [Muhlenbergia emersleyi], 

sacahuista [Nolina microcarpa], and skunkbush 

sumac [Rhus trilobata]) were associated with 10 

or more focal points. The skunkbush sumac was 

the most widespread species; we found it at 71% 

(n = 12) of focal points. 

We found 354 species at the 13 focal 

points where we used both focal-point and 

modified-Whittaker plot survey methods. At 

these focal points, species richness varied 

among the five community types (F3,9 = 21.8, P 

< 0.001, one-way ANOVA). The Conifer Forest 

community had the fewest number of species (26 

+ 8.3 [SE]) and the Sonoran Desertscrub had the 

most species (103 + 5.3). The other communities 

were intermediate: Oak Savannah (81 + 5.9) and 

Pine-oak woodland (64 + 8.3). 

Modified-Whittaker Plots

We recorded 307 species on 13 modifiedWhittaker plots. The mean number of species 

per plot was 60 + 7.8 (SE) with the range from 

97 species in one of the Sonoran Desertscrub 

plots to 20 species in one of the Conifer Forest 

plots. Based on the previous classification 

of plots grouped into community types, we 

compared species richness among communities 

and found differences (F3,9 = 15.9, P < 0.001, 

one-way ANOVA), though sample sizes for each 

community were quite low. The Conifer Forest 

community had the fewest number of species (21 

+ 7.9) and the Sonoran Desertscrub had the most 

species (83 + 5.0). The other communities were 

intermediate: Oak Savannah (55 + 5.6) and Pineoak woodland (50 + 7.9). 

Point-intercept Transects

We found 189 species on 17 point-intercept 

transects. The mean number of species at each 

transect was 28.3 (+ 2.4 [SE]) and ranged from 8 

to 43 observed. Species richness varied among 

the five community types (F3,9 = 25.5, P < 0.001, 

one-way ANOVA) with Oak Savannah having the 

highest species richness (40 + 2.2) and Conifer 

Forest the lowest species richness (10 + 3.2) (Fig. 

3.5). The Sonoran Desertscrub (33 + 1.8) and 

Pine-oak Woodland (24 + 2.0) were intermediate. 

As expected, vertical structure (as 

expressed by the total number of intercepts in 

each of the four height categories), was also 

different among community types (Fig. 3.5). At 

the Sonoran Desertscrub transects, there was 

considerable vegetation close to the ground 

and progressively less vegetation as we moved 

through the other layers of vegetation. Only 

in the most well-developed washes (or with 

the inclusion of saguaro cactus) is there any 

vegetation in the overstory (>4 m). Conversely, 

in the high elevation transects of the Conifer 

Forest community, there is little vegetation in the 

understory vegetation classes and considerably 

more vegetation in the overstory, which consists 

of tall conifer trees. Vertical structure in the 

middle elevation communities shows changes 

in structure toward these two extremes. Ground 

cover type also reflects this gradient, from 

progressively less plant material as one moves up 

the elevational gradient to bare ground that shows 

the opposite pattern (Fig. 3.6). 

Comparison of Modified-Whittaker and Point-intercept 

Transects

Comparing modified-Whittaker plots and pointintercept transects at focal points where we used 

both methods (n = 13), we found a mean of 60% 

(+ 2.7 [SE]) more species on modified-Whittaker 

plots. Differences in species richness between 

the two methods were most pronounced for the 

Sonoran Desertscrub community (67.6 + 2.7) 

and least pronounced for the Oak Savannah 

community (49 + 3.1). The other communities 

were more similar to the Desertscrub community: 

Conifer Forest (61.6 + 4.3) and Pine-oak 

Woodland (62.5 + 4.3). Within each focal point, 

the percent of species that were common to both 

methods was low (23 + 1.7) and did not vary 

significantly among community types (F3,9 = 1.3, 
21

Desertscrub

Oak Savannah

Pine-oak Woodland

Conifer Forest

Number of species

0

10

20

30

40

50

0-0.5 m

0.5-2.0 m

2.0-4.0 m

>4.0 m

Percent

0

20

40

60

80

100

0-0.5 m

0.5-2.0 m

2.0-4.0 m

>4.0 m

Cover

Species Richness

Figure 3.5. Summary (mean + SD) of data from point-intercept transects by community type and 

height class, Saguaro National Park, Rincon Mountain District, 2001. 

P < 0.32, one-way ANOVA). Finally, the number 

of species that we found along transects that we 

did not find in modified-Whittaker plots was 

lowest at the Conifer Forest (3 + 4.5), highest at 

the Oak Savannah (20 + 3.1) and intermediate at 

the Sonoran Desertscrub (13 + 2.8) and Pine-oak 

Woodland (8 + 4.4) plots. 

Inventory Completeness

The district’s flora is perhaps the most complete 

of any large natural area in the Sky Island region 

of southeastern Arizona. In our many days of 

collecting, we found 39 previously undocumented 

species, which represents a 3.3% increase in the 

flora for the district (Appendix A). Almost onehalf of these species were found during the course 

of conducting surveys at focal points. We also 

found a number of species on the east slope of 

the Rincon Mountains. Collectively these areas, 

particularly those away from hiking trails, are the 

least-surveyed areas of the district and finding 

new species there is not surprising.

Assessing overall inventory completeness 

is problematic given the size of the district and 

difficulty accessing many areas because of rough 
22

terrain. Due to the fact that much of the district 

remains unsurveyed, it is possible that we and 

others have not reached the goal of documenting 

90% of the plant species for the entire district. 

However, if we look at inventory effort in 

different areas, the completion estimates are 

mixed. For example, low-elevation, more easily 

accessed areas almost certainly have a species 

list that is close to completion. We found only 

three new species at or near focal points in the 

low-elevation stratum, and only one new species 

in an area near the Loop Drive, a highly visited 

area. The park’s monitoring efforts have had 

similar results in low-elevation areas; in their 25 

long-term monitoring plots (surveyed for seven 

years) park staff have found only 15 new species 

for the district (Appendix A). The flora for the 

high-elevation areas of the district is similarly 

complete. We found only one species in the area 

around Manning Camp, an area that has had 

extensive plot-level research related to the fireeffects program. That program has produced 

only 30 new species in 15 years of surveys of 71 

plots (Saguaro National Park, unpubl. data). By 

contrast, the mid-elevation areas are the least 

surveyed and our results reflect this; we found 

most of our new species at focal points in the 

middle-elevation stratum (e.g., plots 101 and 189 

had four and three new species for the district, 

Desertscrub

Oak Savannah

Pine-oak Woodland

Conifer Forest

Percent

0

20

40

60

80

100

120

Plant 

Rock or Bare ground 

Figure 3.6. Percent (mean + SD) ground cover from point-intercept transects by community type, 

Saguaro National Park, Rincon Mountain District, 2001.

respectively). These plots were among the most 

difficult to reach areas of the district (Fig. 3.3). 

Based on this evidence, we suggest that the floras 

for low- and high-elevation areas are nearly 

complete and that future surveys should focus on 

middle-elevation areas, especially the east slope 

of the Rincon Mountains and the northeastern 

boundary of the district.

Efficacy of Focal Points

Our plot and point-intercept work was insufficient 

to describe all of the vegetation communities of 

the district. Given the size of the district, the 

random location of 17 study sites was certain 

to miss a number of important features and 

areas. These included communities such as the 

semi-desert grasslands and riparian deciduous 

woodland, and many areas such as the east 

and northeast slopes of the Rincon Mountains 

(Fig. 3.3). However, the plots and transects 

were instrumental in (1) establishing long-term 

monitoring plots, (2) getting researchers to areas 

that had never been visited and therefore led to 

the discovery of new species to the district’s flora, 

and (3) providing information used in assessing 

habitat associations for vertebrates. 
23

Discussion

The Rincon Mountain District’s flora is one of the 

most complete floras of the region and it reflects 

extraordinary species richness. Here we review 

some main determinants of species richness, 

though a more thorough analysis can be found 

in Bowers and McLaughlin (1987) and Bowers 

(1984). The most important factors affecting 

species richness are the range of elevations in the 

district and biogeographic factors. 

The Rincon Mountains have an 

elevational range of about 1,800 m (5,900 

feet). Along the gradient from desert floor to 

the highest elevations of the range, temperature 

and rainfall also change, and plants respond to 

these changes. Aspect is also important, where 

high-elevation, north-facing slopes, in particular, 

harbor species that would not otherwise occur 

in the range, such as Rocky Mountain maple 

(Acer glabrum) and Arizona valerian (Valeriana 

arizonica; See Fig. 2.4). Other features that 

play a role in determining local species richness 

include seeps and springs and limestone rock 

outcrops, the latter of which are responsible for 

the presence of at least 35 species in the Turkey 

Creek area (Bowers and McLaughlin 1987). 

The flora of the district is comprised 

of species from a number of biogeographic 

regions, most notably the Sonoran, Chihuahuan, 

and Madrean in the low-elevation areas and the 

Rocky Mountain and Great Plains biogeographic 

regions in the high-elevation areas of the district. 

Bowers and McLaughlin (1987) observed that 

species richness showed an inverse relationship 

to elevation, which was also evident from our 

plot and transect work (Fig. 3.5). This pattern 

is largely the result of the biogeographic 

influence, where species in low-elevation areas 

have distributions that are primarily southern 

(and represented by Madrean, Sonoran, and 

Chihuahuan biogeographical provinces). 

Accordingly, plant species richness increases 

towards the Equator. By contrast, most species 

in the higher-elevation areas of the district have 

greater affinity with northern biogeographical 

provinces; this is consistent with the observed 

decrease in species richness as one moves north 

from the region. These patterns are mirrored in 

other, nearby mountain ranges (e.g., Whittaker 

and Niering 1965). In addition to biogeographic 

influences, there is also high endemism in the 

southwestern United States. McLaughlin (1986) 

analyzed species composition from 50 floras from 

the region and found that over one-half of the 

species occurred in only one or two of the floras. 

Plant species richness in the Rincon 

Mountains is greater than in other nearby 

mountain ranges with relatively complete 

floras. For example, the Huachuca Mountains, 

to the southeast of the Rincon Mountains, 

contains 929 species (Bowers and McLaughlin 

1996), though the Huachuca Range does not 

contain low-elevation Sonoran Desertscrub. 

Similarly, the lower species richness for the 

Pinaleño Mountains (844 species; Johnson 1988, 

McLaughlin 1993, McLaughlin and McClaran 

2004) is likely explained by the lack of species 

from the Sonoran and Chihuahuan desertscrub 

communities, though it is worth noting that the 

elevation range is similar to that of the Rincon 

Mountains. McLaughlin and McClaran (2004) 

also attribute the low species richness in the 

Pinaleños to “comparatively uniform geology and 

topography.”

Bowers and McLaughlin (1987) cited 

41 species that they believed were extirpated 

from the district because of habitat modification. 

Although we looked for them, we did not find 

any of these species, but our review of other 

studies and localized floras within the district 

revealed that six of these species have been 

found since the publication of the Bowers and 

McLaughlin report, including two species of 

rush (Juncus sp.) and the alderleaf mountain 

mahogany (Cercocarpus montanus). These finds 

are encouraging, but as Bowers and McLaughlin 

(1987) note, many of the species that are likely 

extirpated include a number of moisture-loving, 

high-elevation plants that may be permanently 

lost from the range not only due to habitat 

disturbance, but also to global climate change, 

which has reduced the annual winter snowpack 

that enabled many of these species to survive. 

Habitat disturbance may have led to 

the extirpation of a number of species in the 

high-elevation area of the district, and it may 

also be impacting other areas of the district as 
24

well (Swantek et al. 1999). The most prominent 

habitat disturbance in the district is wildland fire. 

Since the NPS began keeping records in 1937, 

there have been 572 fires in the district and the 

park has an active prescribed fire program. As 

part of the program, park personnel monitor 

vegetation responses at 71 plots located in the 

higher elevation areas of the district (Saguaro 

National Park, unpubl. data). Unfortunately, 

there has been no comprehensive report detailing 

the results of that program, so the effects of 

prescribed fire on the abundance and distribution 

of plants in those areas remains largely unknown. 

Historically, there have been 35 major wildland 

fires in the conifer forest near Mica Mountain 

from 1770–1990 (Baisan and Swetnam 1990). 

Other naturally occurring wildland fires have 

burned through the district, and some have been 

in the lower-elevation Sonoran Desertscrub, 

which has not historically been subject to 

fire (Steenbergh and Lowe 1977, Esque et al 

2003). This relatively new phenomenon has 

resulted from an increase in abundance of 

non-native annual grasses (Schwalbe et al. 

1999). Of particular concern to park managers 

are the impacts of fire on saguaro populations 

(Steenbergh and Lowe 1977). These concerns 

are well founded; in the area of the Mother’s Day 

fire of 1994, Schwalbe et al. (1999) found 22% 

mortality of saguaro within four years of the fire. 

This is considered to be a catastrophic event for 

such a long-lived cactus species. Wildland fire 

has important impacts on other resources of the 

park such as soil, air quality, and animals. We 

discuss the impact of fire on vertebrates in the 

respective chapters.

Additional Research and Monitoring Needed

As mentioned earlier, it is likely that most of the 

new species to be added to the district’s flora 

will be found in the middle-elevation areas of 

the district, particularly on the east slope of the 

Rincon Mountains. In addition, invasive, nonnative species will likely become established 

in high-traffic areas such as the Cactus Forest 

Loop Drive and Old Spanish Trail, where the 

park staff have been surveying for them for four 

years. Future funding for the park-based effort 

is uncertain and the SODN I&M program is 

establishing protocols for periodic surveys in 

these areas. Considerable effort has been focused 

on determining the effects of fire on the high 

elevation plant community and we encourage 

the park to analyze and report the results of the 

fire-effects monitoring program. Finally, there 

are a number of long-term monitoring plots for 

saguaros that have not been relocated. These and 

other, recently located plots should be resurveyed 

periodically. Finally, the district is also in need 

of a current, detailed vegetation map, which will 

likely be created in the next few years by the 

I&M program (Andy Hubbard, pers. comm.). 

Vegetation monitoring will be an 

important component of the I&M program at 

Saguaro National Park and other park units in 

the Sonoran Desert Network (Mau-Crimmins 

et al. 2005), yet field methods for vegetation 

monitoring have not been established. Our use 

of the modified-Whittaker and point-intercept 

methods provides data that could inform that 

program. If the goal of the I&M program is to 

monitor species richness or species composition, 

a plot-based method such as the modifiedWhittaker may be more appropriate than the 

point-intercept method because more species 

were observed on plots and the point-interecept 

transects missed many species in the area of the 

transects. However, observer bias in estimating 

species coverage (a measure of dominance) is an 

important limitation of the modified-Whittaker 

and similar methods for monitoring that 

parameter. In fact, estimation of coverage can be 

so great as to obscure trend detection for all but 

the most extreme changes (Kennedy and Addison 

1987). Bias can be minimized by reducing the 

size of the quadrat (Elzinga et al. 2001). With 

regard to observer bias, the point-intercept (or the 

similar line-intercept method) produce less biased 

estimates of species coverage because there is 

less opportunity for interpretation. Elzinga et 

al. (2001) provide an excellent overview of the 

major survey methods for monitoring vegetation 

and they include a good discussion of observer 

bias. 

If the goal of the monitoring program is 

to monitor changes in vegetation structure and 

gross vegetation characteristics (i.e., dominant 
25

plant species), then the point-intercept method 

is likely the more appropriate of the two 

methods. Because we spaced 50 m transects 

systematically throughout the 1 km focal point 

transect, estimates of coverage were likely more 

representative of the study area than the single 

20 m x 50 m modified-Whittaker plot. Further, 

accuracy of estimates from point-intercept 

transects and quantification of the vegetation 

heterogeneity can be assessed by using estimates 

from each 50 m transect section. Estimates 

of accuracy and heterogeneity for modifiedWhitaker plots can also be accomplished by 

establishing multiple plots. 

Powell et al. (2005) and others (I&M 

program, unpubl. data) used similar field 

methods as reported here and found many 

of the same patterns with regards to species 

richness and coverage estimates at nearby 

Tumacácori National Historical Park. Their 

use of “modular” plots (where point-intercept 

transects were established within Braun-Blanquet 

plots [similar to modified-Whitaker plots; BraunBlanquet 1965]) will provide for a more rigorous 

comparison of those two methods. Regardless 

of the field method chosen, the use of plot or 

transect-based field surveys should be used 

in combination with remote sensing, which is 

becoming an invaluable tool for monitoring 

vegetation change (Frohn 1998). 
26
27

Chapter 4: Amphibian and Reptile Inventory

Aaron D. Flesch, Don E. Swann, and Brian F. Powell

Previous Research 

Little information is available on the distribution, 

abundance, and habitat of amphibians and reptiles 

(hereafter herpetofauna) in the Rincon Mountain 

District, though the community composition is 

well known and several species lists exist (Black 

1982, Doll et al. 1986, Lowe and Holm 1991, 

Swann 2004). Because of poor documentation, 

we do not consider the lists of Black (1982) 

or Doll et al. (1986). Lowe and Holm (1991) 

ranked abundance (e.g. rare, uncommon, and 

common) of herpetofauna in the district, but these 

categories were from incidental observations, not 

formal surveys within the district. Lowe (1992) 

summarized some information on distribution of 

herpetofauna in the district but focused mainly 

on providing a regional biogeographic context 

for understanding distribution patterns. Goode 

et al. (1998) inventoried the district’s Expansion 

Area in Rincon Valley and Murray (1996) and 

Swann (1999b) inventoried both the Expansion 

Area and the nearby Rocking K Ranch and 

provided detailed information for these areas. 

Most recently, Bonine and Schwalbe (2003) 

inventoried the Madrona Pools of Chimenea 

Creek but their effort was limited to only five 

days in May. There have also been a number of 

single-species studies in the district, including 

those for the lowland leopard frog (Swann 1997, 

Swann et al. 2003b, Goldberg et al. 2004, Eric 

Wallace, unpubl. data), desert tortoise (Swann 

et al. 2002, Stitt et al. 2003, Edwards et al. 2004, 

Jones et al. 2005), and tiger rattlesnake (Matt 

Goode, unpubl. data). Because most previous 

studies have been limited either spatially or 

temporally, the inventory effort summarized 

in this report represents the first attempt to 

quantify distribution and abundance and provide 

information on habitat of all amphibian and 

reptile species in the district. 

Methods

We surveyed herpetofauna in 2001 and 2002 

using four field methods: (1) plot-based intensive 

surveys, (2) non-plot based extensive surveys 

(Table 4.1), (3) road surveys, and (4) incidental 

observations. We used multiple methods 

to ensure coverage across a broad range of 

environmental features and to facilitate complete 

species lists and estimates of relative abundance. 

We chose the location of intensive surveys (at 

focal-point transects) using a stratified random 

design and stratified by elevation (see Chapter 

1) then constrained surveys by time and area 

(Crump and Scott 1994). We chose the location 

of extensive surveys both randomly and nonrandomly; some extensive surveys were located 

Table 4.1. Characteristics of three major active survey methods used during surveys for herpetofauna, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002. 

Survey type

Characteristic Intensive, plot-based Extensive – Random Extensive – Non-random

Random location Yes Partially No

Area constrained Yes No No

Configuration Plot based visual encounter Non-plot based visual encounter Non-plot based visual encounter

Area (ha) three 1-ha plots per transect Variable Variable

Time constrained Yes, 1 hour No No

Time of day Morning Morning Morning, afternoon, and evening 

Advantages Facilitates comparison with other areas, 

scope of inference to entire park, more 

complete richness and abundance data

Larger scope of inference and potential 

to detect less common species 

Maximum flexibility facilitating 

detection of rare species with 

restricted distributions 

Disadvantages Inefficient for developing complete 

species list

Inefficient for developing complete 

species list

Scope of inference applies only to 

those areas surveyed
28

near intensive plots, but most were in areas we 

thought would have high species richness, species 

of special interest, or species suspected to be in 

the district that had not previously been recorded 

(e.g., the rock rattlesnake). Extensive surveys 

were more flexible and allowed for variation in 

survey time and area. For road and extensive 

surveys, we surveyed in evenings and nights to 

detect species with restricted activity periods 

(Ivanyi et al. 2000). Although we designed 

methods to detect both amphibians and reptiles, 

we detected fewer amphibians because they 

have more limited activity periods and are often 

restricted to aquatic environments, which are rare 

in the district. 

Intensive Surveys

Field Methods

At focal-point transects (hereafter “transects”) 

in 2001, we used plot-based visual encounter 

surveys constrained by time and area (Crump 

and Scott 1994) along 17 transects (Figs. 4.1, 

4.2). Along each transect we surveyed within 

the confines of three 1-ha (100 x 100 m) subplots 

during spring (9 April - 24 May) or two subplots 

during the summer monsoon (18–31 July) and 

searched each subplot for one hour. We surveyed 

only two subplots in summer because there 

was not sufficient time during peak activity 

periods to search all three subplots. Although 

we surveyed all 17 transects in spring only seven 

transects were surveyed in summer and these 

were located only in low (n = 3) and middle (n

= 4) elevation strata. We selected survey times 

that coincided with periods of peak diurnal 

reptile activity because activity levels vary with 

temperature (Rosen 2000). On cooler spring days 

we began our surveys between 0718 and 1421 

hours whereas on hotter, summer days we began 

between 0642 and 1014 hours. To account for 

within-day variation in detectability and to reduce 

variation among observers, we surveyed each 

subplot twice per day by a different observer. We 

did not survey during evenings or nights. 

We searched subplots visually and aurally 

and worked systematically across each subplot 

and used a Garmin E-map GPS to ensure we 

stayed within subplot boundaries during surveys. 

We also looked under rocks and litter and used 

a mirror to illuminate cracks and crevices. For 

each animal detected, we recorded species, sex 

and age/size class (if known), and microhabitat 

(ground, vegetation, rock, edifice, burrow, or 

water). We marked subplot corners with rubbercapped stakes and recorded UTM coordinates 

with a Trimble GPS. We recorded temperature, 

wind speed (km/h), percent relative humidity, 

and percent cloud cover using hand-held Kestrel 

3000 weather meters (Nielson-Kellerman Inc., 

Boothwyn, PA) before and after surveys. We also 

described vegetation and soils. 

Transect

line

1000 m 

1 2 3 4 5 6 7 8 9 10

Plot Number 

Focal

Point

100 m 

100 m 

Figure 4.1. Layout of herpetofauna survey plots along focal-point transects, Saguaro National Park, 

Rincon Mountain District, 2001. We typically surveyed three, 100 x 100 m subplots (dotted boxes) in the 

spring and two subplots (1 and 10) in the summer. When topography prevented surveys in a subplot, we 

surveyed an adjacent subplot.
29

Figure 4.2. Locations of intensive and extensive survey sites for herpetofauna, Saguaro National Park, 

Rincon Mountain District, 2001 and 2002. 

Table 4.2. Herpetofaunal survey effort by year, Saguaro National Park, Rincon Mountain District, 2001 

and 2002. 

2001 2002

Method Elevation range (m)

No. of samples 

(subsamples)a

Survey 

hours

No. of samples 

(subsamples)a

Survey 

hours

Intensive survey 936 – 2,560 17(51) 131.0

Extensive survey – random 850 – 2,119 22 88.0

Extensive survey – non-random 818 – 2,634 58 359.2 5 18.0

Road survey 53 45.8 2 0.5

a No. of subsamples for random surveys equals number of subplots per focal-point transect for intensive surveys, number of survey 

areas for extensive surveys.
30

Effort

We completed 131 surveys at 51 subplots located 

along the 17 focal-point transects (Table 4.2, Fig. 

4.2). In 2002 we discontinued intensive surveys 

because of the relatively low number of species 

detected.

Analysis

We calculated relative abundance of each species 

for each transect by summing all detections 

within the two or three subplots surveyed per 

transect. Because subplots were surveyed twice 

per day, we accounted for within-day variation 

in detectability by using the maximum number 

of individuals detected on either survey for each 

visit because it represented abundance when 

detectability was highest (Rosen and Lowe 1995). 

We estimated relative abundance (no./ha/hr) of 

each species (and all species combined) within 

the district by averaging the maximum number 

of individuals detected on repeated visits to 

each transect, and then averaging results from 

all transects. To compare relative abundance 

of each species (and all individuals combined) 

among elevation strata, we compared the average, 

maximum number detected on all 17 transects 

surveyed in spring among elevation strata using 

ANOVA. To compare relative abundance 

between seasons, we compared the average, 

maximum number detected between seasons for 

the seven transects surveyed in both spring and 

summer (transect nos. 101, 106, 111, 112, 115, 

130, and 139) using paired t-tests. We did not 

compare estimates from summer among strata 

because only low- and-middle elevation transects 

were surveyed and sample sizes were small.

To determine environmental factors that 

explained variation in relative abundance of 

species and species groups and species richness, 

we used multiple linear regression with stepwise 

selection (P < 0.20 to enter, P < 0.05 to stay) and 

22 potential explanatory factors (Table 4.3; from 

point-intercept vegetation sampling; see Chapter 

3). Because data for most species were limited, 

we only considered those with ≥15 observations 

and combined all species of whiptails and all 

other species of lizards except whiptails in 

analyses. We screened explanatory factors before 

modeling and retained only what we judged to 

be the most biologically meaningful factor from 

correlated pairs (r > 0.75) and used Cp statistics 

to guide model selection (Ramsey and Schafer 

2002). Where necessary, we transformed factors 

using log(x) or log(x + 1) to improve normality. 

Extensive Surveys

Non-plot based extensive surveys (referred to 

as “special areas” in Powell et al. 2002, 2003) 

facilitated sampling in areas where we expected 

high species richness, abundance, or species 

not previously detected. Typically, we selected 

areas for extensive surveys in canyons or riparian 

areas, and also included ridgelines, cliffs, rock 

piles, bajadas, summits, or other physiographic 

Table 4.3. Environmental factors considered when modeling variation in relative abundance of species 

and species groups and species richness of herpetofauna, using stepwise multiple linear regression, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002. Data from point-intercept transects 

(height category) and modified-Whittaker plots (plots). 

Height category

Environmental factor (units) basal 0-0.5 m 0.5-2.0 m 2.0-4.0 m >4.0 m plot

Bare ground cover (%) x

Rock cover (%) x

Forb cover (%) x x x

Grass cover (%) x x x

Tree cover (%) x x x x

Shrub cover (%) x x x x

Vegetation cover (all life forms, %) x x x x

Plant species richness (no.) x

Slope (%) x
31

features. We based extensive surveys on visual 

encounters (Crump and Scott 1994) and, in 

contrast to intensive surveys, did not constrain 

surveys by area or time. We focused extensive 

surveys during mornings and also surveyed 

during evenings and nights in low-elevation areas 

when detectability of snakes and amphibians is 

often highest (Ivanyi et al. 2000), and during midday at higher elevations. 

Field Methods

We selected areas randomly and non-randomly 

(Table 4.1). We placed random survey areas 

within approximately 1 to 2 km of focal point 

transects, and surveyed each area once. We 

selected non-random areas by using topographic 

maps and prior knowledge of the district. We 

relied upon visual detection and often looked 

under objects and illuminated cracks to detect 

hidden individuals. We surveyed in spring (4 

April – 24 May) and summer (25 June – 20 

September) of 2001 and 2002. One, two, or three 

observers searched each area simultaneously 

and recorded data separately. Total duration of 

surveys among all observers combined averaged 

5.5 ± 0.4 (± SE) hours per survey (range = 1.2 

- 20.4 hours). We recorded data using similar 

methods as intensive surveys and noted UTM 

coordinates and elevation at the start and end 

points of each survey. 

Effort

We surveyed 85 areas in 2001 and 2002 (Fig. 

4.2), 94.1% of which were surveyed in 2001 

(Table 4.2). Total survey effort was 465.2 hours, 

81% of which was in non-random areas. Survey 

effort was roughly three times greater than 

for other methods and focused mainly during 

daylight except at lower elevations where we also 

surveyed during late evenings and nights. We did 

not survey higher elevation areas in late evenings 

and at night because detectability declined 

markedly with elevation. 

Analysis

We calculated relative abundance for each area 

as the number of individuals detected for each 

species or all species combined per 10 hours of 

effort. For surveys completed by >1 observer 

per area, we summed survey times and detection 

data for all surveyors when calculating effort and 

relative abundance for an area. Although some 

locations were surveyed multiple times, survey 

routes often varied and we therefore considered 

each survey an independent sample despite some 

spatial overlap. To describe general patterns of 

relative abundance for species groups (lizards, 

snakes, and amphibians) and species richness 

across the district, we post-stratified survey 

areas by elevation (low = <4,000 feet, middle = 

4000–6,000 feet, high = >6,000 feet) using the 

median elevation of all animal observations for 

each survey. We then tested for variation among 

strata using one-, two-, or multi-way ANOVA. 

Because relative abundance and species richness 

varied between day and night and no areas 

were surveyed during night at middle and high 

elevations, we limited comparisons only to 

days. To describe patterns of relative abundance 

of individual species across elevation, we used 

multiple linear regression. We transformed 

relative abundance values when necessary 

using log(x) or log(x + 1) to improve normality. 

Because patterns of relative abundance often 

varied with relative humidity (or cloud cover), 

season, and time of day, we adjusted for these 

factors when they explained variation (P ≤ 

0.10) in relative abundance. To describe cloud 

cover, relative humidity, and temperature for 

each area, we averaged measurements taken at 

the beginning and end of each survey. To adjust 

for temporal variation in relative abundance 

and richness across time of day, we considered 

three time periods: day, late evening or night, or 

surveys that spanned portions of both periods 

(day equaled reference level). We considered 

20-min before local sunset time as the cut-point 

between day and late evening or night surveys. 

To adjust for seasonal variation in relative 

abundance and richness, we considered two 

seasons, spring and summer (spring equaled 

reference level). Because relative humidity and 

cloud cover were strongly correlated (r = 0.76, 

P < 0.0001) we only adjusted for the factor that 

explained the most variation in responses. 
32

Road Surveys 

Road surveys involve driving slowly along a 

road, typically after sunset, and watching for 

animals. Such surveys are a common method 

for estimating distribution and abundance 

of herpetofauna and are recommended for 

augmenting species lists (Shaffer and Juterbock 

1994). 

Field Methods

We focused mainly on the Cactus Forest Loop 

Drive and also drove Speedway Boulevard from 

Douglas Spring Trailhead to the intersection with 

Tanque Verde Loop Road and Camino Loma 

Alta from the trailhead to Old Spanish Trail. We 

recorded each individual detected by species 

and whether animals were alive or dead. We 

surveyed 29 April – 18 August 2001 and 9 – 14 

July 2002 during nights and occasionally during 

evenings.

Effort

We conducted 55 road surveys totaling 46.3 hours 

of effort (Table 4.2).

Analysis

Because survey routes varied in length and 

included a number of different segments surveyed 

in various orders, we combined results from 

all routes and road segments. Total mileage 

for each route was not recorded so we scaled 

estimates of relative abundance by time. We 

calculated relative abundance as the number 

of individuals detected for each species (or all 

species combined) per hour of effort. We also 

compared relative abundance of species groups 

across months using ANOVA and linear contrasts. 

We log (x + 1) transformed relative abundance to 

improve normality. 

Incidental Observations

We noted sightings of rare or important species 

by sex and age/size class (if known) and recorded 

time of observations and UTM coordinates 

for all detections. These incidental detections 

were often recorded before or after more formal 

surveys and we use these sightings to determine 

species presence and richness. We also used 

incidental sightings from other field crews (e.g., 

birds). 

Species Identification Challenges

Whiptail lizards (Cnemidophorus [Aspidoscelus 

by some sources] spp.) are notoriously difficult 

to identify in the field because of the similarity 

in appearance for several sympatric species 

(Stebbins 2003). Many parthenogenetic (nonsexually reproducing) whiptails may have arisen 

as hybrids from the same diploid, sexually 

reproducing parent species (Degenhardt et al. 

1996). Several undescribed “parthenospecies” 

may exist in the desert Southwest (Wright and 

Vitt 1993, Cole and Dessauer 1994). Some 

individuals we identified as western (C. tigris) or 

Sonoran spotted (C. sonorae) whiptails may be 

undescribed “species” related to these recognized 

species. 

In the district we saw “classic” Sonoran 

spotted whiptails (adults with six longitudinal 

dorsal stripes, light spots in dark and occasionally 

light dorsal areas; dorsal stripes more yellow 

anteriorly; overall color brown dorsally and 

unmarked white-cream ventrally; tail more 

brownish-orange than bluish as seen in Gila 

spotted whiptails; Degenhardt et al. 1996, Phil 

Rosen pers. obs.). We also observed a variation 

of this classic appearance that superficially 

resembled Gila spotted whiptails, with some 

captured individuals keying out to be this 

species based on characteristics noted in field 

guides, including number of pre-anal scales, 

location of spots in light stripes, and greenish tail 

(Stebbins 2003). Although the Rincon Mountains 

are considered outside the range of the Gila 

spotted whiptail, in this document we report 

these individuals as this species, and report the 

“classic” Sonoran whiptails described above as 

Sonoran spotted whiptails.

Results

We detected 46 species of herpetofauna; seven 

amphibians and 39 reptile species (Appendix 
33

B). Reptilian species included two turtle, 19 

lizard, and 18 snake species. Species richness 

was highest for incidental (n = 43) and extensive 

surveys (n = 39) and lowest for intensive (n = 

25) and road surveys (n = 22). We found seven 

species with only a single survey method, but 

all other species were found with two or more 

methods. Road and extensive surveys each 

yielded detection of one species that was not 

detected by using other methods (Great Plains 

toad, and Great Plains skink, respectively) and 

incidental surveys yielded detection of five 

species not detected by using other methods 

(Mexican spadefoot, canyon spotted whiptail, 

ring-necked snake, western ground snake, and 

Mojave rattlesnake). All 25 species that we 

detected during intensive surveys were detected 

using other methods, although Madrean alligator 

lizard was detected only during intensive and 

extensive surveys.

We detected 4,292 individuals during 

this study – 3,066 during intensive, extensive, 

and road surveys combined, and 1,225 incidental 

observations (Appendix B). Most individuals 

(1,909) were detected during extensive surveys 

and fewest (469) were detected during road 

surveys (Table 4.4). The number of individuals 

detected per unit time was greatest for road 

surveys (mean = 14.9 individuals/hr) markedly 

higher than for extensive (4.1 individuals/hr) 

or intensive (3.6 individuals/hr) surveys. The 

species with the most detections (all methods 

combined) was the ornate tree lizard (n = 750). 

We recorded 11 species <5 times (Appendix B). 

A review of our inventory effort and 

other efforts in the district indicates that the 

district supports 57 species of herpetofauna: nine 

amphibians and 48 reptiles (Appendix B). All but 

five species have been confirmed with a specimen 

and/or photographic voucher (Appendices E, 

F). Our inventory did not result in detection 

of species not already recorded in the district, 

although we produced the first documentation (in 

the form of specimen and photographic voucher) 

for a number of species, including the Mojave 

rattlesnake. 

Intensive Surveys

We detected 469 individuals of 22 species of 

amphibians and reptiles along 17 transects during 

131 hours of effort (Table 4.5). Lizards were 

most common and comprised 50.0% (n = 11 of 

22) of species and 92.8% (n = 435 of 469) of 

individuals whereas snakes comprised 36.4% (n = 

8 of 22) of species and only 4.1% (n = 19 of 469) 

of individuals. We recorded only two species 

of amphibians (Sonoran desert toad and canyon 

treefrog), comprising 2.8% (n = 13 of 469) of 

individuals. Relative abundance averaged 4.6 ± 

0.6 individuals/ha/hr (range = 1.0 – 11.0) for all 

species and strata combined. 

During the spring, when we surveyed all 

17 transects, the ornate tree lizard was the most 

abundant species (2.08 ± 0.41/ha/hr) followed 

by Clark’s spiny and eastern fence lizards (Table 

4.6). Collectively, whiptail lizards were also 

common (mean ± SE = 1.48 ± 0.17/ha/hr), yet 

51.4% could not be identified to species. Of 

whiptails that could be identified to species, 

Gila spotted and Sonoran spotted were equally 

abundant (0.29/ha/hr) and western whiptails were 

less abundant (0.14 ± 0.09/ha/hr). The desert 

tortoise, western banded gecko, and eastern 

collared lizard were rarest (one detection each). 

The Sonoran whipsnake and black-necked garter 

snake were the most common snakes.

Table 4.4. Number of animals and species detected per hour during herpetofaunal surveys by 

year and survey method, Saguaro National Park, Rincon Mountain District, 2001 and 2002. 

2001 2002

Survey type

Animals 

detected

Animals 

per hour 

Species 

detected

Species 

per hour 

Animals 

detected

Animals 

per hour 

Species 

detected

Species per 

hour 

Intensive 469 3.6 22 0.17

Extensive 1818 4.1 39 0.09 91 5.1 16 0.9

Road 654 14.3 18 0.39 34 68.0 5 10.0
34

Species richness and relative abundance 

often varied among elevation strata (Table 4.5). 

Species richness was highest at lower elevation 

(5.2 ± 0.5), moderate at middle elevation (4.3 

± 0.4), and low at the high elevation (2.8 ± 0.5) 

(F2,14 = 6.86, P = 0.0084, ANOVA). Relative 

abundance of all individuals combined varied 

among elevation strata and patterns were similar 

to those for species richness (F2,14 = 5.62, P = 

0.016), yet relative abundance was similar at lowand middle-elevation strata (t

14 = 0.77, P = 0.46, 

linear contrast; Table 4.5). The common sideblotched lizard, greater earless lizard, and western 

whiptail were found only in the low-elevation 

stratum (F2,14 ≥ 2.78, P ≤ 0.096) whereas relative 

abundance of the Clark’s spiny lizard and 

Sonoran spotted whiptail were similar at lowand middle-elevation strata and were either rare 

(Clark’s spiny) or did not occur (Sonoran spotted) 

at the high-elevation stratum (F2,14 ≥ 3.07, P ≤ 

0.079). Eastern fence lizards were not found at 

the low-elevation stratum and relative abundance 

was roughly two times higher at the high- as 

compared to the middle-elevation stratum (F2,14

= 4.63, P = 0.029). Relative abundance seemed 

to vary among elevation strata for other species 

(Table 4.5), though detections were too few for 

quantitative comparisons. 

Species richness and relative abundance 

varied between seasons for some species and 

species groups. Species richness for all taxa 

combined averaged 5.0 ± 0.5 species/ transect 

in both spring and summer (t

6

 = 0.33, P = 0.38, 

paired t-test) yet species richness of lizards in 

spring (4.3 ± 0.4) averaged 0.9 species greater 

than in summer (t

6

 = 2.12, P = 0.039). Relative 

abundance of all species combined did not vary 

between seasons (t

6

 = 0.27, P = 0.40) yet relative 

Table 4.5. Relative abundance (mean + SE; no./ha/hr) of herpetofauna detected during intensive surveys 

in spring (9 April - 24 May) along focal point-transects by elevation strata, Saguaro National Park, Rincon 

Mountain District, 2001. Species for which there are no detections were detected only in summer (18-31 July) in 

low- and/or middle-elevation strata. 

Elevation stratum

Low (n = 5) Middle (n = 7) High (n = 5) All (n = 17)

Species Mean SE Mean SE Mean SE Mean SE

Sonoran Desert toad

canyon treefrog

desert tortoise 0.07 0.07 0.02 0.02

western banded gecko 0.07 0.00 0.02 0.02

eastern collared lizard 0.07 0.07 0.02 0.02

greater earless lizard 0.20 0.13 0.06 0.04

Clark’s spiny lizard 1.13 0.47 1.00 0.29 0.07 0.07 0.76 0.21

eastern fence lizard 0.62 0.29 1.00 0.11 0.55 0.15

common side-blotched lizard 0.20 0.08 0.06 0.03

ornate tree lizard 2.47 0.74 2.62 0.72 0.93 0.37 2.08 0.41

unknown whiptail 1.73 0.52 0.57 0.30 0.07 0.07 0.76 0.24

Sonoran spotted whiptail 0.33 0.18 0.48 0.10 0.29 0.08

Gila spotted whiptail 0.13 0.13 0.33 0.18 0.29 0.09

western whiptail 0.47 0.00 0.14 0.09

Madrean alligator lizard 0.07 0.07 0.05 0.05 0.04 0.03

coachwhip

Sonoran whipsnake 0.07 0.07 0.05 0.05 0.07 0.07 0.06 0.03

western patch-nosed snake

black-necked garter snake 0.07 0.07 0.10 0.06 0.06 0.03

western diamond-backed rattlesnake

black-tailed rattlesnake

tiger rattlesnake

western rattlesnake 0.05 0.05 0.07 0.07 0.04 0.03

all individuals 5.87 0.75 5.00 0.95 2.00 0.42 4.37 0.59
35

abundance of ornate tree lizards and all whiptail 

lizards combined were roughly two times greater 

in the spring (t

6

 ≥ 1.91, P ≤ 0.53) (Table 4.6). The 

desert tortoise, western banded gecko, Madrean 

alligator lizard, and western rattlesnake were 

detected only in spring, whereas the Sonoran 

Desert toad, canyon treefrog, coachwhip, and 

western diamond-backed, black-tailed, and tiger 

rattlesnakes were detected only in summer (Table 

4.6). Eastern collared lizards were not detected 

in spring except in the high-elevation stratum 

(Table 4.5). Some of these patterns may have 

been the result of low sample size, because in the 

cases of western rattlesnake and collared lizards, 

the patterns that we observed are opposite to the 

known natural history of each species. 

Environmental factors that explained 

patterns of species richness and relative 

abundance varied (Table 4.7). Snake richness 

increased with cover of grasses whereas lizard 

richness decreased with increasing cover of 

bare ground. Species richness of snakes and 

lizards increased with shrub cover above 

2 m, though influence of shrub cover was 

much greater for snakes; richness of lizards 

decreased with tree cover between 0.5 and 2.0 

m. Relative abundance (no./ha/hr) of all lizard 

species combined declined with increasing 

cover of bare ground. For all lizards excluding 

whiptails, however, relative abundance decreased 

as grass cover between 0.5 and 2.0 m above 

ground increased, whereas for whiptails relative 

abundance decreased as vegetation cover between 

0.5 and 2.0 m of all plant types combined 

increased. In contrast to patterns for all species 

of lizards combined, relative abundance of 

eastern fence lizards increased with increasing 

cover of bare ground. Relative abundance of 

the Sonoran spotted whiptail and Clark’s spiny 

lizard was positively associated with forb cover 

between 0 and 0.5 m above ground, whereas 

relative abundance of ornate tree lizards was 

positively associated with grass cover in the same 

vegetation stratum. Relative abundance was not 

Table 4.6. Relative abundance (mean + SE; no./ha/hr) of herpetofauna detected during intensive surveys 

along random transects (n = 7) surveyed in both spring (9 April – 8 May) and summer (18 – 31 July), 

Saguaro National Park, Rincon Mountain District, 2001. 

Spring (n = 7) Summer (n = 7) All seasons

Species Mean SE Mean SE Mean SE

Sonoran Desert toad 0.43 0.43 0.21 0.21

canyon treefrog 0.07 0.07 0.04 0.04

desert tortoise 0.05 0.05 0.02 0.02

western banded gecko 0.05 0.05 0.02 0.02

eastern collared lizard 0.07 0.07 0.04 0.04

greater earless lizard 0.14 0.10 0.14 0.14 0.14 0.08

Clark’s spiny lizard 1.24 0.33 1.93 0.70 1.58 0.39

common side-blotched lizard 0.14 0.07 0.14 0.09 0.14 0.05

ornate tree lizard 2.14 0.43 1.14 0.48 1.64 0.34

unknown whiptail 1.43 0.43 0.21 0.15 0.82 0.28

Sonoran spotted whiptail 0.48 0.16 0.50 0.29 0.49 0.16

Gila spotted whiptail 0.14 0.14 0.29 0.15 0.21 0.10

western whiptail 0.33 0.19 0.07 0.07 0.20 0.11

Madrean alligator lizard 0.10 0.06 0.05 0.03

coachwhip 0.07 0.07 0.04 0.04

Sonoran whipsnake 0.10 0.06 0.14 0.09 0.12 0.05

western patch-nosed snake 0.07 0.07 0.04 0.04

black-necked garter snake 0.10 0.06 0.14 0.14 0.12 0.08

western diamond-backed rattlesnake 0.07 0.07 0.04 0.04

black-tailed rattlesnake 0.07 0.07 0.04 0.04

tiger rattlesnake 0.07 0.07 0.04 0.04

western rattlesnake 0.05 0.05 0.02 0.02

all individuals 5.48 0.68 5.07 1.33 5.27 0.72
36

The black-necked garter snake and 

western diamond-backed rattlesnake were the 

most common snakes and western and mountain 

patch-nosed snakes, Sonoran coral snake, and 

common kingsnake were the rarest.

We surveyed most areas during day 

(63.5%) with fewer surveyed in late evening or 

night (10.6%) or spanning both periods (25.9%) 

and all surveys in middle- and high-elevation 

areas were completed during day. At the lowelevation stratum, relative abundance varied 

among survey periods for lizards and amphibians 

(F2,47 ≥ 5.30, P ≤ 0.0084, ANOVA), with 4.7 

times more lizards detected during day (61.8 ± 

7.0) and 3.6 times more amphibians detected 

during late evening and night (31.4 ± 6.3). 

Relative abundance among survey periods did not 

vary for snakes (F2,47 = 0.35, P = 0.71, ANOVA). 

Species richness did not vary among survey 

periods for amphibians, snakes, and all groups 

combined (F2,47 ≤ 2.25, P ≥ 0.12, ANOVA), but 

explained by cover of rock after accounting for 

other factors for all species of lizards combined 

(t

22 ≤ 1.25, P ≥ 0.23), all lizards excluding 

whiptails (t

16 = 0.20, P ≤ 0.85), and all whiptails 

combined (t

16 = 1.15, P = 0.27).

Extensive Surveys

We detected 1,909 individuals of 39 species in 

85 survey areas in 2001 and 2002 (Table 4.8). 

We detected 428 amphibians of five species 

and 1,481 reptiles that included two turtle, 

18 lizard, and 14 snake species, with lizards 

comprising 85.1% (n = 1,261 of 1,481) of all 

reptiles combined. Overall, relative abundance 

averaged 44.6 ± 4.4 individuals/10 hours (range = 

0–177.5) and was highest for lizards (27.0 ± 3.5 

no./10 hours) and lowest for amphibians (4.7 ± 

0.9 no./10 hours). The ornate tree lizard, canyon 

treefrog, Clark’s spiny lizard, and Sonoran Desert 

toad were the most common species (Table 4.8). 

Table 4.7. Environmental factors that explained relative abundance (no./ha/hr) of species (with >15 

observations), species groups, and species richness of lizards and snakes detected during intensive 

surveys, Saguaro National Park, Rincon Mountain District, spring 2001. 

Category

 Species or group

 Environmental factor estimate SE t P

Relative abundance

 Clark’s spiny lizard

 Forb cover 0-0.5 m above ground (%) 0.13 0.05 2.53 0.024

 Tree cover 0-0.5 m above ground (%) -0.25 0.10 2.51 0.025

 eastern fence lizard

 Bare ground basal cover (%) 0.04 0.01 5.93 <0.0001

 Grass cover 0-0.5 m above ground (%) -0.10 0.02 5.14 0.0002

 Grass cover 0.5-2.0 m above ground (%) 0.26 0.06 4.39 0.0007

 ornate tree lizard

 Bare ground basal cover (%) -0.09 0.03 2.59 0.021

 Grass cover 0-0.5 m above ground (%) 0.24 0.08 3.04 0.0088

 Sonoran spotted whiptail

 Forb cover 0-0.5 m above ground (%) 0.08 0.02 3.95 0.0013

 all whiptails

 Bare ground basal cover (%) -0.11 0.02 5.00 0.0002

 Vegetation cover 0.5-2.0 m above ground (%) -0.10 0.04 2.44 0.029

 all lizards excluding whiptails

 Bare ground basal cover (%) -0.15 0.04 3.88 0.0017

 Grass cover 0.5-2.0 m above ground (%) 0.82 0.27 3.01 0.0093

Species richness

 lizards

 Bare ground basal cover (%) -0.03 0.004 6.48 <0.0001

 Tree cover 0.5-2.0 m above ground (%) -0.03 0.01 3.40 0.0047

 Shrub cover 2.0-4.0 m above ground (%) 0.38 0.10 3.94 0.0017

 snakes

 Grass basal cover (%) 0.09 0.02 4.02 0.0013

 Shrub cover >4.0 m above ground (%) 2.05 0.63 3.24 0.0059
37

did vary for lizards (F2,47 = 14.6, P < 0.0001, 

ANOVA), with 2.3 times more species detected 

during day (4.8 ± 0.4) than other periods. 

Relative abundance of amphibians 

increased by 1.2 ± 0.6 individuals/10 hours with 

each 10% increase in cloud cover after adjusting 

for the influence of survey time (t

79 = 1.96, P = 

0.054, test of slope from regression). In contrast, 

relative abundance of lizards decreased by 2.2 ± 

0.7 individuals/10 hours with each 10% increase 

in cloud cover after adjusting for survey time and 

elevation (t

77 = 3.21, P = 0.0019) but did not vary 

Table 4.8. Relative abundance (mean + SE; no./10 hrs) of herpetofauna detected during extensive surveys 

(n = 85), by elevation strata, Saguaro National Park, Rincon Mountain District, 2001 and 2002. Parenthetical 

numbers are sample sizes for number of survey areas. 

Elevation stratum

Low (n = 50) Middle (n = 23) High (n = 12) All (n = 85)

Species mean SE mean SE mean SE mean SE

Couch’s spadefoot 1.36 1.06 0.80 0.63

Sonoran Desert toad 4.97 2.19 2.92 1.31

red-spotted toad 1.90 0.86 0.10 0.10 1.13 0.52

canyon treefrog 2.50 0.53 11.15 5.83 4.29 3.28 5.10 1.69

lowland leopard frog 1.80 1.04 0.33 0.33 1.15 0.62

Sonoran mud turtle 0.91 0.42 0.84 0.48 0.76 0.28

desert tortoise 0.53 0.20 0.35 0.26 0.41 0.14

western banded gecko 0.39 0.15 0.23 0.09

eastern collared lizard 0.10 0.07 0.26 0.26 0.13 0.08

lesser earless lizard 0.10 0.08 0.06 0.05

greater earless lizard 1.43 0.35 0.35 0.26 0.94 0.23

zebra-tailed lizard 2.29 1.14 1.35 0.68

desert spiny lizard 0.87 0.27 0.51 0.17

Clark’s spiny lizard 4.59 0.87 3.41 1.27 0.33 0.33 3.67 0.63

eastern fence lizard 4.16 1.54 6.20 2.37 2.00 0.59

common side-blotched lizard 3.94 1.22 2.32 0.75

ornate tree lizard 10.03 2.61 10.47 2.33 1.65 0.95 8.96 1.69

greater short-horned lizard 0.44 0.30 0.06 0.04

regal horned lizard 0.15 0.11 0.09 0.06

Great Plains skink 0.04 0.04 0.02 0.02

unknown whiptail 1.25 0.48 1.08 0.39 1.03 0.30

Sonoran spotted whiptail 3.40 1.40 1.38 0.61 2.37 0.85

Gila spotted whiptail 0.26 0.13 1.36 0.92 0.42 0.28 0.58 0.26

western whiptail 1.44 0.68 0.84 0.41

Madrean alligator lizard 0.03 0.03 0.52 0.37 0.08 0.05

Gila monster 0.57 0.25 0.33 0.15

coachwhip 0.21 0.12 0.12 0.07

Sonoran whipsnake 0.15 0.08 0.17 0.17 0.13 0.07

western patch-nosed snake 0.04 0.04 0.02 0.02

mountain patch-nosed snake 0.04 0.04 0.02 0.02

gopher snake 0.07 0.05 0.11 0.11 0.06 0.03

common kingsnake 0.02 0.02 0.01 0.01

Sonoran mountain kingsnake 0.08 0.08 0.33 0.33 0.07 0.05

long-nosed snake 0.08 0.05 0.05 0.03

black-necked garter snake 2.85 1.38 1.77 0.76 0.19 0.19 2.18 0.84

Sonoran coral snake 0.03 0.03 0.02 0.02

western diamond-backed rattlesnake 1.62 0.41 0.95 0.26

black-tailed rattlesnake 0.23 0.12 0.64 0.29 0.32 0.32 0.36 0.12

tiger rattlesnake 0.62 0.22 0.36 0.14

western rattlesnake 0.03 0.03 0.56 0.24 0.77 0.66 0.28 0.12

all individuals 53.73 6.05 39.80 7.69 15.89 4.22 44.62 4.37

species richness 34.00 18.00 13.00 39.00
38

with temperature (t

77 = 0.05, P = 0.95). Relative 

abundance of snakes increased with temperature 

(estimate = 0.5/Co, SE = 0.2, t

79 = 2.34, P = 

0.022) and did not vary with relative humidity 

or cloud cover (t

79 ≤ 0.77, P ≥ 0.45). Further, 

species richness decreased with increasing 

cloud cover for lizards (t

77 = 4.92, P < 0.0001) 

and increased with increasing cloud cover for 

amphibians (t

77 = 2.10, P = 0.039) after adjusting 

for the influence of survey time and elevation.

Most surveys were in the low-elevation 

stratum (58.8%) with fewer in the middle- 

(27.1%) and high- (14.1%) elevation strata. 

Relative abundance during daytime surveys 

varied among strata (F2, 51 = 12.9, P < 0.0001, 

ANOVA) and was 2-times lower in the middleand 4.2-times lower in high-elevation strata than 

in the low-elevation stratum (79.9 ± 8.3; Table 

4.7). Species richness for daytime surveys also 

varied with elevation (F2, 51 = 18.3, P < 0.0001, 

ANOVA) and was 1.7 times lower in the middleand 2.5 times lower in the high-elevation strata 

than in the low-elevation stratum (7.3 ± 0.5).

Patterns of species occurrence and 

relative abundance often varied across elevation. 

We detected 17 species in only the low-elevation 

stratum whereas we detected a single species, 

the greater short-horned lizard, in only the 

high-elevation stratum (Table 4.8). Relative 

abundance increased with elevation for eight 

species (Sonoran spotted and western whiptail, 

Clark’s spiny lizard, zebra-tailed lizard, ornate 

tree lizard, greater earless lizard, common sideblotched lizard, and western diamond-backed 

rattlesnake) and decreased with elevation for two 

species (Madrean alligator lizard and eastern 

fence lizard) (P ≤ 0.061, test of slope from 

regression) after adjusting for other important 

factors such as time of day and temperature. 

Canyon treefrogs were most common in the 

middle-elevation stratum (t

82 = 2.15, P = 0.034, 

test of quadratic term from regression).

Relative abundance of many species was 

too low or distribution too restricted to facilitate 

quantitative comparisons of species occurrence 

and relative abundance. Only a single Great 

Plains skink (along lower Chimenea Canyon) and 

lesser earless lizard (along lower Rincon Creek) 

were detected. Only one western patch-nosed 

snake (in a rocky canyon dominated by Sonoran 

desertscrub) and one mountain patch-nosed 

snake (in open pine-oak woodland at ≈1,770 m) 

were detected. Similarly only one Sonoran coral 

snake (in Sonoran desertscrub) and one common 

kingsnake (lower Rincon Creek) were detected. 

All 100 lowland leopard frogs that we observed 

Species mean SE

Couch’s spadefoot toad 1.74 1.38

Sonoran desert toad 5.74 1.52

red-spotted toad 6.04 1.64

Great Plains toad 0.06 0.06

western banded gecko 0.64 0.19

greater earless lizard 0.09 0.05

desert spiny lizard 1.85 0.80

common side-blotched lizard 0.02 0.02

ornate tree lizard 0.06 0.04

regal horned lizard 0.15 0.05

western whiptail 0.05 0.05

Gila monster 0.15 0.08

coachwhip 0.04 0.03

western patch-nosed snake 0.01 0.01

long-nosed snake 0.14 0.06

night snake 0.10 0.04

western diamond-backed rattlesnake 0.25 0.14

black-tailed rattlesnake 0.01 0.01

tiger rattlesnake 0.29 0.19

all individuals 17.48 2.72

Table 4.9. Relative abundance (no./hr) of herpetofauna detected during road surveys, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002. 
39

Sample period

0 20 40 60 80 100 120

Cumulative number of species

10

20

30

40

50

All survey methods 

Extensive and intensive surveys

Figure 4.3. Species accumulation curve for herpetofauna surveys, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. Each sampling period represents batches of 35 individuals, the 

mean number of individuals observed in an eight-hour field day. “All survey methods” includes extensive, 

intensive, road surveys, and incidental observations. The order of all sampling periods was randomized.

were detected in Turkey, Rincon, Chimenea, and 

Wildhorse creeks. 

We detected 24 species during extensive 

surveys that were near randomly-selected 

transects (n = 22) and 38 species in non-random 

areas (n = 63) with only one new species 

detected in random areas. Species richness in 

non-random areas (5.1 ± 0.4) was similar to that 

in random areas (5.3 ± 0.6; t

83 = 0.42, P = 0.81, 

t-test), yet there was some evidence that richness 

of amphibians was greater in non-random 

areas (t

83 = 1.80, P = 0.075, t-test). Relative 

abundance in non-random survey areas (41.9 ± 

5.1 individuals/10 hrs) was also similar to that 

in random areas (52.3 ± 8.6; t

83 = 1.04, P = 0.30, 

t-test), yet there was some evidence that relative 

abundance of lizards was greater in random areas 

(t

83 = 1.89, P = 0.065, t-test). 

Road Surveys

We detected 688 individuals of 19 species during 

55 surveys totaling 46.3 hours of effort (Table 

4.9). We detected four amphibian species (21% 

of all species) totaling 515 individuals, 74.9% 

of all individuals detected and proportionally 

more amphibians than for other survey methods 

(Table 4.7). Reptiles included eight lizard and 

seven snake species; 20.8% (n = 143 of 688) 

of individuals were lizards and 4.4% (n = 30 of 

688) were snakes. Relative abundance averaged 

17.5 ± 2.7 individuals/hr (range = 0– 85.3), the 

majority of which were the Sonoran Desert and 

red-spotted toads. 

Relative abundance averaged 37.1% 

higher in summer than in spring (t

53 = 1.92, P = 

0.060, t-test) but was not necessarily attributable 

to an increase in amphibians during summer 

(t

53 = 0.79, P = 0.43, t-test). The desert spiny 

lizard was the most common lizard detected (89 

detections) and the western diamond-backed 

(eight detections) and tiger (six detections) 

rattlesnakes were the most common snake species 

detected. 

Incidental Observations

We recorded 1,226 incidental detections of 44 

species between 3 April to 5 October 2001 and 

2 May to 7 November 2002 (Appendix B). 
40

All species that we detected incidentally were 

recorded using other methods except for the 

Mexican spadefoot, canyon spotted whiptail, 

ring-necked snake, western ground snake, and 

Mojave rattlesnake. 

Vouchers

We collected 10 specimen vouchers in 2001 and 

2002 (Appendix E) and obtained voucher records 

of 34 species collected by others (Appendix 

F). We obtained 65 photographic vouchers of 

44 species during 2001 and 2002 (Appendix E). 

Photographic vouchers include five amphibian 

and 39 reptile species from three orders and 17 

families. 

Inventory Completeness

We documented 7 species of amphibians and 

39 species of reptiles during this inventory 

(Appendix B). Based on the combined results 

of our inventory and other recent research and 

monitoring efforts for herpetofauna in the district, 

we believe that 9 species of amphibians and 

48 species of reptiles likely occur (Appendix 

B). Therefore, our inventory effort found 81% 

of the species present. Species accumulation 

curves (Fig. 4.3) nearly reached an asymptote for 

extensive and intensive surveys, suggesting that 

additional surveys would have produced few new 

species. In fact, many species that we found only 

incidentally or have been documented few times 

are so rare that encountering them is largely a 

function of chance. 

Species that we did not observe but that 

we believe are present include seven species 

confirmed by previous specimen vouchers 

and/or confirmed by park staff during the past 

decade and two other species believed to be 

present based on nearby specimen vouchers and 

unconfirmed observations. Species confirmed 

by park staff during the time of this study are the 

tiger salamander, American bullfrog, ornate box 

turtle, and Mediterranean house gecko. Of these, 

the non-native American bullfrog is certainly 

incidental; this species has been observed at 

the district in the past decade only during wet 

summers, and then only as dispersing juveniles 

that do not persist (Saguaro NP, unpubl. records). 

A population established in Wildhorse Canyon 

in the 1970s (Kevin Black, pers. comm.) has 

not been present for at least 15 years, possibly 

because of the park’s effort to eliminate it. Tiger 

salamanders are established in stock tanks in 

Reddington Pass (north of the district) and 

Danielle Foster observed one burrowed at the 

base of an exotic grass that she was pulling out 

near Rincon Creek in 2001. It is possible that 

this species breeds in the district, but is difficult 

to find because it spends little time above ground. 

The ornate box turtle is likely established in the 

district and staff found two individuals in 2005, 

though some individuals may be periodically 

released pets (P. Rosen, pers. comm.). The nonnative Mediterranean house gecko occurs in 

buildings in the Administration area only and 

there is no evidence that it is established in other 

areas. 

Based on nearby voucher specimens and 

unconfirmed observations, we believe that three 

other species of reptiles and amphibians occur 

in the district: long-nosed leopard lizard, glossy 

snake, and saddled leaf-nosed snake. A glossy 

snake was collected near the district entrance 

in 1967 and this species may occur along the 

district’s western boundaries. The long-nosed 

leopard lizard has been observed by park staff 

several times in areas such as the Javelina Picnic 

Area (Robert Ellis, pers. comm., Black 1982). It 

occurs on the Rocking K ranch adjacent to the 

district (Murray 1996) and probably occurs in 

the district as well. The checkered garter snake 

(Thamnophis marcianus), a riparian species, 

was reported for the district by Lowe and Holm 

(1991), but we could not find any current or 

historic records for this species. 

Discussion

Biogeography

The Rincon Mountains contain elements of 

several major biogeographic provinces, including 

the Sonoran Desert to the south and west, the 

Rocky Mountains to the north and east, and the 

Chihuahuan Desert and Madrean “sky islands” 

to the south and east (Shreve 1951, Brown 1994, 

Bowers and McLaughlin 1987). The large 
41

elevation range of the district allows it to contain 

many of the reptiles and amphibians associated 

with these very different ecological provinces. 

As a result, many representatives of each of the 

four major regions are present, including a large 

number of species not present in the Tucson 

Mountain District of the park. An interesting 

note is that a few low-desert Sonoran Desert 

species found in the Tucson Mountain District, 

such as the sidewinder (Crotalus cerastes) and 

desert iguana (Dipsosaurus dorsalis) reach 

the eastern edge of their ranges in the Tucson 

Mountains. Thus, the Rincon Mountain District’s 

herpetofauna contains classic Sonoran Desert 

species (e.g., desert tortoise), Rocky Mountain 

species (e.g., ring-necked snake), Chihuahuan 

Desert species (e.g., greater earless lizard), and 

Madrean species (e.g., Madrean alligator lizard). 

Many of these species are on the edge of their 

range in the district. A few taxa, including the 

southern plateau subspecies of the eastern fence 

lizard, occur in the Rincon and nearby Santa 

Catalina Mountains as disjunct populations 

(Stebbins 2003).

There are also a large number of species 

that occur close to the Rincon Mountains but 

that have not been observed in the district. Our 

inventory confirms regional distribution patterns 

of herpetofauna first described by Lowe (1994) 

who noted that many Madrean species reach 

their northern limits along what he referred to as 

the “Madrean Line” that corresponds roughly to 

Interstate-10, which runs just to the south of the 

district (See Fig. 2.1). Lowe (1994) focused on 

several Madrean rattlesnakes that are found in 

the Santa Rita Mountains but not in the Rincon 

or Santa Catalina mountains, including the twinspotted rattlesnake (Crotalus pricei), banded 

rock rattlesnake (C. lepidus), and Arizona ridgenosed rattlesnake (C. willardi). By contrast, the 

western rattlesnake, a “Rocky Mountain” species, 

is found in the Rincon Mountains but not in the 

Santa Rita Mountains. Lowe’s observation has 

been confirmed by biogeographical analyses of 

recent inventories (Swann et al. 2005), including 

ours. Rumors have long persisted that some of 

these Madrean species (especially banded rock 

rattlesnakes) occur in the Rincon Mountains, but 

this inventory provides further evidence that they 

do not. 

Other species found near Tucson that 

do not occur in the district include many mesic 

riparian species, including the Mexican garter 

snake (Thamnophis eques) and Woodhouse toad 

(Bufo woodhousii). The Texas horned lizard 

(Phyrnosoma cornutum), a Chihuahuan Desert 

species, has been found in Mescal (20 km SE 

of the district; Roger Repp, pers. comm.) but is 

unlikely to occur in the district. 

Abundance and Distribution

The Rincon Mountain District has a well-studied 

herpetofauna compared to other areas, due 

mainly to its proximity to Tucson. In particular, 

recent field studies of individual species have 

facilitated incidental observations of reptiles and 

amphibians that are not often seen. In addition, 

the size of the staff at Saguaro in comparison with 

smaller NPS units in the Sonoran Desert Network 

has resulted in better documentation of sightings, 

including collection of roadkill. On the other 

hand, the district is large, mostly roadless, and 

topographically complex, which makes studies 

there difficult. 

Our study is the first to quantify relative 

abundance and distribution of amphibians and 

reptiles in the district and to evaluate patterns of 

these parameters in space and time. Many of the 

patterns that we documented confirm patterns 

observed in previous studies. However, the 

greater rate of detections per hour on extensive 

(4.1 detections/hr) vs. intensive (3.6 detections/

hr) surveys was dramatically different than in 

the Tucson Mountain District, where extensive 

surveys (4.5 detections/hr) produced far fewer 

detections than intensive surveys (6.3 detections/

hr) (Flesch et al. 2006). Tables 4.5 and 4.8 

suggest that this may be due to the effect of 

greater numbers of intensive surveys at higher 

elevations, where detection rates were lower than 

on low-elevation plots. 

In general, both abundance and 

distribution of reptiles and amphibians decreased 

with increasing elevation in the district. This 

pattern is well-known and certainly corresponds 

to declining species richness of reptiles (but 

not amphibians) across an increasing latitudinal 
42

gradient (Stein 2002), and is undoubtedly related 

to the physiology of these taxa. 

The far greater number of diurnal lizards 

detected on both intensive and extensive surveys 

compared to snakes and amphibians reflects the 

diurnal abundance of lizards. Snakes can be 

both diurnal and nocturnal, but are nearly always 

observed less frequently than lizards during 

species inventories in the southwestern United 

States (e.g., Turner et al. 2003, Swann et al. 

2000, Swann and Schwalbe 2001, Powell et al. 

2005). Excluding diurnal frogs in riparian areas, 

most amphibians we observed were toads, which 

are active almost exclusively at night during the 

summer rainy season – clearly evidenced by the 

large increase in the number of toads we detected 

with rising humidity. In contrast, lizard activity 

declines with increasing humidity and cloud 

cover, which is consistent with our observations. 

Study Design

Our major goals for this inventory were to apply 

a repeatable study design that (in some cases) 

allowed inference to the whole district and also 

to detect the maximum number of species per 

unit time of field effort. In general we achieved 

these goals, but clearly some methods were more 

effective than others. 

Intensive surveys were not highly 

successful at the district, in part because of its 

large size and environmental heterogeneity. 

Intensive surveys had relatively low observation 

rates and poor species detections. For 

consistency with other inventories, we stratified 

our study plots based only on elevation, but 

species richness, abundance, and distribution of 

reptiles and amphibians are clearly based on key 

habitat features such as slope, aspect, geology, 

and presence of water. In recognition of this, 

we revised our strategy in 2002, increased the 

number of extensive surveys at lower elevations, 

and were more successful in detecting rare 

species. If intensive surveys are included in 

future species richness monitoring at the district, 

we would recommend a stratification approach 

that includes wet riparian areas. 

Extensive surveys detected many species 

(n = 39), in part because more time was spent 

using this method and areas were surveyed 

in both day and night. However, this method 

did not detect as many species as incidental 

observations (n = 43). As in many previous 

herpetological inventories (see Swann 1999a), 

these results indicate how difficult it is to detect 

many reptile and amphibian species, which tend 

to be rare, extremely cryptic, subterranean in 

their habits, or a combination of these factors. 

Our study confirms that, at least until better 

technology is available for detecting rare species, 

a combination of methods, including incidental 

sightings and collection of roadkilled animals, is 

essential to achieve a complete list of species. 

Management Issues 

We did not observe any federally threatened 

or endangered species. The Sonoran Desert 

population of the desert tortoise is a species 

of conservation concern (Appendix B) and 

has been petitioned for federal listing. This 

species is abundant in and around the district 

(Swann et al. 2002), and the park has both a past 

inventory (Wirt and Robichaux 2000) and current 

monitoring plans for this species. Exotic diseases 

in tortoises, particularly upper respiratory tract 

disease (Jones et al. 2005), is a concern and 

monitoring the health of this species should 

occur periodically. The canyon spotted whiptail 

is another species of conservation concern 

(Appendix B). The only known population in the 

district occurs at Madrona ranger station (Bonine 

and Schwalbe 2003). 

The lowland leopard frog is probably the 

most threatened species of herpetofauna in the 

district, as the park has long recognized (Swann 

1997). Lowland leopard frogs seem to have 

declined in southern Arizona and are extirpated in 

parts of their former range although populations 

in central Arizona seemed to be stable when 

last reported (Clarkson and Rorabaugh 1989, 

Sredl et al. 1997). In addition to habitat loss, a 

major threat to this species is the fungal disease 

chytridiomycosis, an introduced, potentially 

pandemic disease that occurs in the district (D. 

Swann, unpubl. data). The district has several 

small populations of lowland leopard frogs, yet 

at least one major population was extirpated 
43

in recent years due to sedimentation of pools 

following major wildland fires (Swann et al. 

2003). Most other populations seem to be stable, 

yet their small size and isolation may be factors 

that, when combined with stochastic events, may 

threaten their long-term persistence. 

We suspect that the district has a 

relatively stable herpetofauna community. There 

is little evidence that non-native species (reptiles, 

amphibians, mammals, or birds) are having an 

impact on reptile and amphibian populations. 

For example, if Mediterranean geckos were 

capable of establishing themselves in the district, 

they probably would have already done so. 

The greatest threat to herpetofauna from exotic 

species is probably from crayfish and American 

bullfrogs. Crayfish could have a dramatic 

negative impact on populations of lowland 

leopard frogs, canyon treefrogs, Sonoran mud 

turtles, and black-necked garter snakes if they 

were illegally introduced into the Rincon Creek 

watershed. Park personnel should be vigilant to 

prevent establishment.

Reptile poaching may occur in the 

park, but is probably confined to areas along the 

western edge of the district. We suspect that 

individual Sonoran desert toads (Bufo alvarius), 

a species that is traded and used because it 

possesses hallucinogenic qualities, are sometimes 

collected in the district. Roadkill has been welldocumented at the park; park staff estimate that 

literally thousands of reptiles and amphibians 

are killed by cars each year (Kline and Swann 

1998). Species most impacted by roadkill tend to 

be long-lived species such as the desert tortoise 

and Sonoran Desert toad. However, the problem 

is likely more severe in the Tucson Mountain 

District, which is more bisected by roads. 

Finally, habitat loss and fragmentation 

outside the district are major threats to all wildlife 

at Saguaro National Park, although likely a 

greater threat for mammals than for herpetofauna 

(see Chapter 6). The major species impacted 

by habitat loss are desert species with limited 

habitat in the park. These include the lowland 

leopard frog, Mexican spadefoot toad (Spea 

multiplicada), Great Plains toad, canyon whiptail, 

long-nosed leopard lizard, glossy snake, and Pima 

(saddled) leaf-nosed snake. If any herpetological 

species is extirpated from the district in the next 

few decades, we predict it will be a species with 

more specialized habitat requirements, such as 

the canyon spotted whiptail or lowland leopard 

frog. 
44
45

Chapter 5: Bird Inventory

Brian F. Powell 

Previous Research

There has been considerable bird research at the 

Rincon Mountain District, but no comprehensive 

and well-documented inventory has been 

completed. Monson and Smith (1985) compiled 

a checklist for both districts of the park, but there 

is no documentation of the data used to create 

that list. The list includes abundance categories 

for each major vegetation community and this 

information was likely based on Gale Monson’s 

extensive knowledge of the distribution and 

relative abundance of birds in similar vegetation 

communities in the region. 

A few studies have investigated 

songbird community composition in the Sonoran 

desertscrub on the west side of the district near 

the Cactus Forest Loop Drive (Johnson and 

Haight 1991, see also Mannan and Bibles 1989) 

and in the Rincon Valley (Boal and Mannan 1996, 

Freiderici 1998, Powell 1999, 2004). Only two 

multi-species, non-raptor studies have taken place 

in the higher elevations of the district (Marshall 

1956, Short 2002) and no research has taken 

place in the mid-elevation areas of the district or 

on the east slope of the Rincon Mountains. In the 

1980s the park was concerned about the impact of 

non-native cavity-nesting birds on native species 

and they commissioned studies to investigate 

this (Mannan and Bibles 1989, Kerpez and Smith 

1990). Because of the active fire management 

program, park personnel have been interested 

in the effects of fire on the Mexican spotted owl 

(Willey 1998) and songbirds (Short 2002) in the 

high elevation areas of the district. The park 

contracted for periodic raptor surveys (Felley and 

Corman 1993, Berner and Mannan 1992, Bailey 

1994, Griscom 2000). Park personnel surveyed 

three Breeding Bird Atlas blocks within the 

district (Short 1996) and those results are reported 

in Corman and Wise Gervais (2005). The Tucson 

Bird Count includes three low-elevation sites in 

the park, including Rincon Creek (TBC 2005). 

Single species studies have included the elf owl 

(Goad and Mannan 1987, Steidl 2003), Mexican 

spotted owl (Willey 1997, 1998b, Anderson 

and Schon 1999, Steidl and Knipps 1999), buffbreasted flycatcher (Conway and Kirkpatrick 

2001; they also noted other species; Kirkpatrick 

et al. 2006), and purple martin (Stutchbury 1991). 

Park personnel survey periodically for the cactus 

ferruginous pygmy-owl and Mexican spotted 

owl and park staff file annual reports to the U.S. 

Fish and Wildlife Service (Saguaro NP, unpubl. 

reports) on monitoring and relevant management 

activities related to these species.

Methods

We surveyed for birds at the Rincon Mountain 

District from 2001 to 2003, though most of our 

surveys took place in the springs of 2001 and 

2002. We used four field methods: (1) variable 

circular-plot (VCP) counts for diurnal breeding 

and spring migrant birds, (2) nocturnal surveys 

for owls and nightjars (breeding season), (3) line 

transects for diurnal birds in the non-breeding 

season, and (4) incidental observations for all 

birds in all seasons. Although line-transect 

surveys were not included in the original study 

proposal (Davis and Halvorson 2000), we felt 

they were important in our effort to inventory 

birds at the district because many species that 

use the area during the fall and winter may not 

be present during breeding-season surveys. 

Nevertheless, we concentrated our primary 

survey effort on the breeding season because bird 

distribution is relatively uniform in that season 

due to territoriality (Bibby et al. 2002). Our 

survey period included peak spring migration 

times for most species, which added many 

migratory birds to our list.

We sampled vegetation around most 

VCP stations. Vegetation structure and plant 

species composition are important predictors of 

bird species richness or the presence of particular 

species (MacArthur and MacArthur 1961, Rice et 

al. 1984, Strong and Bock 1990). In this report 

we use these data to categorize and describe bird 

communities. These data will also be useful for 

habitat association studies (e.g., Strong and Bock 

1990; see Appendix H for results). 
46

Spatial Sampling Designs

We established study sites based on random 

and non-random criteria. We surveyed at 17 

randomly located focal-point transects (Fig. 5.1; 

see Chapter 1 for additional information). We 

established the locations of all other surveys in 

areas that we believed would have the highest 

species richness or as a matter of convenience 

(Figs. 5.1, 5.2). For all survey methods, we 

collected data at individual stations or sections, 

which we grouped into transects because of 

convenience and efficiency. (An alternative 

approach would have been to establish 

individual stations or sections to maintain greater 

independence, but travel time between stations 

would have reduced the number of stations that 

we were able to visit in a morning.) We placed 

our non-random transects along riparian areas and 

canyons in low-elevation areas (< 4000 ft); in all 

other areas we established non-random transects 

along trails because of accessibility and safety.

VCP Surveys 

We used the variable circular-plot method (VCP; 

Reynolds et al. 1980, Buckland et al. 2001) 

to survey for diurnally active birds during the 

breeding and spring migration seasons (mid April 

through early July). Conceptually, these surveys 

are similar to traditional “point counts” (Ralph 

et. al 1995) during which an observer spends a 

standardized length of time at one location (i.e., 

station) and records all birds seen or heard and 

the distance to each bird or group of birds.

We used three types of VCP surveys 

(Table 5.1). Methods differed primarily by 

the sampling design used to establish their 

location and by the number of visits (see Table 

5.1 for additional information). The following 

description of our survey protocol applies to all 

VCP methods unless otherwise noted. We located 

stations a minimum of 250 m apart to maintain 

independence among observations. On each 

successive visit to a transect we alternated the 

order in which we surveyed stations to minimize 

bias by time of day or direction of travel. We did 

not survey when wind exceeded 15 km/h or when 

precipitation exceeded an intermittent drizzle. 

We attempted to begin surveys approximately 30 

minutes before sunrise and conclude surveys no 

later than three hours after sunrise. 

We recorded a number of environmental 

variables at the beginning of each transect: wind 

speed (Beaufort scale), presence and severity of 

rain (qualitative assessment), air temperature (oF), 

relative humidity (%), and cloud cover (%). After 

arriving at a station, we waited one minute before 

beginning the count to allow birds to resume 

their normal activities. We identified to species 

all birds seen or heard during an eight-minute 

“active” period (5 minutes at reconnaissance 

VCP stations). For each detection we recorded 

the distance (in meters) the bird was from the 

observer (measured with laser range finder 

when possible), time of detection (measured in 

Table 5.1. Characteristics of the three major VCP survey types for birds, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. 

VCP survey type

Repeat-visit

Characteristic Random (focal-point transects) Non-random Reconnaissance 

Randomly located Yes No No

Number of visits per year 4 >2 1

Number of stations 4 variable variable

Count duration at each station 8 minutes 8 minutes 5 minutes

Advantages Scope of inference to larger area, 

vegetation data available 

Flexible, most complete abundance 

data for areas with high species 

richness, uncommon and rare species 

are often accounted for

Maximum flexibility, allows for 

rapid inventories and larger spatial 

coverage, provides good distribution 

data 

Disadvantages Inefficient for developing 

complete species list, transects 

are often in areas of low species 

richness

No spatial inference beyond transect Species lists are less complete, 

because uncommon and rare 

species may be missed
47

Figure 5.1. Locations of VCP survey stations (random [focal-point transects], non-random, 

and reconnaissance), Saguaro National Park, Rincon Mountain District, 2001 and 2002. 
48

one-minute intervals beginning at the start of 

the active period), and the sex and/or age class 

(adult or juvenile), if known. We did not measure 

distances to birds that were flying overhead 

nor did we use techniques to attract birds (e.g., 

“pishing”). We made an effort to avoid doublecounting individuals. If we observed a species 

during the “passive” count period (between 

the eight-minute counts), which had not been 

recorded previously at a station on that visit, we 

recorded its distance to the nearest station. 

Effort

In 2001, we spent more effort surveying at 

focal-point stations (n = 272) than at nonrandom stations (n = 160; Table 5.2). In 2002 

we surveyed exclusively at non-random stations, 

both repeat-visit (n = 130) and reconnaissance (n

= 107). In both years the number of stations and 

visits varied among transects except for random 

transects, which had four stations that we visited 

four times in 2001 (Table 5.2). 

Analysis

Relative Abundance. We calculated relative 

abundance of each species along each transect as 

the number of detections at all stations and visits 

(including zero values) and divided by effort 

(total number of visits multiplied by total number 

of stations). We reduced our full collection of 

Figure 5.2. Location of section breaks for non-breeding season (winter) bird transects and nocturnal 

survey stations, Saguaro National Park, Rincon Mountain District, 2001 and 2002. 
49

observations for each repeat-visit VCP station 

to a subset of data that was more appropriate for 

estimating relative abundance. We used only 

those detections that occurred ≤ 75 m from count 

stations because detectability is influenced by 

conspicuousness of birds (i.e., loud, large, or 

colorful species are more detectable than others) 

and environmental conditions (dense vegetation 

can reduce likelihood of some detections). 

Truncating detections may reduce the influence 

of these factors (Verner and Ritter 1983; for a 

review of factors influencing detectability see 

Anderson 2001a, Pollock et al. 2002). We also 

excluded observations of birds that were flying 

over the station, birds observed outside of the 

eight-minute count period, and unknown species. 

Some observations met more than one of these 

criteria for exclusion from analysis. We report 

Table 5.2. Summary of bird survey effort, Saguaro National Park, Rincon Mountain District, 2001–2003. 

Sample size (n) was used in calculating relative abundance for each transect and year. 

Years(s)

Random or 

Non-random

Number of 

stations 

2001 2002/2003 

Survey type Community type Transect name(s) Visits n Visits n

Repeat-visit VCP Randomb Low Sonoran Desertscrub 112, 115, 130, 138, 139 20 4 80

Middle Sonoran Desertscrub 121 4 4 16

Oak Savannah 101, 106, 111, 189 16 4 64

Pine-oak Woodland 120, 125 8 4 32

High Pine-oak Woodland 107, 128, 155 12 4 48

Conifer Forest 113, 191 8 4 32

Non-random Riparian Lower Rincon Creek 8 4 32 7 55

Upper Rincon Creek 4 4 16 5 35

Box Canyon 7 4 28 5 35

Upper Loma Verde Wash 2-5 4 8 3 15

Pine-oak Woodland Happy Valley Saddle 6 4 24 2 12

Conifer Forest Rincon Peak 4 4 16 2 8

Reconnaissance 

VCP Non-random Low Bridal Wreath Falls 7 1 7

Broadway Trailhead 8 1 8

Chimenea Creek 6 1 6

Freeman Road 8 1 8

Freeman Wash 4 1 4

Loma Verde Wash 8 1 8

Madrona Canyon 6 1 6

Middle Douglas Spring 4-8 2 12

Juniper Spring 5 1 5

High Deer Head Spring 5 1 5

Italian Spring 8 1 8

Manning Cabin 8 1 8

Mica Mountain 5 1 5

Mica Mountain Trail 4 1 4

North Slope Trail 8 1 8

Upper Juniper Basin 5 1 5

Line transect Non-random Douglas Spring 12 4 48

Lower Rincon Creek 13 4 52

Upper Loma Verde 8 4 36

Nocturnal Non-random Low Box Canyon 2 1 2

Loop Drive 6 4 24 1 6

Rincon Creek 4-6 3 18 1 4

Middle Cowhead Saddle 4 2 8

Juniper Basin 3 1 3

High Happy Valley 3 3 9

 Italian Spring 2 1 2

Manning 4 3 12 1 4

Spud Rock Spring 4 1 4

a

 Low = <4,000 feet; Middle = 4,000–6,000 feet; High = >6,000 feet. 

b

 All transects had four stations and were surveyed four times. 
50

the relative abundance by repeat-visit transect and 

year. Because relative abundance is the closest 

index to true population size that we employ (see 

Chapter 1 for more detailed discussion), we use it 

to note the “abundance” of species. 

Relative Frequency of Detection. Relative 

abundance is the least biased index to abundance 

because we control a number of variables that 

account for differences among transects (e.g., 

the ability to see or hear a bird). However, 

we also wanted an index that accounted for all 

of the species observed within a transect (i.e., 

including birds seen > 75 m, flyovers, and birds 

seen outside of the 8-minute count period) and 

which also conveyed some relative abundance 

information. Relative frequency of detections 

incorporates these observations and differs from 

relative abundance in that it is clearly biased 

toward those species that are highly visible or 

vocal. Therefore, it can be thought of as an index 

of the number of birds that we saw and heard at 

typical stations on the transect (i.e., most similar 

to an observer’s “experience”). This method also 

enables us to convey other important information 

regarding species’ presence at the transect level. 

Community Classification. Using data from 

repeat-visit VCP transects, we sought to identify 

bird communities within the district and to 

compare bird characteristics among communities. 

We did not use the original stratification of 

random transects for this analysis because we 

were more interested in identifying communities 

than drawing inference to a larger area. To group 

transects, we used Ward’s hierarchical cluster 

analysis using bird and vegetation data. Cluster 

analysis is a multivariate technique that groups 

like entities (in our case transects) that share 

similar values. We performed separate cluster 

analyses for the bird and vegetation data (see 

Chapter 3 for results of cluster analysis using 

vegetation data). To identify groups from the 

bird data we used mean relative abundance for 

each transect and all visits in both years. We 

attempted to include reconnaissance VCP surveys 

into this analysis but the results were inconsistent, 

most likely because we had no vegetation data for 

these transects and there was likely insufficient 

sampling effort for birds. 

Comparing Communities. We compared 

species richness and relative abundance among 

community types. To compare species richness 

we used a subset of data from all transects so 

that each transect consisted of four visits to 

four stations (n = 16); the minimum number 

of visits and stations to repeated-visit transects 

(see Table 5.2). We used only those detections 

< 75 m from stations (n = 2,476 observations) 

and excluded flyovers and birds seen outside 

of the eight-minute count period. To compare 

relative abundance among communities, we 

used observations from all visits and stations 

and did not choose a subset of observations (as 

for species richness) because relative abundance 

is scaled by survey effort. We tested for 

differences among all communities using oneway analysis of variance (ANOVA) and searched 

for pairwise differences between communities 

using the Tukey-Kramer procedure. We logtransformed relative abundance data to better 

meet assumptions of normality. 

Line-transect Surveys 

Field Methods

We used a modified line-transect method (Bibby 

et al. 2002) to survey for birds from November 

2002 to February 2003. Line transects differ 

from VCP transects in that an observer records 

birds seen or heard while the observer is walking 

an envisioned line rather than while standing at 

a series of stations. The line-transect method is 

more effective during the non-breeding season 

because bird vocalizations are less conspicuous 

and frequent and therefore birds tend to be more 

difficult to detect aurally (Bibby et al. 2002). 

We established three transects in the 

district (Fig. 5.2). Transects were broken into 

sections, each approximately 250 m in length. As 

with VCP transects, we alternated direction of 

travel to reduce biases and did not survey during 

periods of excessive rain or wind (see VCP 

survey methods for details). We began surveys 

about 30 minutes after sunrise and continued until 

we completed the transect. As with VCP surveys, 

we recorded weather conditions at the beginning 

and end of each survey. We timed our travel so 

that we traversed each section in ten minutes, 
51

during which time we assigned all birds seen 

and/or heard into one of the following distance 

categories: ≤ 100 m, > 100 m, or “flyover.” 

When possible, we noted the sex and age class of 

birds. We recorded birds observed before or after 

surveys as “incidentals” (see section below) and 

we did not use techniques to attract birds (e.g., 

“pishing”).

Effort

The number of sections along each transect 

ranged from eight to 13 (Table 5.2). We surveyed 

each transect four times in the winter of 2002 and 

2003. 

Analysis

Due to the low number of observations 

within 100 m of the transect lines, we used 

all observations (except unknown species) to 

estimate relative abundance (see Methods section 

of VCP surveys for more details). 

Nocturnal Surveys

Field Methods

To survey for owls we broadcast commercially 

available vocalizations (Colver et al. 1999) using 

a compact disc player and broadcaster (Bibby et 

al. 2002) and recorded other nocturnal species 

(nighthawks and poorwills) when observed. We 

established nine transects (Fig. 5.2). The number 

of transects per elevation stratum was lowest 

for the middle elevation (n = 2) and highest for 

the high elevation areas (n = 4) (Table 5.2). The 

number of stations per transect varied depending 

on logistical constraints but all stations were a 

minimum of 500 m apart. For transects that we 

visited multiple times, we attempted to reduce 

sampling biases by varying direction of travel 

along transects. We began surveys approximately 

45 minutes after sunset. 

We began surveys at each station with a 

three-minute “passive” listening period during 

which time we broadcast no calls. We then 

broadcast vocalizations for a series of two-minute 

“active” periods. We broadcast vocalizations 

of species that we suspected might be present, 

based on habitat and range information. The 

species that we broadcasted changed based on the 

elevation stratum of the surveys:

•	 Low elevation: elf, western screech, 

burrowing, and barn owls;

•	 Middle elevation: elf, northern pygmy, 

flammulated, and whiskered screech 

owls;

•	 High elevation: northern pygmy, 

flammulated, northern saw-whet, and 

whiskered screech owls. 

We excluded the great horned owl from the 

broadcast sequence because of its aggressive 

behavior toward other owls (though we recorded 

them incidentally). Also, we did not survey for 

the Mexican spotted owl or the cactus ferruginous 

pygmy-owl because that would have required 

use of specific protocols and because park staff 

survey periodically for these species. 

We broadcast recordings of owls in 

sequence of species size, from smallest to largest, 

so that smaller species would not be inhibited by 

the “presence” of larger predators or competitors 

(Fuller and Mosher 1987). During active periods, 

we broadcast owl vocalizations for 30 seconds 

followed by a 30-second listening period. This 

pattern was repeated two times for each species. 

During the count period we used a flashlight to 

scan nearby vegetation and structures for visual 

detections. If we observed a bird during the 

three-minute passive period, we recorded the 

minute of the passive period in which the bird 

was first observed, the type of detection (aural, 

visual or both), and the distance to the bird. If a 

bird was observed during any of the two-minute 

active periods, we recorded in which interval(s) 

it was detected and the type of detection 

(aural, visual, or both). As with other survey 

methods, we attempted to avoid double-counting 

individuals recorded at previous stations. We 

also attempted to use a different observer for each 

visit, alternate direction of travel along transects, 

and not survey during inclement weather. 

Effort

The number of stations and visits to each transect 

differed among transects (Table 5.2). Although 

we had the most transects in the high elevation 

stratum, we had most (56%) of our survey effort 

in the low elevation stratum because of greater 

ease of accessing stations.
52

Analysis

We report relative abundance as the mean number 

of observations. 

Incidental and Breeding Observations 

Field Methods

When we were not conducting formal surveys 

and we encountered a rare species, a species in 

an unusual location, or an individual engaged 

in breeding behavior, we recorded UTM 

coordinates, time of detection, and (if known) 

the sex and age class of the bird. We recorded 

all breeding observations using the standardized 

classification system developed by the North 

American Ornithological Atlas Committee 

(NAOAC 1990), which characterizes breeding 

behavior into one of nine categories: nest 

building, occupied nest, used nest, adult carrying 

nesting material, adult carrying food or fecal sac, 

adult feeding young, adult performing distraction 

display, or fledged young. We made breeding 

observations during standardized surveys and 

incidental observations. 

Analysis

We report frequency counts of incidental and 

breeding observations.

Vegetation Sampling at Non-random VCP Stations

Field Methods

We quantified vegetation characteristics along 

random transects (see Chapter 3 for details). In 

2002 we sampled vegetation associated with 

each of the repeat-visit, non-random transects. 

At each station we sampled vegetation at five 

subplots located at a modified random direction 

and distance. Each plot was located within a 

72° range of the compass from the station (e.g., 

Plot 3 was located between 145° and 216°) to 

reduce clustering of plots. We randomly placed 

plots within 75 m of the stations to correspond 

with truncation of data used in estimating relative 

abundance. 

At each plot we used the point-quarter 

method (Krebs 1998) to sample vegetation 

by dividing the plot into four quadrants along 

cardinal directions. We applied this method to 

plants in one size category: potential cavitybearing vegetation (> 20 cm diameter at breast 

height), and three height categories: sub-shrubs 

(0.5–1.0 m), shrubs (> 1.0–2.0 m), trees (> 

2.0 m). If there was no vegetation for a given 

category within 25 m of the plot center, we 

indicated this in the species column. For each 

individual plant, we recorded distance from the 

plot center, species, height, and maximum canopy 

diameter (including errant branches). Association 

of a plant to a quadrant was determined by the 

location of its trunk, regardless of which quadrant 

the majority of the plant was in; no plant was 

recorded in more than one quadrant. Standing 

dead vegetation was recorded only in the 

“potential cavity-bearing tree” category. On rare 

occasions when plots overlapped we repeated the 

selection process for the second plot. 

Within a 5-m radius around the center of 

each plot, we visually estimated percent ground 

cover by type (bare ground, litter, or rock); 

and percent aerial cover of vegetation in each 

quadrant using three height categories: 0–0.5 m, 

> 0.5–2.0 m, and > 2.0 m. For both estimates we 

used one of six categories for percent cover: 0 

(0%), 10 (1–20%), 30 (21–40%), 50 (41–60%), 

70 (61–80%), and 90 (81–100%). 

Analysis

Using point-quarter data, we calculated mean 

density (number of stems/ha) for all species 

in each of the four height/size categories. We 

used the computer program Krebs to calculate 

density (Krebs 1998). We collected these data 

to characterize gross vegetation characteristics 

around survey stations. 

Results

We made over 15,000 observations of birds and 

found 173 species from 2001 to 2003 (Appendix 

C). We found 10 species that had not previously 

been found in the district including the sulphurbellied flycatcher, elegant trogon, and pinyon 

jay. Among the 173 species that we observed, 

there were a number with special conservation 

designations including the northern goshawk, 

yellow-billed cuckoo, Mexican spotted owl, 
53

and buff-breasted flycatcher. Unusual sightings 

included a nest of the sulphur-bellied flycatcher, 

a singing male buff-breasted flycatcher, and 

sightings of the wild turkey, common black hawk, 

and yellow-breasted chat. We recorded three 

non-native species, including the rock pigeon, 

a new species for the district. We recorded the 

most species during incidental observations (n

= 154) and VCP surveys (n = 149) and fewest 

during nocturnal surveys (n = 9). 

Community Types

We performed cluster analysis on bird and 

vegetation data and found almost complete 

congruency of results for the random transects 

(we did not include vegetation data from nonrandom transects into the cluster analysis for 

plants; see above). Interpreting data from both 

analyses, we found there to be five communities 

(Fig. 5.3). Based on the bird data, we grouped 

the Happy Valley Saddle transect differently than 

we expected; it was originally classified as Oak 

Savannah, but we assigned it to the Pine-oak 

Woodland vegetation community.

•	 Riparian. All low-elevation non-random 

transects (Lower and Upper Rincon 

Creek, Box Canyon, and Loma Verde 

Wash). Creeks and washes lined by thick 

vegetation such as Fremont cottonwood, 

Arizona sycamore, and willow (except 

Loma Verde Wash), velvet ash, and 

bordered by Sonoran Desertscrub.

•	 Sonoran Desertscrub. Five lowelevation random transects (112, 115, 

130, 138, and 139) and one middle 

elevation transect (121). Mixed cactus, 

succulents, and palo verde, with some 

velvet mesquite, especially in the dry 

washes.

•	 Oak Savannah. Four middle-elevation 

random transects (101, 106, 189, and 

111). Open areas dominated by perennial 

grasses with scattered trees, mostly oaks.

•	 Pine-Oak Woodland. Two middle- (125 

and 120) and three high- (107, 155, and 

128) elevation random transects and 

one non-random transect (Happy Valley 

Saddle). Most transects had dense stands 

of manzanita and oaks, interspersed 

101

106

111

189

Happy Valley Saddle

112

115

121

130

138

139

113

191

Rincon Peak

Box Canyon

Lower Rincon Creek

Upper Loma Verde Wash

Upper Rincon Creek

107

120

125

128

155

Oak Savannah 

Riparian 

Sonoran Desertscrub 

Conifer Forest 

Pine/oak Woodland 

Figure 5.3. Dendrogram of bird community groups from Ward’s hierarchical cluster analysis, Saguaro 

National Park, Rincon Mountain District, 2001 and 2002. See text for descriptions of bird communities 

and data used in analysis. 
54

an average of over one individual per station. 

The mourning dove (0.98 + 0.42) and ashthroated flycatcher (0.85 + 0.24) were the only 

other species with relative frequency of detection 

estimates > 0.75.

There were differences in mean relative 

abundance estimates among transects (F4, 263

= 4.2, P = 0.003, ANOVA on log-transformed 

data). Specifically, the Conifer Forest community 

was different from both the Riparian and Pineoak Woodland communities (Table 5.3). Mean 

species richness per visit also varied among 

communities (Table 5.3; F4, 111 = 6.7, P = < 0.001, 

ANOVA). The Riparian community had the most 

species per visit and was significantly different 

from all communities except the Conifer Forest 

community. 

We calculated relative abundance for 120 

species (Table 5.4). The most abundant species 

(based on relative abundance estimates) for each 

community type were:

•	 Riparian: verdin, Lucy’s warbler, and 

mourning dove;

•	 Sonoran Desertscrub: black-throated 

sparrow, cactus wren, and verdin; 

•	 Oak Savannah: Bewick’s wren, rufouscrowned sparrow, and ash-throated 

flycatcher;

•	 Pine-oak Woodland: Bewick’s wren, 

spotted towhee, and black-throated gray 

warbler;

•	 Conifer Forest: yellow-eyed junco, 

mountain chickadee, and spotted towhee 

and cordilleran flycatcher. 

Reconnaissance VCP Surveys

We recorded 75 species during reconnaissance 

VCP surveys in 2002, including two species 

that we did not record during repeat-visit VCP 

with some pine trees, mostly pinyon and 

ponderosa pine (Happy Valley Saddle).

•	 Conifer Forest. Two high-elevation 

random transects (113 and 191) and one 

non-random transect (Rincon Peak). 

Forests of ponderosa pine, Douglas-fir, 

and some Gambel oak. 

Repeat-visit VCP Surveys

We recorded 143 species at all repeat-visit VCP 

stations combined. We found the most species 

in the Riparian community (n = 102 species) and 

fewest species in the Conifer Forest community 

(n = 51; Appendix G), though survey effort 

among communities was unequal (Table 5.2). 

The number of species found in the other three 

communities was intermediate (Appendix G). As 

expected, estimates of species richness (using the 

1st order jackknife procedure) followed the same 

pattern: the Riparian community was the most 

species rich (n = 119 species) and the Conifer 

Forest was the least species rich (n = 69). The 

Sonoran Desertscrub (n = 97 species), Pine-oak 

Woodland (n = 93 species), and Oak Savannah (n

= 79) were intermediate.

We recorded twelve species in all five 

communities and 39 species in only a single 

community (Appendix G). The ash-throated 

flycatcher was the most widespread species; we 

recorded it on 93% (21 of 23) of repeat-visit 

transects. We recorded four other species at 

>75% of transects: rufous-crowned sparrow, 

common raven, brown-headed cowbird, and 

white-winged dove. We recorded an additional 

22 species on >50% of transects and an equal 

number of species on only a single transect. 

The white-winged dove had the highest mean 

frequency of detection (1.25 + 0.44) across strata 

and it was the only species for which we recorded 

Table 5.3. Bird measures by community type and compared using Tukey-Kramer multiple pairwise 

procedure, Saguaro National Park, Rincon Mountain District, 2001 and 2002. Communities with different 

superscripted letter(s) are significantly different (P < 0.05).

Riparian Sonoran Desertscrub Oak Savannah Pine-oak Woodland Conifer Forest

Bird measure Mean SE Mean SE Mean SE Mean SE Mean SE

Relative abundance (log transformed) -3.4a 0.2 -3.1a 0.2 -2.7b 0.2 -3.2 a 0.2 -2.2b 0.2

Species richnessa 25.9 a 0.7 22.1b 0.8 21.2b 1.1 20.6b 0.9 22.1 b 1.2

a

 From 1st order jackknife procedure.
55

Table 5.4. Relative abundance (mean + SD) by community type for birds recorded during repeat-visit VCP 

surveys, Saguaro National Park, Rincon Mountain District, 2001 and 2002. Relative abundance estimates 

exclude flyovers and birds observed >75m from stations. Coefficient of variation (CV) is SD divided by the mean; 

low CV indicates less within-community variability of relative abundance. 

Riparian 

(n = 4)

Sonoran Desertscrub 

(n = 6)

Oak 

Savannah (n = 4)

Pine-oak 

Woodland (n = 6)

Conifer Forest 

(n = 3)

Species Mean SD CV Mean SD CV Mean SD CV Mean SD CV Mean SD CV

Gambel’s quail 0.50 0.24 0.5 0.06 0.10 1.6 0.14 0.24 1.7

Montezuma quail 0.01 0.02 2.4 0.02 0.03 2.0 0.01 0.02 2.4

turkey vulture 0.01 0.02 2.0

Cooper’s hawk 0.02 0.01 0.7

northern goshawk 0.01 0.02 2.4

gray hawk <0.01 0.01 2.0

red-tailed hawk 0.01 0.01 2.0

band-tailed pigeon 0.03 0.05 1.7

white-winged dove 0.79 0.36 0.5 0.40 0.29 0.7 0.35 0.42 1.2 0.04 0.07 1.5

mourning dove 0.87 0.30 0.3 0.49 0.10 0.2 0.16 0.20 1.2 0.05 0.08 1.6

common ground-dove 0.01 0.02 1.2

greater roadrunner 0.02 0.05 2.0 0.01 0.02 2.4

great horned owl 0.05 0.06 1.3

broad-billed hummingbird 0.06 0.06 1.0

black-chinned hummingbird 0.16 0.12 0.8 0.03 0.05 1.7 0.03 0.03 1.2 0.03 0.03 1.1

Anna’s hummingbird 0.02 0.03 1.5 0.03 0.07 2.0 0.02 0.03 1.5

Costa’s hummingbird 0.03 0.03 1.0 0.01 0.02 2.4 0.02 0.03 2.0

broad-tailed hummingbird 0.05 0.13 2.4 0.02 0.03 1.7

rufous hummingbird 0.01 0.01 2.0 0.01 0.02 2.4

elegant trogon 0.01 0.01 2.4

belted kingfisher <0.01 0.01 2.0

acorn woodpecker 0.11 0.18 1.7

Gila woodpecker 0.40 0.26 0.7 0.22 0.20 0.9 0.02 0.03 2.0

ladder-backed woodpecker 0.18 0.11 0.6 0.11 0.09 0.8 0.13 0.12 0.9

hairy woodpecker 0.01 0.01 2.4 0.14 0.07 0.5

Arizona woodpecker 0.04 0.05 1.2

northern flicker 0.02 0.03 1.5 0.05 0.04 0.9

gilded flicker 0.04 0.03 1.0 0.05 0.07 1.4

northern beardless-tyrannulet 0.03 0.03 1.0

greater pewee 0.01 0.03 2.4

western wood-pewee 0.01 0.01 1.2 0.02 0.03 2.0 0.16 0.24 1.5 0.07 0.12 1.7

gray flycatcher 0.01 0.02 2.0

western flycatcher 0.01 0.01 2.0

cordilleran flycatcher <0.01 0.01 2.0 0.02 0.03 1.5 0.40 0.08 0.2

black phoebe 0.01 0.01 0.8 0.02 0.03 2.0

vermilion flycatcher 0.06 0.11 2.0

dusky-capped flycatcher 0.02 0.04 2.0 0.06 0.06 1.1 0.01 0.02 1.7

ash-throated flycatcher 0.27 0.10 0.4 0.37 0.22 0.6 0.60 0.39 0.7 0.33 0.18 0.6

brown-crested flycatcher 0.51 0.35 0.7 0.18 0.16 0.9 0.03 0.03 1.2

sulphur-bellied flycatcher 0.01 0.03 2.4

Cassin’s kingbird 0.05 0.11 2.0 0.02 0.03 2.0

western kingbird 0.02 0.04 2.0

Bell’s vireo 0.59 0.49 0.8

plumbeous vireo 0.10 0.15 1.6 0.23 0.13 0.6

Hutton’s vireo 0.02 0.03 2.0 0.20 0.13 0.7 0.06 0.10 1.7

warbling vireo 0.01 0.01 2.0 0.01 0.02 2.4 0.02 0.03 1.7

Steller’s jay 0.01 0.02 2.4 0.30 0.23 0.8

western scrub-jay 0.02 0.02 1.2 0.01 0.02 2.4 0.06 0.09 1.4

Mexican jay 0.18 0.16 0.9

common raven 0.01 0.01 2.0 0.01 0.02 2.4 0.04 0.08 1.7

purple martin 0.09 0.07 0.8

violet-green swallow 0.04 0.08 1.7

mountain chickadee 0.58 0.16 0.3

bridled titmouse 0.02 0.03 2.0 0.06 0.09 1.4 0.08 0.09 1.0

verdin 1.07 0.37 0.3 0.49 0.28 0.6 0.03 0.07 2.0

bushtit 0.25 0.29 1.1 0.40 0.43 1.1 0.08 0.14 1.7

red-breasted nuthatch 0.03 0.05 1.7
56

Riparian 

(n = 4)

Sonoran Desertscrub 

(n = 6)

Oak 

Savannah (n = 4)

Pine-oak 

Woodland (n = 6)

Conifer Forest 

(n = 3)

Species Mean SD CV Mean SD CV Mean SD CV Mean SD CV Mean SD CV

white-breasted nuthatch 0.02 0.03 2.0 0.08 0.11 1.3 0.33 0.29 0.9

pygmy nuthatch 0.17 0.23 1.4

brown creeper 0.14 0.07 0.5

cactus wren 0.57 0.26 0.5 0.75 0.39 0.5 0.21 0.24 1.2

rock wren 0.12 0.15 1.3 0.14 0.10 0.7 0.06 0.08 1.3

canyon wren 0.04 0.05 1.3 0.06 0.07 1.1 0.08 0.06 0.8 0.11 0.08 0.7 0.03 0.03 0.9

Bewick’s wren 0.48 0.22 0.4 0.01 0.02 2.4 0.77 0.09 0.1 1.29 0.26 0.2 0.04 0.08 1.7

house wren <0.01 0.01 2.0 0.21 0.22 1.1

ruby-crowned kinglet 0.01 0.01 1.3

blue-gray gnatcatcher 0.02 0.01 0.7 0.13 0.13 1.1 0.29 0.33 1.1 0.03 0.05 1.7

black-tailed gnatcatcher 0.12 0.09 0.8 0.25 0.23 0.9

western bluebird 0.01 0.02 2.4 0.17 0.16 0.9

hermit thrush 0.13 0.22 1.7

American robin 0.01 0.01 2.4 0.22 0.15 0.7

northern mockingbird 0.02 0.02 1.4 0.07 0.07 1.1 0.33 0.58 1.7

curve-billed thrasher 0.33 0.19 0.6 0.20 0.19 0.9 0.02 0.03 2.0

crissal thrasher 0.02 0.02 1.2

European starling 0.01 0.01 2.0

phainopepla 0.06 0.06 0.9 0.01 0.02 2.4 0.22 0.30 1.4

olive warbler 0.01 0.01 2.4 0.13 0.18 1.4

orange-crowned warbler <0.01 0.01 2.0 0.01 0.02 2.4 0.01 0.02 2.4

Virginia’s warbler 0.01 0.02 2.0 0.02 0.03 2.0 0.15 0.14 1.0 0.06 0.10 1.7

Lucy’s warbler 0.96 0.35 0.4 0.01 0.02 2.4

yellow warbler 0.05 0.07 1.4

yellow-rumped warbler 0.01 0.01 1.3 0.27 0.11 0.4

black-throated gray warbler 0.01 0.02 2.0 0.02 0.03 1.5 0.27 0.38 1.4 0.58 0.21 0.4 0.14 0.24 1.7

Townsend’s warbler 0.06 0.10 1.7

Grace’s warbler 0.15 0.33 2.3 0.34 0.14 0.4

MacGillivray’s warbler 0.01 0.01 2.0

Wilson’s warbler 0.06 0.03 0.5 0.01 0.02 2.4 0.02 0.03 2.0 0.02 0.03 1.5

red-faced warbler 0.01 0.02 2.4 0.34 0.32 0.9

painted redstart 0.04 0.09 2.4

hepatic tanager 0.02 0.03 2.0 0.15 0.22 1.4 0.07 0.12 1.7

summer tanager 0.11 0.12 1.1

western tanager 0.01 0.01 2.0 0.03 0.07 2.0 0.06 0.09 1.5 0.43 0.43 1.0

green-tailed towhee 0.09 0.13 1.4 0.01 0.02 2.4 0.02 0.05 2.4

spotted towhee 0.01 0.01 2.0 0.13 0.25 2.0 1.25 0.36 0.3 0.40 0.44 1.1

canyon towhee 0.28 0.12 0.4 0.34 0.08 0.2 0.14 0.13 0.9

Abert’s towhee 0.20 0.03 0.1

rufous-winged sparrow 0.16 0.09 0.6 0.03 0.05 1.7

rufous-crowned sparrow 0.01 0.01 1.4 0.31 0.28 0.9 0.71 0.33 0.5 0.28 0.26 0.9

chipping sparrow 0.01 0.02 1.2 0.01 0.02 2.4 0.01 0.02 2.4

Brewer’s sparrow 0.15 0.17 1.1 0.03 0.08 2.4

black-chinned sparrow 0.01 0.02 2.4 0.11 0.22 2.0 0.15 0.17 1.2

Lincoln’s sparrow <0.01 0.01 2.0

lark sparrow 0.02 0.02 1.2

black-throated sparrow 0.18 0.06 0.3 0.79 0.38 0.5 0.05 0.10 2.0

white-crowned sparrow 0.01 0.02 2.0 0.01 0.02 2.4 0.02 0.03 2.0

yellow-eyed junco 0.03 0.05 1.7 1.03 0.47 0.5

northern cardinal 0.46 0.32 0.7 0.06 0.08 1.3

pyrrhuloxia 0.07 0.07 1.1 0.02 0.03 1.5

black-headed grosbeak 0.01 0.01 2.0 0.01 0.02 2.4 0.11 0.18 1.7 0.19 0.24 1.3 0.15 0.17 1.1

blue grosbeak 0.10 0.10 1.1 0.01 0.02 2.4

lazuli bunting 0.04 0.06 1.6

indigo bunting <0.01 0.01 2.0

varied bunting 0.10 0.12 1.2 0.02 0.05 2.4

brown-headed cowbird 0.19 0.13 0.7 0.03 0.05 1.7 0.10 0.12 1.3 0.02 0.03 1.7

hooded oriole <0.01 0.01 2.0

Bullock’s oriole 0.02 0.01 0.7 0.05 0.08 1.6

Scott’s oriole 0.01 0.01 2.0 0.05 0.10 1.9 0.14 0.12 0.8 0.03 0.04 1.1

house finch 0.27 0.13 0.5 0.37 0.42 1.2 0.24 0.43 1.8

pine siskin 0.01 0.02 1.7

lesser goldfinch 0.22 0.18 0.8 0.02 0.05 2.4 0.01 0.01 2.4
57

surveys (yellow-breasted chat and house sparrow; 

Table 5.5). We observed only four species 

(brown-crested flycatcher, mourning and whitewinged doves, and western tanager) at > 50 of the 

transects. This is in contrast to the repeat-visit 

VCP surveys (Appendix G) and is likely because 

by visiting a station only once, we missed species 

that would probably be recorded on subsequent 

surveys. 

Line-transect Surveys

We found 63 species during line-transect surveys 

in the winter of 2002 and 2003 including six 

species that we did not record during VCP 

surveys (Appendix C). We found the most 

species along the Lower Rincon Creek transect 

(n = 45) and fewest along the Douglas Springs 

transect (n = 31; Table 5.6). The chipping 

sparrow was the most abundant species on two 

transects. All three of the most abundant species 

on the Upper Loma Verde transect (chipping 

sparrow, green-tailed towhee, and Brewer’s 

sparrow) did not breed in the Sonoran Desert 

region, whereas the three most abundant species 

along the Lower Rincon Creek transect (blackthroated sparrow, cactus wren, and Gambel’s 

quail) did breed in the district. Two of the three 

most abundant species along the Douglas Springs 

transect (chipping sparrow and western bluebird) 

did not breed in the district. 

Nocturnal Surveys

We recorded nine species during nocturnal 

surveys of nine transects (Table 5.7). We found 

the most species along the Rincon Creek and 

low-elevation transects combined, though survey 

effort was greatest there (Table 5.2). The most 

abundant species within a stratum was the elf owl 

in the low-elevation stratum (Table 5.7). Only 

two species were found only in a single stratum 

and no species were found in all three. The 

great-horned owl was found in the low- and highelevation strata and was certainly missed in the 

middle-elevation stratum.

Incidental and Breeding Observations

We observed 154 species during incidental 

observations, including 13 species that we did 

not record during other surveys (Appendix C). 

We made 288 observations of 78 species that 

confirmed breeding in or near the district (Table 

5.8). Of these we found 104 nests of 48 species 

including a nest of the sulphur-bellied flycatcher 

near Happy Valley Saddle. We found two 

instances of brown-headed cowbird parasitism: 

one blue-gray gnatcatcher feeding a fledgling 

cowbird and one Bell’s vireo nest with a cowbird 

egg. 

Inventory Completeness

The bird survey effort at the Rincon Mountain 

District of Saguaro National Park was the most 

comprehensive of the eight park units surveyed 

by the UA inventory group. We made over 

15,000 observations and found 85% (n = 173) 

of the species that had been found previously 

in the district (Appendix C), and found 10 new 

species. These results are unprecedented in the 

Sonoran Desert Network, and are especially 

important given the large size and diversity of 

communities and difficult access issues in the 

district. Considering all of the other research and 

site-specific inventory efforts in the district (see 

review at the beginning of the chapter), we are 

confident in concluding that at least 90% of the 

species that regularly occur in the district have 

been recorded. 

The species accumulation curve for our 

research (from all surveys combined; Fig. 5.4) 

shows the number of new species dropping off 

significantly at approximately halfway through 

the inventory. After the first half of the surveys, 

we found only an additional 8% (n = 14 species) 

of the species found in the entire effort. A closer 

look at the species accumulation curve for repeatvisit VCP surveys reveals that the Riparian 

community had the most complete inventory, 

though the other communities show signs of 

reaching an asymptote, particularly the Conifer 

Forest community (Fig. 5.5). There is a similar 

pattern for the line-transect surveys (Figure 5.6).
58

Table 5.5. Mean relative abundance of birds from reconnaissance VCP surveys by strata and transect, 

Saguaro National Park, Rincon Mountain District, 2002. 

Elevation Stratum

Lowa Middleb Highc

Species BWF BT LVW FR FW CC MC DS JS JB NST MMT MM M IS DHS

Gambel’s quail 1.5 0.8 1.5 0.3 0.3

zone-tailed hawk 0.4

white-winged dove 0.3 0.6 0.5 0.9 1.5 0.7 0.3 0.2

mourning dove 0.3 1.3 0.3 0.3 0.8 0.3 0.2 0.2 0.1

black-chinned hummingbird 0.3

Anna’s hummingbird 0.1 0.2

broad-tailed hummingbird 0.1 0.1

rufous hummingbird 0.1

acorn woodpecker 0.2

Gila woodpecker 0.6 0.3 0.8 0.3 0.3 0.3

hairy woodpecker 0.6 0.4 0.3 0.1 0.2

northern flicker 0.2 0.1 0.3 0.2

gilded flicker 0.2

northern beardless-tyrannulet 0.1

greater pewee 0.1 0.1

western wood-pewee 0.2 0.4 1.0

cordilleran flycatcher 1.0 0.5 0.2 0.4 0.3

black phoebe 0.1

dusky-capped flycatcher 0.2 0.3

ash-throated flycatcher 0.1 0.3 0.4 0.4

brown-crested flycatcher 0.6 0.1 0.8 0.1 1.0 0.5 0.8 0.2 0.2

Cassin’s kingbird 0.1

Bell’s vireo 0.8 1.2

plumbeous vireo 0.4 0.1 0.4 0.4 0.4

Hutton’s vireo 0.2

warbling vireo 0.3 0.1

Steller’s jay 0.5 0.2 0.3

Mexican jay 0.6 0.2

purple martin 0.4 0.2 0.3

mountain chickadee 0.4 0.5 0.2 0.3 0.3

bridled titmouse 0.2

verdin 0.7 0.9 0.8 0.4 0.8 1.0 1.3

bushtit 1.6

red-breasted nuthatch 0.3

white-breasted nuthatch 0.2 0.6 0.4 0.2 0.8 0.4 0.2

pygmy nuthatch 0.3 1.0 0.6

brown creeper 0.1 0.2

cactus wren 1.3 0.8 0.5 0.9 1.3 0.7 1.3

canyon wren 0.3 0.3

Bewick’s wren 0.1 0.3 0.4 0.4

house wren 0.2 0.5 0.2 0.4 0.4 0.6

blue-gray gnatcatcher 0.6

black-tailed gnatcatcher 0.3 0.8 0.1

western bluebird 0.3 0.4

hermit thrush 0.4 0.4 0.1

American robin 0.3 0.4 0.2

northern mockingbird 0.1

curve-billed thrasher 0.1 0.5 0.1 0.4 0.3 0.3

phainopepla 0.1

olive warbler 0.3 0.4

Virginia’s warbler 0.1

Lucy’s warbler 1.0 0.3 0.5

yellow-rumped warbler 0.6 0.4 0.4 0.1 0.2

black-throated gray warbler 0.5 0.6

Grace’s warbler 0.3 0.8 0.4 0.5 0.2

red-faced warbler 0.4 0.2 0.3 0.4

painted redstart 0.1

yellow-breasted chat 0.1

hepatic tanager 0.5 0.2 0.1
59

Elevation Stratum

Lowa Middleb Highc

Species BWF BT LVW FR FW CC MC DS JS JB NST MMT MM M IS DHS

western tanager 0.3 0.2 0.8 0.5 0.6 0.9 0.5 0.2

spotted towhee 0.8 0.4 0.1 1.4 1.4

canyon towhee 0.3 0.4 0.3

rufous-winged sparrow 0.5 0.4 0.1 0.3 0.2

rufous-crowned sparrow 0.1

black-chinned sparrow 0.1

black-throated sparrow 1.3 0.3 0.2

yellow-eyed junco 0.6 0.5 1.6 0.6 0.3 1.0

northern cardinal 0.6 0.1 0.1 0.3 0.5 0.7

pyrrhuloxia 0.1 0.1 0.1 0.1 0.3

black-headed grosbeak 0.1 0.3 0.2 0.3 0.1 0.2

blue grosbeak 0.1 0.2

varied bunting 0.2 0.2

brown-headed cowbird 0.1 0.1 0.1 0.1 0.3 0.3 0.2

house finch 0.8 0.1 1.1 0.5 0.2

house sparrow 0.3

a

 < 4000 feet elevation: BWF = Bridal Wreath Falls, BT = Broadway Trailhead, LVW = Loma Verde Wash, FR = Freeman Road, 

FW = Freeman Wash, CC = Chimenea Canyon, MC = Madrona Canyon.

b

 4,000 – 6,000 feet elevation: DST = Douglas Springs Trail, JB = Juniper Springs.

c > 6,000 feet elevation: JB = Juniper Basin, NST = North Slope Trail, MMT = Mica Mountain Trail, MM = Mica Mountain, 

M = Manning Cabin, IS = Italian Springs, DHS = Deer Head Spring.

We believe that we recorded all but a 

few species that were breeding in the district at 

the time of the inventory. The breeding status 

of only a few species remains questionable, 

either because we did not record them during 

the breeding season, or because we failed to 

document breeding activity. Species that we 

believe are regular breeders in the district, but 

for which there has been no evidence of breeding 

(Short 1996, Frederici 1998, Powell 1999, 2004) 

include the sharp-shinned hawk, gray vireo, 

northern beardless-tyrannulet, northern roughwinged swallow, loggerhead shrike, juniper 

titmouse, Bendire’s thrasher, European starling, 

yellow-breasted chat, bronzed cowbird, and pine 

siskin. All of these species were seen only a few 

times or not at all during the peak breeding time 

for the species. Most species that we observed 

throughout the breeding season are assumed to 

have bred in the district, even though we found 

no evidence of nesting (Table 5.8; see also 

Appendix C for list of all species that have been 

observed breeding in the district). This group 

includes the greater roadrunner, western scrubjay, red-breasted nuthatch, and brown creeper. 

Also, there are at least two species (wild turkey 

and scaled quail) that we observed only once but 

that we assume nested in the district because they 

maintain year-round home ranges that probably 

include the district. Species that we saw during 

the breeding season, but that were unlikely to 

have nested in the district (because we made an 

effort to determine their breeding status), were 

the yellow-billed cuckoo, buff-breasted flycatcher 

(a single male was observed in the same location 

for four years; Chris Kirkpatrick, pers. comm.), 

and elegant trogon.

Based on nesting records or possible 

nesting attempts in nearby areas (e.g., Corman 

and Wise-Gervais 2005), there are a number of 

species that may have nested in the recent past 

or may nest in the district irregularly. We review 

these species by vegetation community: 

•	 Low-elevation Sonoran Desertscrub/ 

Southwestern Deciduous Riparian: ruddy 

ground dove (Columbina talpacoti), 

buff-collared nightjar (Caprimulgus 

ridgwayi), violet-crowned hummingbird 

(Amazilia violiceps), northern roughwinged swallow, thick-billed kingbird 

(Tyrannus crassirostris), and indigo bunting 

(Passerina cyanea).

•	 Semi-desert Grassland and/or Oak 

Savannah: northern harrier and Swainson’s 

hawk. 

•	 Pine-oak and/or Conifer Forests: northern 

saw-whet owl (Aegolius acadicus), 

long-eared owl (Asio otus), white-eared 
60

Transect

Upper Loma Verde 

(n = 36)

Lower Rincon Creek 

(n = 52)

Douglas Spring 

(n = 48)

Species Mean SE Mean SE Mean SE

Gambel’s quail 0.86 0.372 0.94 0.436 0.19 0.132

Cooper’s hawk 0.04 0.027

red-tailed hawk 0.06 0.033 0.04 0.029

American kestrel 0.03 0.028 0.02 0.019 0.04 0.029

mourning dove 0.17 0.085 0.12 0.045

great horned owl 0.06 0.039

Anna’s hummingbird 0.03 0.028

Gila woodpecker 0.86 0.160 0.71 0.133 0.02 0.021

red-naped sapsucker 0.02 0.019

ladder-backed woodpecker 0.31 0.104 0.29 0.084 0.06 0.035

northern flicker 0.17 0.053 0.08 0.050

gilded flicker 0.08 0.061 0.23 0.081

black phoebe 0.02 0.019

Say’s phoebe 0.03 0.028 0.02 0.019

ash-throated flycatcher 0.02 0.019

loggerhead shrike 0.04 0.029

western scrub-jay 0.19 0.078 0.08 0.037 0.25 0.082

Mexican jay 0.13 0.092

common raven 0.06 0.039 0.02 0.019 0.23 0.189

violet-green swallow 0.11 0.111

bridled titmouse 0.46 0.193

juniper titmouse 0.04 0.042

verdin 0.42 0.101 0.48 0.105 0.27 0.077

bushtit 0.69 0.455

white-breasted nuthatch 0.02 0.021

cactus wren 0.78 0.155 1.04 0.162 0.17 0.069

rock wren 0.03 0.028 0.12 0.045

canyon wren 0.04 0.029

Bewick’s wren 0.42 0.092 0.33 0.081 0.56 0.094

house wren 0.15 0.051

ruby-crowned kinglet 0.39 0.121 0.42 0.104 0.42 0.102

black-tailed gnatcatcher 0.25 0.092 0.13 0.062

western bluebird 0.06 0.058 0.79 0.339

Townsend’s solitaire 0.35 0.109

hermit thrush 0.04 0.027

American robin 0.06 0.058 0.04 0.029

northern mockingbird 0.08 0.047

curve-billed thrasher 0.75 0.151 0.63 0.113

crissal thrasher 0.03 0.028 0.17 0.062

cedar waxwing 0.42 0.297

phainopepla 0.11 0.066 0.08 0.037 0.04 0.029

yellow-rumped warbler 0.08 0.083

green-tailed towhee 1.22 0.165 0.08 0.037

spotted towhee 0.17 0.063 0.02 0.019 0.44 0.084

canyon towhee 0.58 0.175 0.37 0.087 0.81 0.165

Abert’s towhee 0.03 0.028 0.13 0.048

rufous-winged sparrow 0.89 0.182 0.88 0.144

rufous-crowned sparrow 0.52 0.115

chipping sparrow 2.36 0.785 0.40 0.279 2.54 0.932

Brewer’s sparrow 1.03 0.477 0.10 0.096

black-chinned sparrow 0.06 0.033

vesper sparrow 0.08 0.046

Lincoln’s sparrow 0.12 0.052

black-throated sparrow 0.33 0.120 1.17 0.329

white-crowned sparrow 0.72 0.162

dark-eyed junco 0.06 0.028 0.25 0.082 0.25 0.053

northern cardinal 0.19 0.087 0.37 0.126

pyrrhuloxia 0.03 0.028 0.10 0.050

eastern meadowlark 0.10 0.074

house finch 1.00 0.298 0.62 0.135 0.06 0.035

pine siskin 0.10 0.096

lesser goldfinch 0.52 0.295

Lawrence’s goldfinch 0.03 0.028

Table 5.6. Relative abundance (mean + SE) of birds from line-transect surveys, Saguaro National Park, Rincon 

Mountain District, 2002 and 2003. 
61

Table 5.7. Mean relative abundance of birds from nocturnal surveys by elevation strata and transect, 

Saguaro National Park, Rincon Mountain District, 2001 and 2002. 

Low elevation Middle elevation High elevation

Species

Cactus Forest 

Loop Drive

Rincon

Creek

Box

Canyon

Cowhead

Saddle

Happy Valley 

Saddle

Juniper 

Basin Manning

Italian

Spring

Spud

Rock

barn owl 0.05

flammulated owl 0.33 0.25

western screech-owl 0.7 0.55 1.38 1.67

whiskered screech-owl 1 0.33 0.19

great horned owl 0.37 0.05 0.5 0.06

elf owl 2.37 1.77 2 0.13 0.33

lesser nighthawk 0.07 0.05

common poorwill 0.57 0.59 0.13

whip-poor-will 0.67 1.13 0.5 0.25

 

Nest

Adults carrying 

objects Other 

Species Building

With 

eggs

With 

young

Occupied FoodNesting 

material

Distraction 

displays

Feeding recently 

fledged young

Recently 

fledged 

young Totals

Gambel’s quail 1 3 4

Cooper’s hawk 1 1

northern goshawk 2 2

Harris’s hawk 1 1

zone-tailed hawk 2 2

red-tailed hawk 1 1 2

band-tailed pigeon 1 1

white-winged dove 4 1 5

mourning dove 4 1 5

great horned owl 2 2

lesser nighthawk 1 1

common poorwill 1 1 2

whip-poor-will 1 1

black-chinned hummingbird 1 1 2

Costa’s hummingbird 2 2

acorn woodpecker 1 1

Gila woodpecker 4 2 3 1 10

hairy woodpecker 1 1

Arizona woodpecker 1 1

gilded flicker 1 2 3

western wood-pewee 1 1 1 3

cordilleran flycatcher 1 1 2

Say’s phoebe 1 1

vermilion flycatcher 1 1

ash-throated flycatcher 3 3 3 9

brown-crested flycatcher 4 3 1 1 9

sulphur-bellied flycatcher 1 1 2

Cassin’s kingbird 2 2

western kingbird 1 1

Bell’s vireo 3 8 1 12

plumbeous vireo 1 1 3

Hutton’s vireo 2 2

warbling vireo 1 1

Mexican jay 1 1

common raven 1 1

purple martin 8 2 10

bridled titmouse 1 1

Table 5.8. Number of breeding behavior observations for birds from all survey types, Saguaro National Park, 

Rincon Mountain District, 2001 and 2002. Breeding behaviors follow standards set by NAOAC (1990). 
62

Discussion

The bird community in the Rincon Mountain 

District of Saguaro National Park is diverse and 

is a function of the many biotic communities 

present in the Rincon Mountains, from Sonoran 

Desertscrub to Conifer Forest. Vegetation 

responds to the extreme differences in elevation, 

soils, and rainfall (see Chapter 3), and vegetation 

is one of the most important predictors of 

bird community structure (James 1971). This 

relationship is supported by the results of our 

inventory; the 23 repeat-visit VCP transects 

were classified into five distinct communities 

(Fig. 5.3). Important vegetation characteristics 

Nest

Adults carrying 

objects Other 

Species Building

With 

eggs

With 

young

Occupied FoodNesting 

material

Distraction 

displays

Feeding recently 

fledged young

Recently 

fledged 

young Totals

verdin 2 1 1 2 6

bushtit 1 1 2

white-breasted nuthatch 1 1 2

pygmy nuthatch 1 1 2

cactus wren 1 2 3

rock wren 1 1 2 4

canyon wren 1 1

Bewick’s wren 1 1 3 1 6

house wren 1 1 2

blue-gray gnatcatcher 1 1 1 2 5

black-tailed gnatcatcher 1 2 3

western bluebird 1 2 3

hermit thrush 1 1

American robin 1 1

northern mockingbird 1 1

curve-billed thrasher 3 1 1 3 8

phainopepla 1 1 2

Virginia’s warbler 3 2 5

Lucy’s warbler 2 4 1 7

yellow-rumped warbler 1 1 1 1 4

black-throated gray warbler 1 5 6

Grace’s warbler 2 1 2 5

red-faced warbler 2 1 3 6

painted redstart 1 1 2

hepatic tanager 1 1 1 3

summer tanager 1 1

western tanager 1 2 1 4

spotted towhee 1 3 1 8 13

canyon towhee 1 1 1 3

Abert’s towhee 1 1 2

rufous-winged sparrow 1 1

rufous-crowned sparrow 3 1 7 11

black-chinned sparrow 2 3 4 9

black-throated sparrow 1 1 13 15

yellow-eyed junco 2 1 4 7 14

northern cardinal 2 4 6

pyrrhuloxia 1 1

blue grosbeak 1 1

brown-headed cowbird 2 2

Scott’s oriole 1 1 3

house finch 1 1 2

hummingbird (Hylocharis leucotis), 

blue-throated hummingbird (Lampornis 

clemenciae), golden-crowned kinglet 

(Regulus satrapa), Townsend’s solitaire 

(Myadestes townsendi), flame-colored 

tanager (Piranga bidentata), red crossbill 

(Loxia curvirostra), and evening grosbeak 

(Coccothraustes vespertinus). 

The district will likely gain some nesting 

species in the coming few years. For example, 

one non-native species, the Eurasian collareddove (Streptopelia decaocto) has recently 

established breeding populations in the region 

(Corman and Wise-Gervais 2005).
63

Sample period

0 10 20 30 40 50

Cumulative number of species

60

80

100

120

140

160

180

200

Figure 5.4. Species accumulation curve for all survey methods for birds, Saguaro National 

Park, Rincon Mountain District, 2001 and 2002. Each sample period is a randomized 

combination of approximately 250 observations. 

that consistently predict occurrence of bird 

species include vertical structure (MacArthur 

and MacArthur 1961, Cody 1981), horizontal 

patchiness (heterogeneity; Roth 1976, Kotliar 

and Weins 1990), and floristics (Rice et al. 1984, 

Strong and Bock 1990). To even the most casual 

observer, there are extreme changes in all of the 

characteristics from the valley floor to the highest 

points of the Rincon Mountains. This pattern 

of vegetation change across altitude and aspect 

is typical of the “sky island” mountain ranges 

of southern Arizona and adjacent Mexico (e.g., 

Whittaker and Niering 1965). 

Although the district contains a number 

of biotic communities that are characteristic 

of the sky island mountains, it shares one of 

the biogeographic traits with the herpetofauna 

community: it is not as species rich as the sky 

island ranges to the south. In particular, ranges 

in the U.S., such as the Chiricahua (Taylor 1997) 

and Huachuca mountains regularly host breeding 

species that have strictly Madrean distributions 

including the Lucifer (Calothorax lucifer), 

Berylline (Amazilia beryllina), and violetcrowned (Amazilia violiceps) hummingbirds, 

eared trogon (Euptilotis neoxenus), Mexican 

chickadee (Poecile sclateri), and flame-colored 

tanager (Piranga bidentata), to name a few. 

Although it likely that some of these species 

(e.g., blue-throated hummingbird [Lampornis 

clemenciae]) occasionally appear in the Rincon 

Mountains (see Inventory Completeness), our 

surveys provide further evidence that these 

species do not regularly occur there. Two species 

that reach their northern breeding distribution 

in the district (or nearby mountain ranges) are 

the buff-breasted flycatcher and sulphur-bellied 

flycatcher. We found the first confirmation of 

breeding for the sulphur-bellied flycatcher in 

the district, and the buff-breasted flycatcher 

may breed there occasionally. A third Madrean 

species, the elegant trogon, may also occasionally 

breed in the Rincon Mountains, but there has 

been no confirmation of this. 

An important resource for birds in the 

district is the riparian corridor along Rincon 

Creek, which had higher species richness than 

any other area in the district (Appendix G). 
64

Powell (2004) compared the bird community 

along Rincon Creek to adjacent upland sites and, 

partially using data contained in this report, found 

the riparian area to have more than twice as many 

species. Studies elsewhere in the Southwest 

have found similar patterns (Carothers et al. 

1974, Whitmore 1975). Even among riparian 

areas of the district, the Lower Rincon Creek 

transect stands out as the most species-rich area 

of the district for both VCP (Appendix G) and 

line-transect surveys (Table 5.6). We found four 

species that were restricted to riparian areas in 

the Southwest (Rosenberg et al. 1991) and that 

were consistent members of the bird community 

along Rincon Creek: Bell’s vireo, Abert’s towhee, 

summer tanager, and yellow warbler (Table 

5.4). Other riparian species that we observed 

along Rincon Creek included the mallard, gray 

hawk, belted kingfisher, and northern beardlesstyrannulet. 

Although riparian areas in the 

Southwest, such as Rincon Creek, are home to 

a disproportionate number of bird species, these 

areas are decreasing in both size and habitat 

quality (Rosenberg et al 1991, Russell and 

Monson 1998). This is evident along Rincon 

Sample period

0 10 20 30

Cumulative number of species

0

20

40

60

80

100

Sonoran Desert Upland

Oak Savanna

Pine-oak Woodland

Conifer Forest

Sample period

0 20 40 60 80

Cumulative number of species

0

20

40

60

80

100

Riparian

Figure 5.5. Species accumulation curves for repeat-visit VCP transects from the five communities, Saguaro 

National Park, Rincon Mountain District, 2001 and 2002. Each sample period is a randomized combination 

of approximately 50 observations. Data include all observations from VCP surveys including flyovers and bird 

observed within 300 m of stations. Note difference in scale for sampling period. 
65

Creek where many of the large riparian trees 

are in poor condition (Powell 2004); some 

loss of riparian bird species may have already 

occurred. For example, the yellow-billed cuckoo 

and song sparrow, both riparian obligate birds 

are common in nearby Cienega Creek (Corman 

and Magill 2000), and there is no reason to 

believe that with healthier riparian vegetation 

these species would not be found along Rincon 

Creek as well. The current drought has certainly 

affected the health of the riparian system. The 

decline in the amount and timing of surface water 

availability is also likely affected by the recent 

increase in groundwater pumping supplying the 

explosive growth of housing and commercial 

development in the Rincon Valley (see Chapter 

2). Because birds are so closely tied to vegetation 

characteristics, the loss and degradation of large 

riparian trees will mean a reduction in the number 

of species and abundance of some riparianobligate birds along the creek. The threat of 

losing groundwater and (by extension) surface 

water to development, recently prompted the park 

to file in-stream flow water rights in an effort to 

ensure the long-term viability of the riparian area. 

They have also initiated studies of the plant and 

vertebrate communities of the area (e.g., Powell 

2004). 

Housing developments in the Rincon 

Valley, in particular, will also likely affect the 

bird community through an increase in nonnative (rock pigeon, European starling, and house 

sparrow) and human-adapted species (e.g., the 

great-tailed grackle, mourning dove, house finch, 

and brown-headed cowbird). The increase in 

density of human-adapted species invariably 

follows housing developments and these changes 

usually lead to the decline in densities of nonhuman-adapted species, especially in the areas 

immediately adjacent to development (Mills et 

al. 1989, Germaine et al. 1998). Mannan and 

Bibles (1989) suggest a number of ways to reduce 

the impact of non-native bird species on the 

district’s wildlife including (1) limiting density of 

housing near the district boundary, (2) reducing 

the number of horses, (3) limiting sources of open 

water, and (4) limiting landscaping with nonnative plants, especially lawns. Many of these 

measures are effective in reducing native, humanadapted species as well. 

An increase in nearby housing may 

facilitate the spread of non-native plants, which 

Sample period

0 5 10 15 20

Cumulative number of species

10

20

30

40

50

60

70

Figure 5.6. Species accumulation curve for line-transects for birds, Saguaro National 

Park, Rincon Mountain District, 2002-2003. Each sample period is a randomized 

combination of approximately 50 observations. 
66

can impact other native plant and vertebrate 

communities through alteration of vegetation 

structure and ecosystem function. Also 

associated with housing developments are 

increases in the number of free-roaming feral 

pets, which kill and harass native wildlife (Clarke 

and Pacin 2002). Finally, with development 

come roads, which act as barriers to movement 

of wildlife because of direct mortality and 

modification of behavior (e.g., Kline and Swann 

1998, Trombulak and Frissell 2000, Clark et al. 

2001, Cain et al. 2003). 

Wildland fire has always played a major 

role in shaping pine-oak woodlands and conifer 

forests of western North America. At the Rincon 

Mountain District, the forests have experienced 

low- to moderate-intensity burns approximately 

every decade since the 15th century (Swetnam 

and Baisan 1996). Recently (last 100 years) 

active fire suppression has reduced the frequency 

of these low- and moderate-intensity burns, and 

increased the occurrence of high-intensity burns 

(Allen 1996, Pyne 1996, Swetnam et al. 1999) 

that radically alter forest structure (Swetnam and 

Baisan 1996). Using data (in part) from surveys 

in the district, Kirkpatrick and Conway (2006) 

found a number of bird species to be positively 

associated with the occurrence of fire in pine-oak 

woodlands. In particular, they found the hairy 

woodpecker, greater pewee, western wood pewee, 

white-breasted nuthatch, Virginia’s warbler, house 

wren, spotted towhee, and yellow-eyed junco to 

be positively associated with moderate- to highintensity fires. These species were common in 

the Conifer Forest community (Table 5.4) and 

may reflect the recent fire history of these areas. 

Short (2002) studied the effects of prescribed 

fire on the high-elevation bird community of 

the district. She found inconsistent results 

with regard to population changes of the most 

common species, but nest success of the groundnesting yellow-eyed junco declined dramatically 

the year following fires. Recent large standreplacing fires in the nearby Santa Catalina 

Mountains should reinforce to park managers the 

vital role of an active prescribed-burning program 

and a fire management program that allows for 

some natural fires to burn their course. The park 

has both of these programs and they should be 

commended for using fire to restore the district’s 

high-elevation communities. We encourage 

managers to include bird monitoring in these 

programs (see below). 

The district’s bird community has 

undoubtedly undergone significant changes in the 

recent past. In addition to a changed fire regime 

in the high-elevation areas of the district, the 

low-elevation and semi-desert grassland areas 

have experienced an increase in shrubs and cacti. 

Unfortunately, there are no baseline data to which 

we can compare our results. There are a number 

of species that probably occurred in the district 

and that have undergone range-wide population 

declines. Based on its distribution in the 

nearby mountain ranges, the thick-billed parrot 

(Rhynchopsitta pachyrhyncha) was probably 

resident in the Rincon Mountains at the turn of 

the 20th century (Phillips et al. 1964). Similarly 

the Aplomado falcon (Falco femoralis) was 

considered common in the semi-desert grasslands 

of southeastern Arizona in the late 19th and early 

20th centuries, but no longer occurs in the region. 

The district lies within the historical range for 

this species (Keddy-Hector 1998), and based 

on its habitat requirements, it would have been 

likely to occur on the north side of the district 

near Douglas Springs. The eastern bluebird 

probably bred in the district; it bred in Happy 

Valley (just east of the park boundary) and in the 

nearby Santa Catalina Mountains in the 1970s 

(Corman and Wise-Gervais 2005) but no longer 

nests in these areas. There were a few incidental 

records of the California condor (Gymnogyps 

californianus) in the sky island region in the 

1880s (Phillips et al. 1964). 

Additional Research Needed

The bird community along Rincon Creek is 

likely to change more than any other community 

in the district if the drought and groundwater 

pumping continue. The inclusion of birds in the 

I&M program is encouraging and we suggest 

that emphasis be put on important areas such as 

Rincon Creek. Courtney Conway (University of 

Arizona) is preparing to determine reproductive 

success of riparian birds along Rincon Creek and 

similar nearby areas to investigate the impacts of 
67

surface water availability on habitat quality (i.e., 

reproduction). Additional monitoring should be 

focused in the middle- and high-elevation areas 

of district and it may be possible to combine 

some of this monitoring with the fire-effects 

monitoring program. 

Because birds are highly mobile, we 

expect new species will be added to the district 

list for years to come. Surveys in areas that are 

difficult to access (e.g., Douglas Springs area) 

will be most likely to yield new species. Also, 

surveys during the fall, winter, and early spring 

will likely add species to the list. We also 

encourage the breeding-status clarification of a 

number of species that we expect breed in the 

district, but that we were not able to confirm (see 

Inventory Completeness).
68
69

Chapter 6: Mammal Inventory

Don E. Swann and Brian F. Powell

Previous and Ongoing Research 

Saguaro National Park has never had a 

comprehensive survey of its mammals, and 

surprisingly little research has been conducted 

on mammals in the Rincon Mountain District 

considering the park’s long history as a national 

park. However, a few studies provide valuable 

information on mammals, particularly Lowell 

Sumner’s work in the mid-20th Century (Sumner 

1951) and Russell Davis and Ronnie Sidner’s 

survey of mammals in the high country of the 

Rincons in the early 1990s (Davis and Sidner 

1992). H. Brown and L. Huey (unpubl. data) 

made collecting trips to the Rincons in 1911 and 

1932, respectively (Davis and Sidner 1992). In 

addition, the park’s administrative records at the 

Western Archaeological and Conservation Center 

contain invaluable files (dating from the 1940s 

and 1950s) on mammal sightings and species of 

concern including the Mexican gray wolf and tree 

squirrels. 

More recently, M’Closkey (1980 and 

citations therein) and Duncan (1990) trapped 

small mammals in desert areas of the district. 

Albrecht (2001) and Flesch (2001), using the 

small-mammal trapping data from this inventory 

effort, analyzed patterns of species richness 

and relative abundance for both units of the 

district. (Copies of these reports are available 

in the archive locations cited in Chapter 1). 

Small mammals were also included in surveys 

of the Rincon Valley expansion area in the 

1990s (Fitzgerald 1996, Bucci 2001) and in the 

recent PULSE study of the Madrona Pools area 

(Swann 2003). Both large and small mammals 

were included in surveys of the Rocking K 

Ranch adjacent to the district during the early 

1990s, but most of the large mammals recorded 

in these surveys (Lynn 1996) are based on 

sightings by local residents that may not be 

credible. The small mammal report by Fitzgerald 

(1996) contains a species (hispid pocket mouse 

[Perognatus hispidis]) not previously known to 

occur in the Rincon Mountains and Fitzgerald did 

not collect a specimen voucher. Similarly, a large 

mammal report for the expansion area (Fitzgerald 

1996) is based largely on identification of scat 

and burrows, which we do not consider reliable. 

The inventory of bats is probably nearly complete 

because of Ronnie Sidner’s extensive surveys 

for the last 15 years (Sidner 1991, Sidner and 

Davis 1994, Sidner 2003). Finally, park staff 

have been collecting observations of wildlife 

for several decades. Most of these sightings, 

while not entirely reliable, have been entered 

into a database and mapped in a GIS, and are 

available in a supplement to this report. Other 

sightings remain uncataloged in logbooks from 

the Manning Camp Ranger Station and other 

sources; many of these uncataloged sightings 

were summarized by Davis and Sidner (1992). 

Methods 

We surveyed for mammals using five field 

methods: (1) trapping for rodents and ground 

squirrels (primarily nocturnal; hereafter referred 

to collectively as small mammals), (2) infraredtriggered photography for medium and large 

mammals, (3) netting for bats, (4) pitfall traps 

for shrews and pocket gophers, and (5) incidental 

observations for all mammals. 

Small Mammals

Field Methods

We trapped small mammals using Sherman 

live traps (folding aluminum or steel, 3 x 3.5 

x 9”; H. B. Sherman, Inc., Tallahassee, FL) 

set in grids (White et al. 1983) along focalpoint transects; Figs. 6.1, 6.2). We opened 

and baited (one tablespoon: 16 parts dried 

oatmeal to one part peanut butter) traps in the 

evening, then checked and closed traps the 

following morning. We placed a small amount 

of polyester batting in each trap to prevent trap 

deaths due to cold nighttime temperatures. We 

marked each captured animal with a permanent 

marker to facilitate recognition; these “batch 
70

marks” appeared to last for the duration of the 

sampling period. For each animal we recorded 

species, sex, age class (adult, subadult, or 

juvenile), reproductive condition, weight, and 

measurements for right-hind foot, tail, ear, head, 

and body. For males we recorded reproductive 

condition as either scrotal or non-reproductive; 

for females we recorded reproductive condition 

as one or more of the following: non-reproducing, 

open pubis, closed pubis, enlarged nipples, small 

or non-present nipples, lactating, post-lactating, 

or non-lactating.

Spatial Sampling Design

The majority of our trapping effort in 2001 

was at focal-point transects set throughout the 

district (Fig. 6.3; see Chapter 1). We trapped 

at a subset of nine random transects that were 

surveyed for other taxonomic groups (two, four, 

and three transects in the low-, medium-, and 

high-elevation strata, respectively). We visited 

seven of these transects twice in 2001; repeat 

visits were two to four months apart (Appendix 

I). In 2002 we trapped only at non-random sites 

in areas that we believed would have high species 

richness: two sites along Rincon Creek and one 

site each at Juniper Basin and Douglas Springs 

(Fig. 6.4). We did not revisit non-random sites.

Along each focal-point transect we 

established three grids (Fig. 6.1) with either a 

3x7 or a 5x5 trap configuration (Fig. 6.2). Traps 

set in a 3x7 arrangement had 16.7 m spacing 

among traps and traps in a 5x5 arrangement had 

12.5 m spacing among traps. Occasionally we 

also placed traps “preferentially,” meaning that 

we set traps in locations that the field crews felt 

contained areas with high species richness rather 

than in grids. Typically these “preferential” sites 

were near the random grids; the crew set out

5 to 70 additional traps after setting up the 

random grids (Figs. 6.3, 6.4). At non-random 

sites the layout of traps was variable, but typically 

1000 m 

focal point 50 m 

50 m 

Figure 6.1. Layout of small-mammal trapping grids along focal-point transects, Saguaro National Park, 

2001. See Fig. 6.2 for more details. 

Figure 6.2. Detailed layout of small-mammal trapping grids at Saguaro National Park, 2001 and 

2002. We used 3x7 trap grids in 50x100 m plots (A) from mid‐April through mid‐June and 5x5 trap grids in 

50x50 m plots (B) from mid‐June through October.
71

was in a 5x5 or a 2x10 configuration. The 2x10 

configuration was usually along both edges of a 

wash because we believed that these areas would 

host more animals. 

Temporal Sampling Design

The total number of nights that we trapped each 

grid was variable, but was typically two or three 

nights per visit (see Appendix I). Occasionally 

we trapped for as many as four nights or as few 

as one night. Because our goal was to maximize 

the number of individuals and species trapped, we 

varied the number of nights trapped based on the 

trapping results in the first few nights of trapping; 

if we were catching few animals, we moved to a 

different location. We always trapped at multiple 

plots on the same night to maximize efficiency. 

At focal points we always trapped all the grids 

along the transect on the same nights and 

typically trapped other, nearby non-random areas. 

In some non-random areas (e.g., Douglas Spring) 

we trapped on multiple grids. In this report we 

summarize results by “plot group” which is the 

collection of trapping grids that represent an area. 

Effort

We trapped for 4,589 trap-nights (Table 6.1). 

We had the most trapping effort in the middleelevation stratum (2,195 trap nights), less in the 

high-elevation stratum (1,390 trap nights), and 

the least in the low-elevation stratum (1,004 trap 

nights). In non-random areas, the percentage of 

the total number of trap nights was 36%, 50%, 

and 37% for the low-, middle-, and high-elevation 

strata, respectively (Table 6.1).

Figure 6.3. Locations of random (focal-point transect) small-mammal trapping sites, pitfall traps for 

shrews, and bat-trapping stations, Saguaro National Park, Rincon Mountain District, 2001 and 2002.
72

Analysis

We expressed effort as the number of trap nights 

(number of traps multiplied by number of nights 

they were open) after accounting for sprung traps 

(misfired or occupied; Beauvais and Buskirk 

1999). Sprung traps reduce trap effort because 

they are no longer “available” to capture animals; 

Figure 6.4. Locations of non-random small-mammal trapping sites, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002.

Table 6.1. Summary of small-mammal trapping effort, Saguaro National Park, 

Rincon Mountain District, 2001 and 2002. See Appendix I for additional detail. 

Elevation stratum Location type Number of trapping sites Total trap nights

Low Random 4 721

Non-random 3 284

Middle Random 7 1,094

Non-random 5 1,102

High Random 5 869

Non-random 4 521

we account for this by multiplying the number of 

sprung traps by 0.5 (lacking specific information, 

we estimate sprung traps were available for 

half of the night; Nelson and Clark 1973). We 

calculated relative abundance for species by 

dividing the number of captures by the number 

of trap nights times 100. For this report we 
73

abundant surface water (Fig. 6.3). At most 

sites we set mist nets directly over water and 

varied the number of net hours among sites and 

visits depending on field conditions. We used 

monofilament nylon nets of three net sizes (5-m, 

9-m, or 12-m) depending on the site and set nets 

singly or stacked, depending on conditions. For 

each bat captured, we recorded time of capture, 

species, and sex. When appropriate, we also 

recorded reproductive condition, forearm length, 

mass, body condition, tooth wear, presence 

of parasites, and other measurements. We 

determined whether individuals were adult, 

subadult (by closure of epiphyses), or juvenile (by 

appearance). We estimated age by tooth wear. 

For females, we recorded reproductive condition 

as pregnant (palpation for fetal bones), currently 

lactating (mammary gland with milk), previous 

evidence of lactation (misshapen or scarred 

nipples), or nulliparity (non-use of nipples). We 

determined reproductive condition for males by 

the degree of swelling of testes or the presence of 

black epididymides and used this information to 

determine if the male was not reproductive, semireproductive, or reproductive. We marked all 

captured bats with a temporary, non-lethal marker 

to prevent counting the same individual more 

than once in the same evening. We used sonar 

detectors (Anabat and/or QMC Mini) at all sites 

to aid in determining bat presence/absence and 

relative activity as compared to the visual or mistnet results. We listened passively for the call of 

pallid bats, the only species in the area that can be 

definitively identified by its directive call.

Effort

We visited three roosts that were known, or were 

likely, to have bats. We netted bats at six sites 

for a total of 13 nights of netting in 2001 and four 

nights of netting in 2002 (Appendix J). Most of 

our netting effort was at lower Rincon Creek and 

at Manning Camp Pond; we netted at each site 

for five nights. Deer Creek was the only site at 

which we netted on the east slope of the Rincon 

Mountains. 

Analyses

We report the number of species and individuals 

caught by site, but because of the extreme 

differences in trapping effort among sites we 

calculated relative abundance by plot group, type 

of plot (random or non-random), and visit. 

Pitfall Trapping

It is possible that the Arizona shrew (Sorex 

arizonae) and vagrant shrew (Sorex vagrans) 

occur in the Rincon Mountains; they have been 

found in adjacent mountain ranges in southern 

Arizona (Hoffmeister 1986). Also, pocket 

gophers (Thomomys spp.) are very difficult to 

capture using Sherman traps. To survey for 

shrews and pocket gophers we placed pitfall 

traps (3-quart buckets [19 cm tall x 14 cm wide]) 

in moist, north-facing slopes of the Rincon 

Mountains in 2001. We placed traps adjacent to 

a natural feature such as a fallen log or rock. We 

attempted to check traps every 10 days to two 

weeks.

Effort

We placed traps in three areas: North Slope Trail, 

Italian Spring, and Spud Rock Spring (Fig. 6.3). 

We placed 10 traps (22 May to 24 September) at 

the North Slope Trail site, and four traps each at 

Italian Spring and Spud Rock Spring (6 June to 

10 October). 

Bats

We surveyed for bats using two field methods: 

roost-site visits and netting. For netting, we 

concentrated our survey effort in areas that were 

most likely to have bats, mostly riparian areas 

with surface water present. We did not survey 

for bats near focal points because of the low 

probability of success in these areas. 

Roosts 

We visited roosts that were known to have bats, 

based on historic records, or were likely to have 

bats based on habitat characteristics. At roosts, 

we observed bats with the aid of infrared-filtered 

light and night-vision equipment or red-filtered 

light. When bats were present, we worked 

quickly to identify them to species, but if there 

were no bats we used bright light, then searched 

for and collected skeletal material. 

Mist Netting 

Because most insectivorous bats congregate 

at water sites, we selected sites known to have 
74

calculated percent netting success (PNS) for 

comparisons among sites. We calculated PNS 

as the number of animals caught divided by 

effort (total length of net coverage multiplied 

by amount of time nets were open multiplied 

by 100). We do not attempt to present percent 

netting success as a measure of relative 

abundance because netting bats is somewhat a 

function of chance; many more individuals and 

species can be present in an area than are caught. 

Large and Medium Mammals

Saguaro National Park initiated a medium and 

large mammal inventory in 1999, prior to the 

initiation of the UA inventory effort. In addition 

to support from the NPS, this inventory effort has 

been funded by several small grants to the park, 

and reports have been generated for each of these 

projects (Aslan 2000, Wolf and Swann 2002, 

Swann et al. 2003a, Swann 2003). This report 

combines data presented in these previous reports 

with new data not previously reported. 

Spatial Sampling Design

We used infrared-triggered cameras to detect 

medium and large mammals at a combination of 

random and non-random sites from January 1999 

to June 2005 (Figs. 6.5, 6.6). We located nonrandom sites (Fig. 6.5) primarily at known water 

sources and animal trails. We chose the location 

of these sites to be in areas that we believed 

would have the highest species richness. The 

location of random sites was primarily based on 

the random coordinates chosen as focal points 

for the plant and animal inventories (see Chapter 

1), though many of these focal points were not 

surveyed for the other taxonomic groups. To 

avoid interference with other inventory activities 

at sites where there was other inventory work 

and to maintain consistency among all focal 

points, we offset all camera locations from the 

focal point by using the same coordinates but 

with the NAD 27 map datum instead of NAD 83; 

this moved the focal points approximately 200 

m from the original location. We also generated 

additional random camera locations to increase 

sampling in some areas that were not represented 

by focal points, particularly at high elevations and 

on the east slope of the Rincon Mountains. When 

possible, we placed three camera units at each 

location focal point using the following criteria 

(Fig. 6.7): 

(1) within 50 m of the random point

(2) at a random drainage point nearby 

(selected either randomly within a 1-km 

area; Aslan 2000) or at a random point 

located at the nearest measured point in a 

mapped drainage (Wolf and Swann 2002, 

Swann et al. 2003a), and 

(3) at a non-random point chosen by the field 

technician, usually located between 80 

and 500 m from the random point. 

Temporal design

We generally returned to each camera one week 

after initial setup to check that it was functioning 

properly and to make repairs and change film, 

if necessary. We then left the camera in place 

for approximately two weeks, though the length 

of time varied, especially in remote areas that 

required long days of hiking to reach the camera. 

Field methods

We primarily used the Trailmaster camera 

system at focal points. The system (model 1500, 

Goodson and Associates, Inc., Lenaxa, KS; 

Kucera and Barrett 1993) consists of a transmitter 

that emits an infrared beam, a receiver that 

detects the beam, and a camera that is connected 

to the receiver with a cable (Fig. 6.8). The 

receiver triggers the camera to take a picture 

when an animal breaks the beam. At all nonrandom sites, and occasionally at focal points, 

we also used the DeerCam (model DC-100, Nontypical, Inc., Park Falls, WI) and the Trailmaster 

500 and 1550 models. Because they have 

identical functions, we do not further differentiate 

equipment we used.

We baited each focal-point camera using 

a fish-based canned catfood and a commercial 

trapping lure that attracted predators. Generally, 

we baited with catfood the first week, then the 

trapping lure the second week, but for highelevation surveys in 1999 we randomly selected 

only one bait and used it for two weeks. We 

occasionally baited non-random sites. For visitor 

safety reasons, we did not locate baited stations 

within 200 m of a trail.
75

Effort

We placed cameras at 74 non-random and 40 

random sites throughout the district (Appendix K; 

Figs. 6.5, 6.6). At focal points we had 24 points 

with three cameras, 13 points with two cameras, 

and three points with one camera (Appendix K; 

see Spatial Sampling Design section above for 

more information). Considering both types of 

camera locations (random and non-random), we 

placed most cameras in the low-elevation stratum 

(54%; Table 6.2). Twenty eight percent of the 

cameras were in the middle-elevation stratum, 

and 18% were in the high-elevation stratum. 

The total number of camera nights at all sites 

was 3,895 and the percent of camera nights, by 

elevation stratum, was higher in the low-elevation 

stratum and lower in the other strata: 69%, 18%, 

and 13% in the low-, middle-, and high-elevation 

strata, respectively (Table 6.2).

Analysis

We analyzed all photos and identified the 

animal(s) present. We excluded from analysis 

all non-mammals (birds, reptiles, and blank 

pictures), unknowns that could not be identified 

to genus, humans, horses with riders, and 

nocturnal rodents (mostly woodrats). A 

few species pairs (black-tailed and antelope 

jackrabbits, hooded and striped skunks, and 

Figure 6.5. Locations of non-random infrared-triggered cameras, Saguaro National Park, Rincon 

Mountain District, 2000-2005. 
76

Figure 6.6. Locations of random infrared-triggered cameras, Saguaro National Park, Rincon Mountain 

District, 2000-2005. 

Figure 6.7. Example of three-camera placement at one of the random points, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. Location “R” is the random point, “D” is at the nearest mapped drainage to the 

random point, and “A” is a point chosen by the technician (in this case a natural water hole).
77

white-tailed and mule deer) are difficult to 

distinguish under poor light conditions or if 

only part of the animal is visible; for these we 

made the best possible attempt to distinguish 

them, and sometimes identified the individual 

to genus only. 

We entered these and other data 

(species, number of individuals, film number, 

location, date, time if available, bait, etc.) 

into an Access database. For each random 

area and for each point we summarized the 

number of species and number of individuals 

photographed. To create species distribution 

maps, we converted UTM coordinates to NAD 

83 datum and imported them into ArcView.

Comparing species abundance and 

presence among locations using infraredtriggered photography is problematic. As 

with all methods, animals may not be detected 

because they are absent, or because they were 

present and not detected. In addition, rates 

of detection undoubtedly vary greatly among 

species. Determining relative abundance can 

also be difficult. Infrared-triggered camera 

units often do not operate continuously 

between the time they are set and when they 

are next checked because the roll of film may 

be entirely exposed, or because the unit may 

fail due to technical problems or field errors. 

To estimate rates of detection based on effort, 

we used dates on photographs to determine as 

closely as possible how many days a camera 

unit was operating for each roll of film, then 

summed the number of operational days at 

each location. Where dates were not available 

for a roll of film, we substituted the mean 

number of days it took to fill a 36-exposure roll 

of film (11.8 days).

We compared species richness among 

the three elevation strata and between random 

Receiver Transmitter 

(2) Receiver

triggers

camera to 

take picture 

Infrared beam Camera

(1) Animal blocks infrared 

beam from getting to receiver 

Figure 6.8. Typical configuration for an active infrared-triggered camera system. Image based on Swann 

et al. (2004).

Table 6.2. Summary of infrared-triggered camera effort, Saguaro National Park, Rincon 

Mountain District, 1999–2005. See Appendix K for more complete summary. 

P= Number of camera nights

Location type Elevation stratum Number of cameras Sum Mean SD

Non-random Low 58 2162 37 40.4

Middle 5 200 40 33.6

High 11 294 27 18.8

Random Low 36 515 14 5.9

Middle 44 523 12 8.2

High 21 201 10 6.1
78

did not confirm the banner-tailed kangaroo rat 

(Dipodomys specatabilis), previously confirmed 

by specimen voucher (Hoffmeister 1986), and 

did not observe any of the distinctive sign of this 

very large kangaroo rat. Three species of bats 

that we did not observe, the western small-footed 

myotis, Yuma myotis, and western pipistrelle, 

have been confirmed recently (Davis and Sidner 

1992; Sidner 2003) and undoubtedly still occur 

at the district. One species of rodent (southern 

grasshopper mouse) is also present; a roadkilled 

individual found by Don Swann in 1997 was 

confirmed by Yar Petryzyn at the University of 

Arizona mammal collection. Four species are 

extirpated from the district (grizzly bear [Ursus 

arctos], jaguar [Panthera onca], Mexican gray 

wolf [Canis lupus], and bighorn sheep [Ovis 

canadensis]), and a fifth species (North American 

porcupine) may be extirpated, though it remains 

on the species list. 

Small Mammals

We trapped 544 individual rodents (including 

recaptures) in 2001 and 2002, and documented 

13 species through our trapping effort, as well as 

three species of diurnal squirrels (Table 6.3). One 

species, the fulvous harvest mouse (4 captures) 

was a new species for the district. We did not 

capture two species that have been previously 

documented for the district (the southern 

grasshopper mouse and banner-tailed kangaroo 

rat). 

Small mammal species richness was 

highest in the middle-elevation stratum (Table 

6.3), though sampling effort was also greater 

in that stratum. Therefore, after accounting for 

differences in sampling effort, species richness 

did not vary among strata (F2,35 = 0.16, P = 

0.86, one-way ANOVA, log-transformed data). 

Species richness was higher on non-random plots 

than on random plots in all strata (Table 6.3). 

At both high- and low-elevation strata, relative 

abundance of all rodents combined was higher 

on non-random plots than on random plots, but at 

middle elevations, relative abundance was higher 

on random plots (Table 6.3). In general, relative 

abundance was higher at both low and high 

elevations than at middle elevations. 

and non-random camera areas using one-way 

analysis of variance. Because cameras were open 

for differing lengths of time (Appendix K), we 

standardized effort for each camera by dividing 

observed species richness by the number of days 

that a camera was open. We then log-transformed 

these data to meet assumptions of normality. At 

random sites, we tested for differences in species 

richness among strata and type of camera (e.g., 

directly on random point, in nearest mapped 

drainage, and at site chosen by field personnel) 

using one-way analysis of variance.

Results

Species Richness

We confirmed a total of 59 species of mammals 

in the Rincon Mountain District (Appendix D). 

This included 12 species confirmed through 

specimens, 32 species confirmed through 

photographs, nine species captured for which 

a voucher specimen previously existed, five 

species confirmed through a combination of 

voucher specimens and photos, and one species 

confirmed through reliable observation. One 

species included in this total (eastern cottontail) 

was confirmed by photographs in appropriate 

high-elevation habitat, but requires further 

documentation. We confirmed three species 

of mammals not previously confirmed for the 

district: western red bat, fulvous harvest mouse, 

and Virginia opossum. The latter two species 

represent significant range extensions. We 

observed only one species listed by the U.S. Fish 

and Wildlife Service as endangered, the southern 

long-nosed bat. Three species of non-native 

animals were documented for the district (feral 

cat, domestic dog, and domestic cattle) but we 

do not believe that any of these species have 

established feral populations in the district. 

There have been a total of 66 species 

observed or documented in the district in the last 

few decades based on this and previous studies 

(Appendix D). We did not document the presence 

of 11 species that were previously documented 

for the Rincon Mountain District. We did not 

confirm the deer mouse, captured in the early 

1950s near Manning Camp (Appendix F). We 
79

Excluding the results for the whitethroated wood rat, whose identification may have 

been confused with the Mexican woodrat in some 

instances, there were important patterns among 

strata (Table 6.3). In particular, we trapped only 

one species (rock squirrel) in a single-elevation 

stratum, and only one species (brush mouse) in 

all three strata. The remainder of the species we 

found in two strata, either in the low- and middleor the middle- and high-elevation strata. We 

trapped no species solely in the middle-elevation 

stratum.

Bats

We confirmed 15 species, including one species 

that was not previously found at the district 

(western red bat; Table 6.4, Appendix D). We 

observed bats in only one roost site, where 500-

1000 cave myotis and six southern long-nosed 

bats were found. This was the only site at which 

we confirmed the southern long-nosed bat.

Lower Rincon Creek had the highest 

species richness of any site, and Manning Camp 

had the highest percent netting success and 

the most individuals captured (Table 6.4). We 

captured five species at Lower Rincon Creek 

that we did not capture in any other site and one 

species at Manning Camp Pond that we did not 

capture at any other site. At no other site did we 

capture species that were not found elsewhere. 

Wild Horse Canyon was the least productive 

site; we only caught one bat in three consecutive 

nights of trapping there. Three nights of netting 

were the most productive for species richness 

– two at Lower Rincon Creek and one at 

Manning Camp Pond – during this time we found 

seven species. There were extreme differences 

in the number of individuals caught and species 

richness within sites, particularly for Lower 

Rincon Creek and Manning Camp Pond, the two 

most sampled sites. At Lower Rincon Creek, the 

number of bats captured ranged from zero to 16 

and species richness ranged from zero to seven. 

Similar differences were observed for Manning 

Camp Pond. 

The big brown bat was the most 

widespread and abundant species; it was found 

at five of the six sites and in all elevation strata 

(Table 6.4). Big brown bats were captured in 

80% of the visits to Lower Rincon Creek and 

Manning Camp Pond. The Brazilian free-tailed 

bat was the next most-captured bat; we captured 

16 individuals at three sites. Of the 14 species 

that we captured at the Rincon Mountain District, 

10 were represented by four or fewer individuals. 

Low Middle High

Species R NR R NR R NR

rock squirrel 0.2

cliff chipmunk 0.2 2.2 3.5

Abert’s squirrel 0.2

Sonoran Desert pocket mouse 14.8

rock pocket mouse 3.9 5.4 2.5

Bailey’s pocket mouse 0.3 0.3 2.5

Merriam’s kangaroo rat 0.4 7.3

western harvest mouse 0.2 0.4

fulvous harvest mouse 1.4

cactus mouse 0.8 4.2 1.2 0.4

brush mouse 0.3 2.8 1.9 2.6 5.0 11.9

western white-throated woodrat 1.9 2.8 2.3 0.5 0.2 a 4.0 a

Mexican woodrat 0.2 1.7 1.3

yellow-nosed cotton rat 0.6 0.8

Arizona cotton rat 0.7 0.2

Species richness 5 7 7 9 5 8 a Identification at high elevations was not certain and further trapping is required to confirm this species. 

Table 6.3. Relative abundance of small mammals by strata and site type (R = random [focal-point 

transects]; NR = non-random), Saguaro National Park, Rincon Mountain District, 2001 and 2002. See 

Appendix I for summary of trapping effort. 
80

Even the cave myotis, for which we found a roost 

of >500 individuals, was represented by only a 

few individuals captured by netting. 

Medium and Large Mammals

In 3,895 estimated camera nights, 2,939 

photographs captured at least one mammal (not 

including nocturnal rodents, people, and horses 

with riders) and a total of 3,407 individual 

mammals that could be identified to genus. We 

photographed 27 species, including two nonnative species, domestic dog and cattle (Table 

6.5, Appendix D). We documented one species 

(Virginia opossum) not previously reported 

for the district and a large number of species 

for which there had previously been only 

observational records. 

The largest number of photographs 

was of the gray fox (1018 photos), followed 

by collared peccary (588 photos), and ringtail 

(229 photos). Species richness among elevation 

strata was highest in the low elevation (n = 24) 

and progressively lower through the elevation 

strata (n = 15, 13 at medium- and high-elevation 

stratum, respectively; Table 6.5), though effort 

was disproportionate in the low-elevation stratum 

(Table 6.2). After accounting for camera effort, 

there was no difference in species richness among 

strata (F2,170 = 2.0, P = 0.13, one-way ANOVA 

on log-transformed data), but random cameras 

did have slightly higher species richness than 

non-random camera sites (t

173 = 3.0, P = 0.003, 

two-tailed t-test). Among random sites where we 

placed three cameras, there were no differences 

in species richness among strata (F2,67 = 1.5, P = 

0.23, one-way ANOVA on log-transformed data), 

and within these sites there were no differences 

among the type of camera placement (at the 

focal-point transects; F2,67 = 1.1, P = 0.34, oneway ANOVA on log-transformed data). 

Pitfall Trapping

We trapped eight animals in pitfall traps: six 

desert shrews at the North Slope site, one western 

harvest mouse, and one Botta’s pocket gopher 

at Italian Spring. We trapped no animals at 

Spud Rock Spring. In this report we assume 

the desert shrews we captured during this study 

are Crawford’s desert shrew, but further genetic 

work would be necessary to confirm that it is this 

Table 6.4. Results of netting for bats, by elevation strata, site, and visit, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. 

Low Middle High

Chimenea 

Creek Lower Rincon Creek 

Wild Horse 

Canyon 

Deer 

Creek Manning Camp Pond 

Devil’s 

Bathtub

1 2 1 2 3 4 5 1 2 3 1 1 2 3 4 5 1

Mexican long-tongued bat 1

unknown myotis 1 1

southwestern myotis 2 1 1

cave myotis 1 1 1

fringed myotis 1 1

long-legged myotis 1 2

California myotis 2 1 1 3 1

silver-haired bat 1 2 1

big brown bat 4 4 2 2 2 2 5 10 19 4 1

western red bat 1

hoary bat 1 2 2 1 1

Townsend’s big-eared bat 1

pallid bat 1

Brazilian free-tailed bat 8 2 1 1 3 1

pocketed free-tailed bat 1 1

total detections by visit 2 4 16 4 0 8 9 0 1 0 7 1 12 17 21 5 3

total detections by site 6 37 1 7 56 3

percent netting success 3.4 7.1 0.4 7.4 19.1 11.1

species richness by site 2 12 1 3 7 3
81

No. photos

Relative

abundance No. photos

Relative 

abundance No. photos

Relative 

abundance

Virginia opossum 2 0.1

American black bear 2 0.1 21 2.9 10 2.0

white-nosed coati 17 0.6 8 1.1 3 0.6

ringtail 142 5.3 78 10.8 9 1.8

common raccoon 5 0.2

striped skunk 134 5.0 21 2.9 7 1.4

hooded skunk 160 6.0 20 2.8 2 0.4

white-backed hog-nosed skunk 20 0.7 4 0.6 3 0.6

western spotted skunk 3 0.1 3 0.4

coyote 97 3.6

domestic dog 2 0.1

common gray fox 602 22.6 283 39.3 133 27.0

mountain lion 46 1.7 16 2.2 11 2.2

bobcat 50 1.9 2 0.3 4 0.8

round-tailed ground squirrel 1 0.0

rock squirrel 13 0.5 3 0.4

Harris’ antelope squirrel 7 0.3

Abert’s squirrel 8 1.6

Arizona gray squirrel 2 0.3 1 0.2

antelope jackrabbit 7 0.3

black-tailed jackrabbit 10 0.4

desert cottontail 48 1.8 3 0.4

eastern cottontail 3 0.6

domestic cattle 3 0.1

collared peccary 561 21.0 27 3.8

mule deer 28 1.0

white-tailed deer 104 3.9 23 3.2 63 12.8

Total photographs 2064 0.81 514 0.73 257 0.53

Species richness 24 15 13

Table 6.5. Number of photographs of mammals from infrared-triggered photography by elevation strata, 

Saguaro National Park, Rincon Mountain District, 1999–2005. “Abundance” equals the number of photographs 

of that species per estimated number of working camera-nights. Does not include individuals that could be 

identified to genus but not species (e.g., some photos of deer, skunks, rabbits, and squirrels).

species and not Cockrum’s desert shrew; both 

species potentially occur in the Rincon Mountains 

(Baker et al. 2003b). 

Inventory Completeness

We confirmed a total of 59 species of mammals 

in the Rincon Mountain District and failed to 

confirm 11 species that have been previously 

documented for the Rincon Mountains. Of these 

11, four species (grizzly bear, jaguar, Mexican 

gray wolf, and bighorn sheep) are certainly 

extirpated from the district and two others 

(deer mice, North American porcupine, and 

banner-tailed kangaroo rat) may be extirpated. 

We believe that three species of bats and one 

rodent that were documented in the past are still 

present and would be confirmed with additional 

effort. Based on these records, if we assume that 

four species still present went undetected, our 

inventory confirmed 93% of mammals known for 

the district. The species accumulation curves for 

small mammal trapping (Fig. 6.9) and bats (Fig. 

6.10) as well as for infrared-triggered cameras 

(Fig. 6.11) also suggest that our inventory was 

fairly complete. These results make this effort 

one of the most comprehensive of its kind in 

the region for mammals. The infrared-triggered 

effort, in particular, is unprecedented. 

The three “new” species reported during 

this study may not have been observed before 

simply due to lack of survey effort. This situation 

 Low Middle High
82

Sampling period

0 2 4 6 8 10 12 14 16

Cumulative number of species

2

4

6

8

10

12

Low elevation 

Middle elevation 

High elevation 

Figure 6.9. Species accumulation curve for small-mammal trapping by elevation stratum, Saguaro 

National Park, Rincon Mountain District, 2001 and 2002. Each sampling period represents 10 observations 

(excluding recaptures).

Sampling period

0 2 4 6 8 10 12 14

Cumulative number of species

0

2

4

6

8

10

12

14

16

Figure 6.10. Species accumulation curve for bat trapping, Saguaro National Park, Rincon 

Mountain District, 2001 and 2002. Each sampling period represents one night of netting. 
83

seems unlikely in the case of the Virginia 

opossum, which has been extending its range 

northward; the record from this study represents 

a significant range extension (Babb et al. 2004). 

The red bat and fulvous harvest mouse were both 

found only along Rincon Creek, in the expansion 

area that was added to Saguaro National Park 

during the 1990s. The red bat is a riparian 

obligate species and may occur in the district only 

in this area. Less is known about the fulvous 

harvest mouse, but the finding of this species in 

the district is noteworthy. 

Additional inventory work could also 

increase the number of bat species detected. In 

total, 18 bat species have been confirmed for the 

Rincon Mountain District. Ronnie Sidner, who 

collected data for this effort and is a regional 

expert on the distribution and ecology of bats, 

believes that an additional four species could be 

found with additional survey effort: California 

Sample period

0 5 10 15 20 25 30

Cumulative number of species

5

10

15

20

25

Middle Elevation

High Elevation

Sample period

0 10 20 30 40

Cumulative number of species

5

10

15

20

25

Low elevation

Figure 6.11. Species accumulation curve for infrared-triggered cameras, Saguaro National Park, 

Rincon Mountain District, 1999-2005. Each sample period for the low-elevation stratum represents a 

randomized combination of 50 observations. Each sample period for medium- and high-elevation strata 

represents 20 observations. 
84

leaf-nosed bat (Macrotus californicus), western 

mastiff bat (Eumops perotis), little brown bat 

(Myotis lucifugus), and western yellow bat 

(Lasiurus xanthinus).

Discussion

Biogeography

As noted in the other chapters, the biology of 

the district reflects a fascinating geography. The 

district is located within two major watersheds, 

the Santa Cruz River on the west side of the 

Rincons and the San Pedro River on the east 

side. More importantly, the Rincon Mountains 

contain elements of several major biogeographic 

provinces, including the Sonoran Desert to the 

south and west, the Rocky Mountain region to 

the north and east, the Chihuahuan Desert to 

the east, and the Madrean region to the south. 

The Rincon Mountain District also hosts a 

significant elevational range, from 814 m (2,670 

ft) to 2,641 (8,665 ft), and a number of different 

plant communities. As a result, the district 

contains mammals that represent several different 

biogeographic origins, including a large number 

of species not present in the Tucson Mountain 

District. Thus, the Rincon Mountain District’s 

mammals include classic Sonoran Desert species 

(e.g., the round-tailed ground squirrel); species 

strongly associated with the Madrean region 

and central America (e.g., the white-nosed coati, 

collared peccary, and southern long-nosed bat); 

“northern” species (e.g., the American black bear 

and northern raccoon); and typically western 

species (e.g., the Botta’s pocket gopher and 

American badger).

A number of species, particularly bats 

and rodents, are on the edge of their range in 

the district. Our documentation of the fulvous 

harvest mouse is the furthest northwest location 

ever recorded for this species (Hoffmeister 

1986). Similarly, the Rincon-Catalina complex 

represents the northwestern-most site for the 

yellow-nosed cotton rat and the northeasternmost site for the Arizona pocket mouse. In 

contrast, several species we did not capture 

are found just southeast of the district in the 

Santa Rita Mountains and nearby sky island 

mountain ranges including the pygmy mouse 

(Baiomys taylori), fulvous cotton rat (Sigmodon 

fulviventor), hispid pocket mouse (Chaetodipus 

hispidus), and others (Hoffmeister 1986). It is 

possible that with more intensive effort these 

species might be found in the district. Indeed, 

Davis and Dunford (1987) suggest that the 

yellow-nosed cotton rat has only recently 

migrated into the Rincon Mountains. Lowe 

(1992) and Swann et al. (2005) have discussed 

the biogeography of reptiles and amphibians in 

the Rincon Mountains and factors that possibly 

influence distribution; it seems possible that these 

same patterns occur for smaller mammals as well. 

Habitat Associations

Despite its close proximity to Tucson, the district 

has had only a few mammal studies. Our study 

represents the first comprehensive inventory of 

the district below the high country (which was 

studied by Davis and Sidner 1992), and the first 

to quantify relative abundance and distribution 

of species. We trust that it will provide a good 

baseline for evaluating future changes in the 

mammal community at the district. 

Our study indicates that the Rincon 

Mountains have a typical assemblage of other 

sky island mountain ranges, with the exception 

of some semi-desert grassland species and 

the addition of a strong desert component. It 

is noteworthy that species richness for small 

mammals was similar between middle and high 

elevations. Overall, species richness was highest 

at the lowest elevations and decreased at higher 

elevations. There is a strong desert component 

in the mammalian community of the district, 

with a large number of species, ranging from 

the Sonoran Desert pocket mouse to mule deer, 

found only at lower elevations. However, a few 

species such as the Abert’s squirrel and Mexican 

woodrat were found only at high elevations. The 

middle elevations are richest overall, containing 

components of both deserts and forests. 

We did not attempt to separate riparian 

from upland species richness in this study. 

However, as would be expected, wet riparian 

areas at all elevations stand out as hotspots of 

mammal diversity. Davis and Sidner (1992) 
85

point out that the pond at Manning Camp has 

a remarkable diversity of bats. Davis and 

Sidner captured 12 species in just a few nights, 

compared to 12 species over many years of 

intensive netting at the Southwestern Research 

Station pond in Portal, Arizona, and nine species 

over many years at Quitobaquito Pond in Organ 

Pipe Cactus National Monument. Our netting 

results supported this statement; we recorded 

extraordinary species richness at both Manning 

Camp Pond and Rincon Creek (Table 6.4) 

Twenty-nine species of terrestrial mammals have 

now been documented at the Madrona Pools area 

of Chimenea Creek, and Sidner (2003) noted that 

a remarkable total of 17 species of bats have now 

been recorded along Chimenea Creek. 

Differences in habitat associations among 

species are similar to previous studies in the 

region. As in the Huachuca Mountains, the brush 

mouse is the most common small mammal in 

brushy and wooded vegetation above semi-desert 

grasslands (Hoffmeister and Goodpaster 1954). 

As previously described in collections made 

by Huey in 1932, Collins in 1954, and Davis 

and Sidner in 1984 and 1985 (Davis and Sidner 

1992), the brush mouse appears to be the only 

species of Peromyscus known to occur in the high 

country of the Rincon Mountains. However it is 

unclear if the deer mouse occurs in the Rincons. 

The yellow-nosed cotton rat was first documented 

in the district in 1984 in Manning Camp Meadow 

(Sidner and Davis 1994) and according to Davis 

and Dunford (1987) has recently colonized 

isolated montane grasslands in southern Arizona 

over the last 60 years. Although not previously 

known above 1,860 m (Cockrum 1960) or oak 

woodland (Hoffmeister 1986), these cotton rats 

now inhabit montane meadows in southeast 

Arizona where the longtail vole (Microtus 

longicaudus) is absent (Davis and Ward 1988). 

This is the case in the Rincon Mountains. 

We found the yellow-nosed cotton rat to be 

uncommon in montane meadows and adjacent 

pine forest in 2001; accurate assessment of their 

status would require a more focused multi-year 

study. The two lower-elevation records we 

obtained constitute the first documentation of 

their occurrence in more typical oak woodland/

grassland habitat.

The Mexican woodrat is perhaps more 

common in the Rincon Mountains than previously 

thought. Only four localities were previously 

known: Spud Rock Cabin, documented in 1932; 

Happy Valley Saddle, documented in 1968; 

Manning Camp Meadow, documented in 1984 

and 1985; and Spud Rock Summit documented in 

1985 (Davis and Sidner 1992). In 2001 we found 

this species near Mica Meadow, at and around 

Italian Spring, and east of Happy Valley Lookout. 

We found the western white-throated woodrat 

at all elevation strata, which is unusual because 

it is generally found below the conifer belt 

(Hoffmeister 1986). We believe this discrepancy 

may have been an artifact of poor identification 

by our field crews rather than a shift in habitat for 

this species.

Changes in the Mammal Community

Some of the patterns in distribution and 

abundance of mammals observed during this 

study contrast with historic records of mammals 

at the district. There is strong evidence that 

major changes have occurred in the mammal 

community of the district during the past seven 

decades, although lack of data precludes a full 

understanding of them. The greatest apparent 

changes since the park’s establishment include 

the extirpation of several large mammals, 

population increases for some other species, and 

significant changes in distribution of deer and 

(probably) some small mammals. Some of these 

changes are well-documented (e.g., we know a 

great deal about deer because of Sumner’s [1951] 

work and other records), but most others are not. 

The reasons for these changes are not at all clear, 

but there is some evidence for why they may 

have occurred. 

Of the extirpated species, the Mexican 

gray wolf and bighorn sheep appear to have been 

established at the time of the park’s creation 

(1933), though they were not common. In 

subsequent decades they slowly disappeared. 

The Mexican gray wolf was likely extirpated 

due to predator control programs, which were 

implemented throughout the southwestern United 

States. To its credit, the NPS made an effort to 
86

keep predator-control activities out of the district 

during the 1930s and 1940s, but it is possible 

that bounty hunters entered the district anyway 

(Saguaro NP, unpubl. records). Ironically, the 

effort to keep bounty hunters out of the district 

was led by Don Egermayer, the park custodian; 

but Egermayer himself shot a wolf on the X-9 

Ranch in 1947 (Saguaro NP, unpubl. records). 

Bighorn sheep occurred in the district 

through the 1940’s (Davis and Sidner 1992). A 

herd of 14 were observed south of Rincon Peak 

in 1942 (Coss 1969), and a weathered horn was 

collected on Tanque Verde Ridge in 1957. This 

species may have been eliminated by illegal 

hunting, although there may have been other 

factors as well. 

At least two, and probably five, jaguars 

were shot in the Rincon Mountains (in 1902, 

1912, 1920, and two in 1932) prior to the 

establishment of the park (Girmendonk 1994; 

Davis and Sidner 1992; specimen records 

in Appendix F), and there were occasional 

sightings of this species in the park’s early years. 

Currently, there are several jaguars known to 

be resident in southern Arizona, close to the 

Mexican border (Jack Childs, pers. comm.). We 

attempted to photograph jaguars during this 

study, placing cameras at high elevations along 

game trails where cat scat and scrapes were 

found, but obtained no photographs and found no 

evidence of this species. Although grizzly bears 

were once present in the Rincons, it is doubtful 

that any were present by the time the park 

was established; the last record for the Rincon 

Mountains was in 1921 (cited in Davis and Sidner 

1992). Both jaguars and grizzly bears would 

have been hunted aggressively well before the 

establishment of the park. 

The last known sighting of a North 

American porcupine was near Juniper Basin in 

the mid-1990s by District Ranger Bob Lineback. 

We made a concerted effort to search for this 

species during this study, but with no success. 

Porcupines appear to be declining throughout 

southern Arizona, possibly due to habitat 

changes, although Harley Shaw (pers. comm.) has 

suggested that it is due to the large increase in the 

population of mountain lions. 

While hunting and range-wide factors 

appear to be important in the loss of some 

species, significant changes in habitat for 

mammals at lower elevations, as well as habitat 

loss, may be responsible for other changes in the 

mammal community. Habitat changes include 

the large increase in shrubs and forbs since the 

cessation of grazing at the district. Active fire 

suppression and drought may have also played 

important roles in the increase of woody shrubs 

(Brown 1994, Bahre 1995, Van Auken 2000), 

particularly in the middle-elevation areas of the 

district. Habitat loss includes the significant 

loss of open space outside the district due 

to residential and commercial development, 

which has reduced low-lying desert habitat to a 

relatively thin strip along the west and south sides 

of the Rincon Mountains (see Chapter 2). 

Mule deer appear to be declining in 

the district for at least the past five decades. 

Sumner (1951) reported that mule deer were the 

dominant deer species below 6500’ in the Rincon 

Mountains, while white-tail deer occurred above 

7000’. Today, white-tail deer are commonly seen 

in the vicinity of the Cactus Forest Loop Drive 

(Don Swann, pers. obs.), and in this study mule 

deer were only photographed below 4000’ in 

elevation. Mule deer are declining throughout 

the western United States, and the cessation 

of cattle grazing at the district in the 1950s 

and 1960s has led to important changes in the 

vegetation community, such as growth of shrubs, 

that may favor the white-tail deer. The loss of 

mule deer habitat outside the district (due to 

increases in the adjacent housing developments) 

is probably also a major factor in their declining 

population at the district. Similarly, American 

badgers were sighted often in the early years 

of the park (Saguaro NP, unpubl. records), but 

were not photographed or collected during this 

study. Two reliable observations (one inside the 

Cactus Forest Loop Drive, and one on the district 

boundary near Freeman Road) and one recent 

photograph of a American badger by Ranger John 

Williams at the Wildhorse Gate on Speedway in 

March, 2006, indicate that this species still occurs 

in the district, but is definitely now rare. 
87

There is some suggestion that population 

declines have occurred, or are occurring, in the 

small mammal community. Kangaroo rats tend 

to prefer open-canopy areas with few shrubs, 

and were often mentioned in early accounts of 

Saguaro National Park. Today they are relatively 

uncommon and one species (the banner-tailed 

kangaroo rat) may be extirpated. The bannertailed kangaroo rat was present in low-elevation 

areas at some time prior to the mid-1980s 

(Hoffmeister 1986). Changes in the small 

mammal community might be expected to follow 

the significant changes in desert vegetation in 

the district that have occurred since the 1930s. 

Changes included a dramatic increase in shrubs 

and forbs following the cessation of grazing, 

as a well as reduction in the number of saguaro 

cacti. More obviously, there have certainly been 

changes in the status of the Arizona gray squirrel 

in the park since the introduction of the nonnative Abert’s squirrel. These changes are not yet 

well-understood, and the Arizona gray squirrel 

still occurs in the Rincons. However, ongoing 

research suggests that the Aberts’s squirrel is 

successfully established throughout the high 

country, even where Arizona gray squirrels occur, 

and that the native species is now uncommon 

(Koprowski 2006). 

It is noteworthy that three species 

(American black bear, mountain lion, and whitenosed coati) have exhibited the opposite trend 

and have increased in recent decades. Sumner 

(1951) noted that mountain lions and American 

black bears were absent during his wildlife 

survey in 1951, and did not mention the whitenosed coati. We believe that mountain lions 

and American black bears have increased due to 

decreases in hunting pressure outside the district 

as well as due to improvements in habitat inside 

the district. White-nosed coati may be moving 

northward and expanding their population size 

(Davis and Callahan 1992). However, this 

species is known to undergo dramatic population 

fluctuations (Chris Hass, pers. comm.). Because 

there are no records of coati prior to 1957, it is 

likely that they are new arrivals to the district 

(Davis and Sidner 1992). At any rate, the large 

number of photographs during our study, as well 

as a number of sightings of breeding groups, 

suggest that this species is doing very well at the 

present time. 

Management Implications and Additional 

Research Needed

Like many national parks (Newmark 1995, 

Powell et al. 2004), Saguaro National Park 

has seen the loss of mammal species since it 

was created in 1933. Our study indicates that 

these losses may be continuing at the district. 

We believe that the loss of habitat outside the 

district is the primary concern for large mammals 

at the present time. It seems that significant 

management efforts, with a proactive political 

effort outside the district, are needed to prevent 

the future extirpation of species like American 

badger and mule deer. Because the district is 

a relatively large natural area, it will provide 

habitat for many more species than will smaller 

areas, including the Tucson Mountain District.

While some species have declined or 

disappeared over the district’s history, many have 

increased. The park deserves credit for instituting 

land management practices that have improved 

habitat for many species. NPS policies, including 

cessation of cattle grazing, banning of hunting 

and trapping, restoration of natural fire regimes, 

elimination of off-road vehicles, and restriction 

of road-building have all helped to improve 

conditions for mammals and other wildlife at 

the district. In addition, while the lack of highprofile encounters between humans and mountain 

lions at the district has probably been a matter 

of good luck; the district’s few American black 

bear incidents are probably the result of good 

bear management policies, including installation 

of bear boxes in all campsites and diligent 

housekeeping at Manning Cabin. 

Future research should focus on 

learning more about those mammals for which 

very little data are available. Our inventory 

suggests that these species include the American 

badger, eastern cottontail, grassland rodents, 

pocket gophers, mule deer, and North American 

porcupine. With the exception of grassland 

rodents, all of these species may occur in low 
88

populations in the district and may be sensitive to 

future extirpation. We recommend a monitoring 

program for mule deer, a high-profile species; 

loss of this once-common species from a national 

park would be very unfortunate. We also 

recommend continued research on forest squirrels 

and increased research on small mammals. 

Pocket gophers, an ecologically significant group 

of animals at the park about which almost nothing 

is known, would also be an excellent candidate 

for additional research. 

Additional small trapping may increase 

the number of species documented in the Rincon 

Mountain District. The Rincon Mountains are 

a rugged and remote mountain range. Packing, 

setting, checking, and removing live-traps 

is difficult and time-consuming work. We 

believe that complete understanding of the 

genus Peromyscus (white-footed mice) in the 

Rincons remains elusive. We confirmed cactus 

mouse and brush mouse, but two deer mouse 

specimens exist from the park (Appendix F), 

and white-footed mice may also occur in the 

district based on records from nearby mountain 

ranges (Hoffmeister 1986, Lange 1960). In 

addition, mesquite mouse is also a possibility 

at lower elevations. Species in this genus are 

very difficult to distinguish in the field, and 

specimens (or genetic samples) are required. In 

addition, we failed to detect several semi-desert 

grassland rodents that have been recorded in 

nearby mountain ranges where better access 

facilitates more comprehensive surveys. Whether 

our failure to capture more semi-desert grassland 

species was due to insufficient effort or to 

interesting aspects of biogeography remains to be 

seen; there is evidence that many of these species 

simply do not occur in the Rincon Mountains. 

Nevertheless, we encourage the park to promote 

additional studies of small mammals in the 

district, particularly in the semi-desert grasslands 

at elevations between 4000 and 6000 feet. 

We also suggest that the park encourage 

visitors to the backcountry to report sightings of 

porcupines, which we believe may be extirpated 

from the district. Because porcupines are difficult 

to confuse with other species and because many 

park visitors now carry digital cameras, it would 

be prudent to enlist their assistance to report 

sightings of this species. We suggest posting 

requests for information at prominent trailheads 

or attaching such a request to each backcountry 

permit. 
89

Chapter 7: Literature Cited

Albrecht, E. W. 2001. Influence of microhabitat 

features on species richness and relative 

abundance in small mammal communities of 

Saguaro National Park, Arizona. Unpublished 

report to the University of Arizona Inventory 

and Monitoring Program, Tucson. Available in 

archive locations cited in Chapter 1. 

Allen, L. S. 1996. Ecological role of fire in the 

Madrean province. Pp. 5–10. In Effects of fire 

on Madrean province ecosystems. USDA Forest 

Service General Technical Report RM-GTR-289. 

Rocky Mountain Forest and Range Experiment 

Station, Ft. Collins, CO.

American Ornithologists’ Union (AOU). 1998. 

Checklist of North American birds, seventh 

edition. American Ornithologists’ Union and 

Allen Press Inc., Lawrence, KS.

American Ornithologists’ Union (AOU). 2003. 

Forty-second supplement to the American 

Ornithologists’ Union checklist of North 

American birds. Auk 117:847–858.

Anable, M. E., M. P. McClaran, and G. B. Ruyle. 

1992. Spread of introduced Lehmann’s lovegrass 

Eragrostis lehmanniana Nees. in southern 

Arizona, USA. Biological Conservation 61:181–

188.

Anderson, D. R. 2001a. The need to get the basics 

right in wildlife field studies. Wildlife Society 

Bulletin 29:1294–1297.

Anderson, G. 2001b. Rocking K Ranch and 

Expansion Area: vegetation surveys 2001.

Unpublished report to Saguaro National Park, 

Tucson.

Anderson, R., and K. Schon. 1999. Fire effects 

monitoring in Mexican spotted owl habitat in 

the Rincon Mountains of Saguaro National Park, 

Arizona. Pp. 7–8. In L. Benson and B. Gebow, 

editors. A century of parks in southern Arizona: 

second conference on research and resource 

management in southern Arizona national parks, 

extended abstracts. National Park Service, 

Southern Arizona Office and USGS Sonoran 

Desert Field Station, University of Arizona, 

Tucson.

Aslan, C. 2000. Mammalian predators in the Rincon 

Mountain District of Saguaro National Park: a 

high-elevation survey using remote photography. 

Unpublished report to University of Arizona 

Conservation Biology Internship Program and 

Saguaro National Park, Tucson, Arizona.

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. 

Bahre, C. J. 1995. A legacy of change: historic human 

impact on vegetation in the Arizona borderlands. 

University of Arizona Press, Tucson. 

Bailey, R. G. 1998. Ecoregions: The ecosystem 

geography of the oceans and continents. 

Springer-Verlag, New York, NY.

Bailey, S. J. 1994. Saguaro National Monument 

sensitive raptor surveys. Unpublished notes (for 

12 through 17 April 1994) to Saguaro National 

Park, Tucson. 

Baird, K. J., R. MacNish, and D. P. Guertin. 2000. An 

evaluation of hydrologic and riparian resources 

in Saguaro National Park, Tucson, Arizona. 

Department of Hydrology and Water Resources, 

Research Laboratory for Riparian Studies, 

University of Arizona, Tucson.

Baisan, C. H., and T. W. Swetnam. 1990. Fire history 

on a desert mountain range: Rincon Mountain 

Wilderness, Arizona, USA. Canadian Journal of 

Forestry Research 20:1559–1569.

Baker, R. J., L. C. Bradley, R. D. Bradley, J. W. 

Dragoo, M. D. Engstrom, R. S. Hoffmann, C. A. 

Jones, F. Reid, D. W. Rice, and C. Jones. 2003a. 

Revised checklist of North American mammals 

north of Mexico, 2003. Occasional Papers of the 

Museum of Texas Tech University 229:1–23.

Baker, R. J., M. B. O’Neill, and L. R. McAliley.

2003b. A new species of desert shrew, 

Notiosorex, based on nuclear and mitochondrial 

sequence data. Occasional papers, Museum of 

Texas Tech University 222:1–12.

Beauvais, G. P., and S. W. Buskirk. 1999. Modifying 

estimates of sampling effort to account for 

sprung traps. Wildlife Society Bulletin 27:39–

43. 

Berner, L. R., and R. W. Mannan. 1992. Survey for 

sensitive raptors in the Rincon Mountains of 

Saguaro National Monument, Arizona. Final 

report to Saguaro National Park, Tucson.

Bibby, C. J., N. D. Burgess, and D. A. Hill. 2002. 

Bird census techniques. Academic Press, 

London, England.

Black, K. L. 1982. An annotated checklist of the 

herpetofauna of Saguaro National Monument’s 

Rincon Mountain Unit. Unpublished report to 

Saguaro National Monument, Tucson, Arizona. 

Boal, C. W., and R. W. Mannan. 1996. Avian 

inventory. In L. Harris and W. Shaw, editors. 

Wildlife Inventory of the Rincon Valley. 
90

Unpublished report to Saguaro National Park, 

Tucson.

Bonham, C. D. 1989. Measurements for terrestrial 

vegetation. John Wiley and Sons Inc., New 

York, NY.

Bonine, K. E., and C. R. Schwalbe. 2003. 

Herpetofauna inventory of the Madrona Pools 

area. Pp. 44–61. In T. Edwards and D. E. 

Swann, editors. Madrona Pools “Pulse” study. 

Final report to Saguaro National Park, Tucson. 

Bowers, J. E. 1984. Woodland and forest flora and 

vegetation of Saguaro National Monument. 

Unpublished report to Saguaro National Park, 

Tucson. 

Bowers, J. E., and S. P. McLaughlin. 1987. Flora 

and vegetation of the Rincon Mountains, Pima 

County, Arizona. Desert Plants 8:50–95. 

Bowers, J. E., and 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.

Braun-Blanquet, J. 1965. Plant sociology: the study 

of plant communities. Hafner Inc., London, 

England.

Briggs, M. K., L. Harris, J. Howe, and W. Halvorson.

1996. Using long-term monitoring to understand 

how adjacent land development affects natural 

areas: an example from Saguaro National Park, 

Arizona (USA). Natural Areas Journal 16:354–

361.

Brown, D. E. 1994. Biotic communities: 

southwestern United States and northwestern 

Mexico. University of Utah Press, Salt Lake 

City, UT.

Bucci, M. 2001. Final report: small mammal 

trapping, Saguaro National Park Expansion Area 

and Rocking K Ranch. Unpublished report to 

Saguaro National Park, Tucson.

Buckland, S. T., D. R. Anderson, K. P. Burnham, J. L. 

Laake, D. L. Borchers, and L. Thomas. 2001. 

Introduction to distance sampling: Estimating 

abundance of biological populations. Oxford 

University Press, London, England. 

Cain, A. T., V. R. Touvila, D. G. Hewitt, and M. 

E. Tewes. 2003. Effects of a highway and 

mitigation projects on bobcats in southern Texas. 

Biological Conservation 114:189–197.

Carothers, S.W., R. R. Johnson, and S.W. Atchinson. 

1974. Population structure and social 

organization of southwestern riparian birds. 

American Zoologist 14:97–108.

Clark, B. K., B. S. Clark, L. A. Johnson, and M. T. 

Haynie. 2001. Influence of roads on movements 

of small mammals. Southwestern Naturalist 

46:338–344.

Clarke, A. L., and T. Pacin. 2002. Domestic cat 

“colonies” in natural areas: a growing exotic 

species threat. Natural Areas Journal 22:154–

159.

Clarkson, R. W., and J. C. Rorabaugh. 1989. Status 

of leopard frogs (Rana pipiens complex: 

Ranidae) in Arizona and southeastern California. 

Southwestern Naturalist 34:531–538.

Cockrum, E. L. 1960. The recent mammals of 

Arizona: their taxonomy and distribution. 

University of Arizona Press, Tucson. 

Cody, M. L. 1981. Habitat selection in birds: the 

roles of vegetation structure, competitors, and 

productivity. BioScience 31:170–113.

Cole, C. J., and H. C. Dessauer. 1994. Unisexual 

lizards (genus Cnemidophorus) of the Madrean 

Archipelago. Pp. 267–273. In Biodiversity and 

management of the Madrean Archipelago: the 

Sky Islands of Southwestern United States and 

northwestern Mexico. General Technical Report 

RM-GTR-264. USDA Forest Service, Rocky 

Mountain Research Station, Ft. Collins, CO. 

Colver, K. J., D. Stokes, and L. Stokes. 1999. Stokes 

field guide to bird songs. Time Warner, New 

York, NY.

Conway, C. J., and C. Kirkpatrick. 2001. Population 

status, detection probability, and effects of fire 

on buff-breasted flycatchers. Final report to the 

Arizona Game and Fish Department, Phoenix, 

AZ. 

Corman, T. E., and R. T. Magill. 2000. Western 

yellow-billed cuckoo in Arizona: 1998 and 1999 

survey report. Technical report 150. Arizona 

Game and Fish Department, Nongame and 

endangered wildlife program. Phoenix, AZ. 

Corman, T. E., and C. Wise-Gervais, editors. 2005. 

Arizona breeding bird atlas. University of New 

Mexico Press, Albuquerque, MN.

Coss, H. 1969. Rincon Mountain bighorn. 

Unpublished report in Saguaro National Park 

files, Tucson. 

Crump, M. L., and N. J. Scott. 1994. Visual 

encounter surveys. Pp. 84–92. In W. R. Heyer, 

M. A. Donnelly, R. W. McDiarmid, L. C. 

Hayek, and M. S. Foster, editors. Measuring 

and monitoring biodiversity: Standard methods 

for amphibians. Smithsonian Institution Press, 

Washington, D.C.

Davis, R., and J. R. Callahan. 1992. Post-Pleistocene 

dispersal in the Mexican vole (Mictrotis 

mexicanus): an example of an apparent trend in 

the distribution of southwestern mammals. Great 

Basin Naturalist 52:262–268. 
91

Davis, R., and C. Dunford. 1987. An example of 

contemporary colonization by small, nonflying 

mammals of montane islands of the American 

Southwest. American Naturalist 129:398–406. 

 Davis, K., and W. L. Halvorson. 2000. A study plan 

to inventory vascular plants and vertebrates: 

Sonoran Desert Network. National Park Service, 

Southern Arizona Office, Phoenix, AZ.

Davis, R., and R. Sidner. 1992. Mammals of 

woodland and forest habitats in the Rincon 

Mountains of Saguaro National Monument, 

Arizona. Technical Report 47. Cooperative 

Park Resources Study Unit, School of Natural 

Resources, University of Arizona, Tucson. 

Davis, R., and O. G. Ward. 1988. The vacant 

“Microtis niche” now occupied by the yellownosed cotton rat (Sigmodon ochrognathus) on 

an isolated mountain in southeastern Arizona. 

Journal of Mammalogy 69:362–365.

Degenhardt, W. G., C. W. Painter, and A. H. Price. 

1996. Amphibians and reptiles of New Mexico. 

University of New Mexico Press, Albuquerque, 

NM.

Dimmitt, M. A. 2000. Biomes and communities of 

the Sonoran Desert region. In S. J. Phillips and 

P. W. Comus, editors. A natural history of the 

Sonoran Desert. Arizona-Sonora Desert Museum 

Press, Tucson.

Doll, M. H., R. V. Ellis, R. L. Hayes, R. Sidner, H. 

McCrystal, R. Davis, and R. L. Hall. 1986. A 

checklist of the herpetofauna and mammals of 

Saguaro National Monument. Southwest Parks 

and Monuments Association, Tucson.

Drewes, H. 1977. Geologic map and sections of 

Rincon Valley Quadrangle, Pima County, 

Arizona. U.S. Geological Survey, Miscellaneous 

Investigations Series Map I-997. 

Duncan, D. K. 1990. Nocturnal rodent populations 

and associated vegetation with implications 

of human use at Saguaro National Monument, 

Arizona. Technical Report 35. Cooperative 

National Park Service Resources Study Unit, 

School of Natural Resources, University of 

Arizona, Tucson.

Edwards, T., C. R. Schwalbe, D. E. Swann, and C. S. 

Goldberg. 2004. Implications of anthropogenic 

landscape change on inter-population movements 

of the desert tortoise (Gopherus agassizii). 

Conservation Genetics 5:485–499. 

Elzinga, C. L., D. W. Salzer, J. W. Willoughby, and J. 

P. Gibbs. 2001. Monitoring plant and animal 

populations. Blackwell Science, Malden, MA. 

Esque, T.C., A. M. Búrquez, C. R. Schwalbe, T. R. 

Van Devender, M. J. M. Nijhuis, and P. Anning. 

2003. Effects of fire on desert tortoises and their 

habitats. In T. R. Van Devender, editor. The 

Sonoran Desert tortoise: natural history, biology 

and conservation. Arizona-Sonora Desert 

Museum Press, Tucson.

Felley, D. L., and T. E. Corman. 1993. Spring 

1993 cactus ferruginous pygmy-owl surveys in 

Arizona. Unpublished report. Nongame and 

endangered wildlife program, Arizona Game and 

Fish Department, Phoenix, AZ.

Fishbein, M. 1995. Flora of Madrona Canyon. 

Unpublished report to Saguaro National Park, 

Tucson.

Fishbein, M., and J. E. Bowers. 1996. Additions and 

corrections to the Flora of the Rincon Mountains, 

Pima County, Arizona. Unpublished report to 

Saguaro National Park, Tucson. 

Fishbein, M., S. McMahon, G. Ferguson, V. 

Steinmann, and A. Johnson. 1994a. Flora of 

Chimenea Canyon, Saguaro National Monument, 

East Unit. Unpublished report to Saguaro 

National Park, Tucson.

Fishbein, M., V. Steinmann and A. Johnson. 

1994b. Floristic Survey of the proposed Box 

Canyon Protected Natural Area: Final report. 

Unpublished report to Saguaro National Park, 

Tucson.

Fitzgerald, C. S. 1996. Large mammal inventory. 

Pp. 20–23. In L. K. Harris, editor. Wildlife 

inventory of the Saguaro National Park 

expansion area. Unpublished report to Saguaro 

National Park, Tucson. 

Flesch, A. D. 2001. Distribution, relative abundance, 

and microhabitat associations of small mammals 

in Saguaro National Park, southeast Arizona. 

Unpublished report to the University of Arizona 

Inventory and Monitoring Program, Tucson. 

Flesch, A. D, D. E. Swann, and B. F. Powell. 2006. 

Amphibians and reptiles of Saguaro National 

Park, Tucson Mountain District. In B. F. Powell, 

C. A. Schmidt, and W. L. Halvorson, editors. 

Plant and vertebrate inventory of Saguaro 

National Park, Tucson Mountain District. 

Unpublished report to Sonoran Desert Network 

I&M Program, Tucson. 

Frederici, P. 1998. An evaluation of avian resources 

and habitats in the Saguaro National Park 

expansion area, Rincon Mountain District. 

Unpublished report to Saguaro National Park, 

Tucson. 

Frohn, R. C. 1998. Remote sensing for landscape 

ecology: new metric indicators for monitoring, 

modeling and assessment of ecosystems. Lewis 

Publishers, Boca Raton, FL. 
92

Fuller, M. R., and J. A. Mosher. 1987. Raptor 

survey techniques. Pp. 37–66. In B. A. GeronPendleton, B. A. Millsap, K. W. Cline, and D. M. 

Bird, editors. Raptor management techniques 

manual. National Wildlife Federation, 

Washington, D.C.

Funicelli, C. S., P. J. Anning, and D. S. Turner. 2001. 

Long-term vegetation monitoring at Saguaro 

National Park: A decade of change. Technical 

Report No. 70. USGS Sonoran Desert Research 

Station, University of Arizona, Tucson.

Germaine, S. S., S. S. Rosenstock, R. E. 

Schweinsburg, and W. S. Richardson. 1998. 

Relationships among breeding birds, habitat, 

and residential development in greater Tucson, 

Arizona. Ecological Applications 8:680–691.

Girmendonk, A. L. 1994. Ocelot, jaguar, and 

jaguarundi sighting reports: Arizona and Sonora, 

Mexico. Unpublished report, Arizona Game 

and Fish Department, Nongame and Endangered 

Wildlife Program, Phoenix, AZ.

Goad, M. S., and R. W. Mannan. 1987. Nest site 

selection by elf owls in Saguaro National 

Monument, Arizona. Condor 89:659–662.

Goldberg, C., D. E. Swann, and J. E. Wallace. 2004. 

Genetic structure of lowland leopard frog 

populations at Saguaro National Park. Final 

report to Western National Parks Association and 

Saguaro National Park, Tucson. 

Goode, M. J., D. Turner, and S. Breeden. 1998. 

Herpetofauna of Saguaro National Park 

expansion area: Habitat assessment, inventory, 

threats and management recommendations. 

Unpublished report to Saguaro National Park, 

Tucson. 

Griscom, H. 2000. Saguaro National Park, Arizona, 

sensitive raptor survey, 2000. Unpublished 

report to Saguaro National Park, Tucson.

Guertin, P. P. 1998. A vegetation and plant survey of 

the newly added lands of Saguaro National Park, 

Rincon Mountain District. Unpublished report to 

Saguaro National Park, Tucson.

Halvorson, W. L., and B. S. Gebow, editors. 2000. 

Floristic surveys of Saguaro National Park 

protected natural areas. Technical Report No. 

68. USGS Sonoran Desert Research Station, 

University of Arizona, Tucson. 

Halvorson, W. L., and P. P. Guertin. 2003. USGS 

Weeds in the West project: status of introduced 

plants in southern Arizona Parks. USGS, 

Sonoran Desert Research Station, University of 

Arizona, Tucson.

Hayek, L. C., and M. A. Buzas. 1997. Surveying 

natural populations. Columbia University Press, 

New York, NY. 

Heritage Data Management System (HDMS). 2004. 

Arizona Game and Fish Department. Accessed 

5 March from: http://www.gf.state.az.us/w_c/

edits/hdms_species_lists.html.

Hoffmeister, D. F. 1956. Mammals of the Graham 

(Pinaleño) Mountains, Arizona. American 

Midland Naturalist 55:257–288.

Hoffmeister, D. F. 1986. Mammals of Arizona. 

University of Arizona Press and Arizona Game 

and Fish Department, Tucson. 

Hoffmeister, D. F., and W. W. Goodpaster. 1954. 

The mammals of the Huachuca Mountains, 

southeastern Arizona. Illinois Biological 

Monograph 24. 

Integrated Taxonomic Information System (ITIS). 

2005. Accessed on 20 March 2005 from: http://

www.itis.usda.gov/index.html. 

Ivanyi, C., J. Perry, T. R. Van Devender, and H. 

Lawler. 2000. Reptile and amphibian accounts. 

Pp. 533–585. In S. J. Phillips and P. W. Comus, 

editors. A natural history of the Sonoran Desert. 

Arizona-Sonora Desert Museum Press, Tucson. 

James, F. C. 1971. Ordinations of habitat 

relationships among breeding birds. Wilson 

Bulletin 83:215–236.

Johnson, R. R., and L. T. Haight. 1991. The 

importance of xeroriparian ecosystems to 

the wintering avifauna of Saguaro National 

Monument. Pp. 105–110. In Proceedings of 

the symposium on research in Saguaro National 

Monument. Cooperative Park Studies Unit, 

University of Arizona, Tucson.

Johnson, W. T. 1988. Flora of the Pinaleño 

Mountains, Graham County, Arizona. Desert 

Plants 8:147–162.

Jones, C. A., C. R. Schwalbe, D. E. Swann, D. P. 

Prival, and W. W. Shaw. 2005. Mycoplasma 

agassizii in desert tortoises. Final report to the 

Arizona Game and Fish Department Heritage 

Grant Project #U03005. Phoenix, AZ. 

Keddy-Hector, D. P. 1998. Aplomado falcon (Falco 

femoralis). In R. Glinski, editor. Raptors of 

Arizona. University of Arizona Press, Tucson. 

Kennedy, K. A., and P. A. Addison. 1987. Some 

considerations for the use of visual estimates of 

plant cover in biomonitoring. Journal of Ecology 

75:151-157. 

Kerpez, T. A., and N. S. Smith. 1990. Competition 

between European starlings and native 

woodpeckers for nest cavities in Saguaros. Auk 

107:367–375.

Kirkpatrick, C., and C. J. Conway. In prep. Effects 

of fire on montane forest birds in southeastern 

Arizona. 
93

Kirkpatrick, C., C. J. Conway, and P. B. Jones. 2006. 

Distribution and relative abundance of forest 

birds in relation to burn severity in southeastern 

Arizona. Journal of Wildlife Management 70: 

1009-1012.

Kline, N. C., and D. E. Swann. 1998. Quantifying 

wildlife mortality on roads in Saguaro National 

Park. Pp. 23–31. In G. L. Evink, P. Garrett, 

D. Zeigler, and J. Berry, editors. Proceedings 

of conference on transportation and wildlife 

ecology, Fort Meyers, FL, February 10–12 1998.

Koprowski, J. 2006. Ecology of introduced Abert’s 

squirrels. Final report to Western National Parks 

Assocation and Saguaro National Park, Tucson, 

AZ. 

Kotliar, N. B., and J. A. Weins. 1990. Multiple scales 

of patchiness and patch structure: a hierarchical 

framework for the study of heterogeneity. Oikos 

59:253–260.

Krebs, C. J. 1998. Ecological methodology. Second 

edition. Addison-Welsey Educational, Menlo 

Park, CA.

Kucera, T. E., and R. H. Barrett. 1993. The 

Trailmaster camera system for detecting wildlife. 

Wildlife Society Bulletin 21:505–508.

Lange, K. I. 1960. Mammals of the Santa Catalina 

Mountains, Arizona. American Midland 

Naturalist 64:436–438.

Latta, M. J., C. J. Beardmore, and T. E. Corman. 

1999. Arizona Partners in Flight conservation 

plan. Technical Report 142. Nongame and 

Endangered Wildlife Program, Arizona Game 

and Fish Department, Phoenix, AZ.

Levy, P. S., and S. Lemeshow. 1999. Sampling of 

populations: methods and applications. John 

Wiley and Sons, New York, NY.

Lowe, C. H. 1992. On the biogeography of the 

herpetofauna of Saguaro National Monument. 

Pp. 91–104. In Proceedings of the symposium 

on research in Saguaro National Monument. 

Cooperative Park Studies Unit, University of 

Arizona, Tucson. 

Lowe, C. H. 1994. An overview of the herpetofauna 

of the Sonoran Desert. In: Herpetology of 

the North American deserts: Proceedings of 

a symposium. Southwestern Herpetologists 

Society, Special Publication No. 5:71-78. 

Lowe, C. H., and P. A. Holm. 1991. The amphibians 

and reptiles at Saguaro National Monument, 

Arizona. Technical Report No. 37. Cooperative 

National Park Resources Studies Unit, University 

of Arizona, Tucson. 

Lynn, S. 1996. Large mammal survey. Pp. 29-40. 

In L. K. Harris and C. R. Schwalbe, editors. 

Wildlife inventory of the Rincon Valley. Arizona 

Game and Fish Heritage Project U93007, 

Phoenix, AZ. 

MacArthur, R. H., and J. W. MacArthur. 1961. On 

bird species diversity. Ecology 42:594–598.

Mannan, R. W., and B. Bibles. 1989. Impacts of 

exotic cavity-nesting birds on native cavitynesting birds in Saguaro National Monument. 

Unpublished report to Saguaro National Park, 

Tucson.

Marshall, J. T., Jr. 1956. Summer birds in the Rincon 

Mountains, Saguaro National Monument, 

Arizona. Condor 58:81–97.

Mau-Crimmins, T., A. Hubbard, D. Angell, C. 

Filipone, and N. Kline. 2005. Sonoran Desert 

Network vital signs monitoring plan. Technical 

report NPS/IMR/SODN-003. National Park 

Service. Denver, CO.

McAuliffe, J. R. 1993. Case study of research, 

monitoring, and management programs 

associated with the saguaro cactus (Carnegia 

gigantea) at Saguaro National Monument, 

Arizona. Technical report NPS/WRUA/NRTR93/01. National Park Service, Cooperative 

National Park Resources Studies Unit, University 

of Arizona, Tucson. 

M’Closkey, R. T. 1980. Spatial patterns in sizes of 

seeds collected by four species of heteromyid 

rodents. Ecology 61:486–489. 

McColly, R. A. 1961. Geology of the Saguaro 

National Monument Area, Pima County, Arizona. 

Masters Thesis, Department of Geology, 

University of Arizona, Tucson.

McLaughlin, S. P. 1986. A floristic analysis of 

the southwestern United States. Great Basin 

Naturalist 46:46–55.

McLaughlin, S. P. 1993. Additions to the flora of the 

Pinaleño Mountains, Graham County, Arizona. 

Journal of the Arizona-Nevada Academy of 

Science 27:5–32. 

McLaughlin, S. P., and M. P. McClaran. 2004. Recent 

additions to the flora of the Pinaleño Mountains, 

Graham County, Arizona. Journal of the 

Arizona-Nevada Academy of Science 37:91–93.

McPherson, G. R. 1993. Effects of herbivory and 

herb interference on oak establishment in a semiarid temperate savanna. Journal of Vegetation 

Science 4:687–692.

Mills, G. S., J. B. Dunning Jr., and J. M. Bates. 

1989. Effects of urbanization of breeding bird 

community structure in southwestern desert 

habitats. Condor 91:416–428. 

Monson, G., and S. Smith. 1985. Bird checklist for 

Saguaro National Monument. Southwest Parks 

and Monument Association, Tucson. 
94

Mott, D. N. 1997. Saguaro National Park, Arizona 

Water Resources Report. Unpublished report by 

the National Park Service, Fort Collins, CO.

Murray, R. C. 1996. Amphibian and reptile inventory. 

Pp. 31–46. In Wildlife inventory of the Saguaro 

National Park expansion area. L. K. Harris, 

editor. Unpublished report to Saguaro National 

Park, Tucson. 

National Park Service (NPS). 1992a. NPS-75: 

Natural resources inventory and monitoring 

guidelines. U.S. Dept. of Interior, Washington, 

D.C.

National Park Service (NPS). 1992b. Update to the 

natural/cultural resources management plan. 

Saguaro National Monument, Tucson. 

National Park Service (NPS). 2005. NPS visitation 

database reports. Accessed on 09 January 2005 

from:. http://www2.nature.nps.gov/NPstats/

npstats.cfm.

Nelson, L., and F. W. Clark. 1973. Correction 

for sprung traps in catch/effort calculations 

of trapping results. Journal of Mammology 

54:295–298. 

Newmark, W. D. 1995. Extinction of mammal 

populations in western North-American national 

parks. Conservation Biology 9:512–526.

North American Ornithological Atlas Committee 

(NAOAC). 1990. Handbook for atlasing 

North American breeding birds. C. Smith, 

editor. Accessed 13 July 2001 from: http://

americanbirding.org/norac/atlascont.htm.

Phillips, A., J. Marshall, and G. Monson. 1964. The 

birds of Arizona. The University of Arizona Press, 

Tucson.

Pima Association of Governments (PAG). 2005. 

Population estimates 1980–2004. Accessed 

on Aug 22, 2005 from: http://www.pagnet.org/

population/data/Est1980-2004.htm.

Pima County Flood Control District (PCFCD). 2005. 

Unpublished data from Manning Camp ALERT 

rain gauge. Accessed 23 June 2005 from: http://

rfcd.pima.gov/alertsys/.

Pollock. K. H., J. D. Nichols, T. R. Simons, G. L. 

Farnsworth, L. L. Bailey, and J. R. Sauer. 

2002. Large scale wildlife monitoring studies: 

statistical methods for design and analysis. 

Environmetrics 13:105-119. 

Powell, B. F. 1999. Birds of the Rocking K 

Development and Saguaro National Park. 

Unpublished report to Saguaro National Park, 

Tucson. 

Powell, B. F. 2004. Assessment of the bird 

community along the Middle Reach of Rincon 

Creek, Saguaro National Park. Unpublished 

report to Saguaro National Park, Tucson. 

Powell, B. F., E. W. Albrecht, W. L. Halvorson, 

and K. Docherty. 2003. Biological inventory 

report for the Sonoran Desert Network: 

2002. Annual Report No. 2. Sonoran Desert 

Network Inventory Program. USGS, Sonoran 

Desert Research Station and School of Natural 

Resources, University of Arizona, Tucson. 

Powell, B. F., E. W. Albrecht, W. L. Halvorson, C. 

A. Schmidt, P. Anning, and K. Docherty. 2005. 

Vascular plant and vertebrate inventory of 

Tumacácori National Historical Park. USGS 

Open File Report 2005-1142. U.S.Geological 

Survey, Southwest Biological Sciences Center, 

Sonoran Desert Research Station, University of 

Arizona, Tucson. 

Powell, B. F., E. W. Albrecht, W. L. Halvorson, C. A. 

Schmidt, and K. Docherty. 2004. Vascular plant 

and vertebrate inventory of Casa Grande Ruins 

National Monument. Final report to the National 

Park Service, Sonoran Desert Inventory and 

Monitoring Program, Tucson.

Powell, B. F., K. Docherty, and W. L. Halvorson. 

2002. Biological inventory report for the 

Sonoran Desert Network: 2000 and 2001 field 

seasons. Annual Report No. 1. Sonoran Desert 

Network Inventory Program. USGS, Sonoran 

Desert Research Station and School of Natural 

Resources, University of Arizona, Tucson.

Pyne, S. J. 1996. Nouvelle southwest. Pp. 10–16. 

In W. Covington, and P. K. Wagner, editors. 

Conference on adaptive ecosystems restoration 

and management: restoration of cordilleran 

conifer landscapes of North America. General 

Technical Report RM-GTR-278. USDA Forest 

Service, Rocky Mountain Forest and Range 

Experiment Station, Ft. Collins, CO.

Ralph, C. J., J. R. Sauer, and S. Droege, technical 

editors. 1995. Monitoring bird populations 

by point counts. General Technical Report 

PSW-GTR-149. USDA Forest Service, Pacific 

Southwest Research Station, Albany, CA.

Ramsey, F. L., and D. W. Schafer. 2002. The 

statistical sleuth: a course in methods of data 

analysis. 2nd Edition. Duxbury, Inc., Pacific 

Grove, CA.

Rappole, J. H. 1995. The ecology of migrant birds: a 

neotropical perspective. Smithsonian Institution 

Press, Washington, D.C. 

Reynolds, R. T., J. M. Scott, and R. A. Nussbaum. 

1980. A variable circular-plot method for 

estimating bird numbers. Condor 82:309–313.
95

Rice, J., B. W. Anderson, and R. D. Ohmart. 1984. 

Comparison of the importance of different habitat 

attributes to avian community organization. 

Journal of Wildlife Management 48:895–911.

Rondeau, R., and R. Van Devender. 1992. Floristic 

survey of the proposed Wildhorse Canyon 

Protected Natural Area. Unpublished report to 

Saguaro National Park, Tucson.

Roseberry, R. D., and N. E. Dole, Jr. 1939. The 

vegetation type survey of Saguaro National 

Monument. Unpublished report. 

Rosen, P. C. 2000. A monitoring study of vertebrate 

community ecology in the northern Sonoran 

Desert, Arizona. Unpublished PhD dissertation, 

Department of Ecology and Evolutionary 

Biology, University of Arizona, Tucson.

Rosen, P. C., and C. H. Lowe. 1995. Lizard 

monitoring protocol for the ecological 

monitoring program in Organ Pipe Cactus 

National Monument, Arizona. Special Report 11, 

Section 4. Cooperative Ecosystems Study Unit, 

University of Arizona, Tucson. 

Rosenberg, K. V., R. D. Ohmart, W. C. Hunter, and 

B. W. Anderson. 1991. Birds of the Lower 

Colorado River Valley. University of Arizona 

Press, Tucson.

Roth, R. L. 1976. Spatial heterogeneity and bird 

species diversity. Ecology 57:773–782. 

Russell, S., and G. Monson. 1998. The birds of 

Sonora. University of Arizona Press, Tucson.

Saguaro National Park (Saguaro NP). 2005. Sonoran 

upland vegetation monitoring in Saguaro 

National Park, 1999–2005. In-park report.

Schwalbe, C. R., T. C. Esque, M. J. Nijhuis, D. F. 

Haines, J. W. Clark, and P. J. Swantek. 1999. 

Effects of fire on Arizona Upland Desertscrub 

at Saguaro National Park. Pp. 107–109. In L. 

Benson and B. Gebow, editors. A century of 

parks in southern Arizona: second conference on 

research and resource management in southern 

Arizona national parks, extended abstracts. 

National Park Service, Southern Arizona Office 

and USGS Sonoran Desert Research Station, 

University of Arizona, Tucson.

Shaffer, H. B., and J. E. Juterbock. 1994. Night 

driving. Pp. 163–166. In W. R. Heyer, M. A. 

Donnelly, R. W. McDiarmid, L. C. Hayek, and 

M. S. Foster, editors. Measuring and monitoring 

biodiversity: standard methods for amphibians. 

Smithsonian Institution Press, Washington, D.C.

Shmida, A. 1984. Whittaker’s plant diversity 

sampling method. Israel Journal of Botany 

33:41–46.

Short, K. C. 1996. Results of the Arizona Breeding 

Bird Atlas effort in Saguaro National Park. 

Unpublished report to Saguaro National Park, 

Tucson.

Short, K. C. 2002. The effects of low-severity 

prescribed fires on vegetation, arthropods, and 

breeding birds in southwestern ponderosa pine 

forest. PhD. Dissertation and technical report, 

University of Montana, Missoula, MT. 

Shreve, F. 1951. Vegetation of the Sonoran Desert. 

Carnegie Institution of Washington Publication 

no. 591. Washington, D.C.

Sidner, R. 1991. Survey for the endangered lesser 

long-nosed bat (Leptonycteris curasoae) on 

Saguaro National Monument: May–September 

1991. Unpublished report to Saguaro National 

Park, Tuscon.

Sidner, R. 2003. Bat diversity at the Madrona Pools 

area. Pp 74–83. In T. Edwards and D. E. 

Swann, editors. Madrona Pools PULSE study. 

Unpublished report, Saguaro National Park, 

Tucson.

Sidner, R., and R. Davis. 1994. Bats of Saguaro 

National Monument, 1992–1994. Unpublished 

report to Saguaro National Park, Tucson. 

Sprouse, T., R. Emanuel, and B. Tellman. 2002. 

Final report: surface water quality monitoring 

overview and assessment for the Sonoran Desert 

Network, National Park Service. Unpublished 

report. Water Resources Research Center, 

University of Arizona, Tucson. 

Sredl, M. J., J. M. Howland, J. E. Wallace, and L. 

S. Saylor. 1997. Status and distribution of 

Arizona’s native Ranid frogs. Pp. 37–89. In

M. J. Sredl, editor. Ranid frog conservation 

and management. Technical Report no. 121, 

Nongame and Endangered Wildlife Program, 

Arizona Game and Fish Department, Phoenix, 

AZ.

Stebbins, R. C. 2003. A field guide to western reptiles 

and amphibians. Third edition. Houghton 

Mifflin, New York, NY.

Steenbergh, W. F., and C. H. Lowe. 1977. Ecology 

of the saguaro: II reproduction, germination, 

establishment, growth, and survival of the young 

plant. National Park Service Monograph Series 

No. 8, Washington, DC. 

Steidl, R. J. 2003. Monitoring elf owls in and near 

Saguaro National Park, 2000–2002. Unpublished 

report to the Western National Parks Association 

and Saguaro National Park, Tucson. 

Steidl, R. J., and A. Knipps. 1999. Effects of fire on 

Mexican spotted owls in Saguaro National Park. 
96

Unpublished progress report to Saguaro National 

Park, Tucson.

Stein, B. A. 2002. States of the union: Ranking 

America’s biodiversity. Natureserve, Arlington, 

VA.

Stitt, E. W., C. R. Schwalbe, D. E. Swann, R. C. 

Averill-Murray, and A. K. Blythe. 2003.

Sonoran Desert tortoise ecology and 

management: effects of land use change and 

urbanization on desert tortoises. Final report to 

Saguaro National Park.

Stohlgren, T. J., M. B. Falkner, and L. D. Schell. 

1995a. A modified-Whittaker nested vegetation 

sampling method. Vegetatio 117:113–121.

Stohlgren, T. J., J. F. Quinn, M. Ruggiero, and G. S. 

Waggoner. 1995b. Status of biotic inventories 

in U.S. national parks. Biological Conservation 

71:97–106. 

Strong, T. R., and C. E. Bock. 1990. Bird species 

distribution in riparian habitats in southeastern 

Arizona. Condor 92:866–885.

Stutchbury, B. J. 1991. Coloniality and breeding 

biology of purple martins (Progne subis 

hesperia) in saguaro cactus. Condor 93:666–675.

Sumner, L. 1951. A biological survey of Saguaro 

National Monument. Unpublished report to 

Saguaro National Park, Tucson.

Swann, D. E. 1997. Status of leopard frogs (Genus 

Rana) in the Rincon Mountain Unit of Saguaro 

National Park: Results of surveys, summer 1996. 

Draft final report to Saguaro National Park, 

Tucson. 

Swann, D. E. 1999a. Evaluating approaches for 

monitoring terrestrial vertebrates in U.S. 

national parks: an example from Tonto National 

Monument. MS Thesis, School of Natural 

Resources, University of Arizona, Tucson. 

Swann, D. E. 1999b. Surveys of reptiles and 

amphibians, Rocking K and Saguaro National 

Park Expansion Area. Unpublished report to 

Saguaro National Park, Tucson. 

Swann, D. E. 2003. Mammal inventory of the 

Madrona Pools area. Pp. 88–102. In T. Edwards 

and D. E. Swann, editors. Madrona Pools 

PULSE study. Unpublished report, Saguaro 

National Park, Tucson.

Swann. D. E. 2004. A checklist of herpetofauna and 

mammals of Saguaro National Park, Arizona. 

Western National Parks Association, Tucson. 

Swann, D. E., R. C. Averill-Murray, and C. R. 

Schwalbe. 2002. Distance sampling for Sonoran 

desert tortoises. Journal of Wildlife Management 

66:943–949. 

Swann, D. E., V. Gempko, and B. F. Powell. 2003a. 

Special surveys for lowland leopard frogs, 

mammals, and plants on the remote east slope 

of the Rincon Mountains. Unpublished report 

to the Sonoran Desert Inventory and Monitoring 

Program, Saguaro National Park, and Desert 

Southwest Cooperative Ecosystems Study Unit, 

Tucson.

Swann, D. E., M. J. Goode, and C. R. Schwalbe. 

2000. Inventory and recommendations for longterm monitoring of reptiles and amphibians at 

Fort Bowie National Historic Site, Arizona. 

Report to the National Park Service, Southern 

Arizona Office, Phoenix, AZ. 

Swann, D. E., C. C. Hass, D. C. Dalton, and S. A. Wolf. 

2004. Infrared-triggered cameras for detecting 

wildlife: an evaluation and review. Wildlife 

Society Bulletin 32:1–9. 

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. 

Pp. 149-153. In G. J. Gottfied, B. S. Gebow, 

L. G. Eskew, and C. B. Edminster, editors. 

Connecting mountain islands and desert seas: 

biodiversity and management of the Madrean 

Archipelago II. 2004 May 11-15; Tucson, AZ. 

Proceedings RMRS-P-36. USDA Forest Service, 

Rocky Mountain Research Station, Fort Collins, 

CO. 

Swann, D. E., K. Schon, and R. E. Wallace. 2003b. 

Effects of wildland fire on lowland leopard 

frogs at Saguaro National Park, Arizona. Poster 

presentation to American Geomorphological 

Society Annual Meeting, Denver, CO., October 

2003. 

Swann, D. E., and C. R. Schwalbe. 2001. Inventory 

and recommendations for long-term monitoring 

of reptiles and amphibians at Coronado National 

Memorial, Arizona. Unpublished report to 

National Park Service, Southern Arizona Office, 

Phoenix, AZ.

Swantek, P. J., W. L. Halvorson, and C. R. Schwalbe. 

1999. GIS database development to analyze 

fire history in southern Arizona and beyond, an 

example from Saguaro National Park. Technical 

report no. 61. U.S. Geological Survey, Arizona 

National Park Resources Unit, University of 

Arizona, Tucson, AZ. 

Swetnam, T. W., C. D. Allen, and J. Betancourt. 1999. 

Applied historical ecology: using the past to 

manage the future. Ecological Applications 

9:1189–1206.

Swetnam, T. W., and C. H. Baisan. 1996. Fire 

histories of montane forests in Madrean 
97

borderlands. Pp. 15–16. In effects of fire 

on Madrean province ecosystems. General 

Technical Report RM-GTR-289. USDA Forest 

Service, Rocky Mountain Forest and Range 

Experiment Station, Ft. Collins, CO.

Taylor, R. C. 1997. Location checklist to the birds 

of the Chiricahua Mountains. Borderland 

Productions, Tucson, AZ.

Trombulak, S. C., and C. A. Frissell. 2000. Review 

of ecological effects of roads on terrestrial and 

aquatic communities. Conservation Biology 

14:18–30.

Tucson Bird Count (TBC). 2005. Accessed from web 

on August 4, 2005: http://www.tucsonbirds.

org/index.html.

Turner, D. S., and C. S. Funicelli. 2000. Ten-year 

resurvey of epidermal browning and population 

structure of saguaro cactus (Carnegiea gigantea) 

in Saguaro National Park. Technical Report No. 

69. USGS, Sonoran Desert Research Station, 

University of Arizona, Tucson.

Turner, D. S., P. A. Holm, E. B. Wirt, and C. R. 

Schwalbe. 2003. Amphibians and reptiles 

of the Whetstone Mountains, Arizona. The 

Southwestern Naturalist 48:347–355. 

United States Department of Agriculture (USDA). 

2005. The PLANTS Database, Version 3.5. 

National Plant Data Center, Natural Resources 

Conservation Service, Baton Rouge, LA. 

Accessed on 10 March 2003 from: http://plants.

usda.gov. 

Van Auken, O. W. 2000. Shrub invasions of North 

American semiarid grasslands. Annual Review 

of Ecology and Systematics 31:197–215.

Verner, J., and L. V. Ritter. 1983. A comparison of 

transects and point counts in oak-pine woodlands 

of California. Condor 87:47–68.

Warshall, P. 1994. The biopolitics of the Mt. 

Graham red squirrel (Tamiasciuris hudsonicus 

grahamensis). Conservation Biology 8:977–988.

Western Regional Climate Center (WRCC). 2005. 

Arizona climate summaries for the University of 

Arizona. Accessed on 15 February 2005 from: 

http://www.wrcc.dri.edu/summary/climsmaz.

html.

White, G. C., K. P. Anderson, K. P. Burnham, and D. 

L. Otis. 1983. Capture-recapture and removal 

methods for sampling closed populations. Los 

Alamos National Laboratory, Los Alamos, NM.

Whitmore, R. C. 1975. Habitat ordination of breeding 

passerine birds of the Virgin River valley, 

southwestern Utah. Wilson Bulletin 87:65–74. 

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.

Willey, D. W. 1997. Life history attributes of 

Mexican spotted owls (Strix occidentalis lucida) 

in Saguaro National Park. Unpublished report to 

Saguaro National Park, Tucson.

Willey, D. W. 1998a. Effects of fire on Mexican 

spotted owls (Strix occidentalis lucida) at 

Saguaro National Park. Arizona Game and Fish 

Department Heritage Project #I96029 Final 

Report. Phoenix, AZ.

Willey, D. W. 1998b. Monitoring Mexican spotted 

owl occupancy and reproductive status in 

Saguaro National Park, 1998 field season final 

report. Unpublished report to Saguaro National 

Park, Tucson.

Wirt, E. B, and R. H. Robichaux. 2000. Survey 

and monitoring of the desert tortoise Gopherus 

agassizii at Saguaro National Park. Unpublished 

report to Saguaro National Park, Tucson.

Wolf, S. A., and D. E. Swann. 2002. Infraredtriggered photography of mammals in the Rincon 

Mountain District of Saguaro National Park, 

Arizona. Final report to Western National Parks 

Association and Saguaro National Park, Tucson. 

Wright, J. W., and L. J. Vitt. 1993. Biology of 

the whiptail lizards (genus Cnemidophorus). 

Oklahoma Museum of Natural History, Norman, 

OK. 
98
99

Appendix A. List of plant species that were observed (O) or collected (X) at Saguaro National Park, Rincon Mountain District. Species list derived from species seen 

or collected by UA Inventory personnel from this study (UAI), specimens located in the University of Arizona herbarium (from 1909–1996; UAH), Bowers and McLaughlin (1987; 

B&M), Rondeau and Van Devender (1992; R&D), Fishbein et al. (1994a; FIa), Fishbein et al. (1994b; FIb), Fishbein (1995; FI), Fishbein and Bowers (1996; F&B), Guertin (1998; 

GU), Halvorson and Guertin (2003; H&G), long-term monitoring plots 1998–2004 (SNP in prep; LTM), fire-effects monitoring (Saguaro National Park, unpublished data; FEM). 

Species in bold-faced type are non-native (from USDA 2004).

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Acanthaceae Anisacanthus thurberi (Torr.) Gray Thurber’s desert honeysuckle X X X O O O O O

Carlowrightia arizonica Gray Arizona wrightwort X X X O O O O

Elytraria imbricata (Vahl) Pers. purple scalystem O

Justicia candicans (Nees) L. Benson Arizona water-willow O

Ruellia nudiflora (Engelm. & Gray) Urban violet wild petunia X O

Siphonoglossa longiflora (Torr.) Gray longflower tubetongue X X X O O O O O

Tetramerium nervosum Nees hairy fournwort X X X O O X

Aceraceae Acer glabrum Torr. Rocky Mountain maple X

Acer glabrum var. neomexicanum (Greene) Kearney & Peebles New Mexico maple X

Acer negundo L. boxelder X

Acer negundo var. interius (Britt.) Sarg. boxelder X

Agavaceae Agave chrysantha Peebles goldenflower century plant O X O O

Agave palmeri Engelm. Palmer’s century plant X X O O O O O O

Agave parryi Engelm. Parry’s agave O

Agave schottii Engelm. Schott’s century plant X O O

Agave schottii Engelm. var. schottii Schott’s century plant X O O O

Yucca baccata var. brevifolia (Schott ex Torr.) L. Benson & Darrow Spanish dagger X O

Yucca elata (Engelm.) Engelm. soaptree yucca O

Yucca elata (Engelm.) Engelm. var. elata soaptree yucca X O

Yucca schottii Engelm. Schott’s yucca O X X O O O O

Aizoaceae Trianthema portulacastrum L. desert horsepurslane X X

Amaranthaceae Amaranthus albus L. prostrate pigweed X X O O

Amaranthus blitoides S. Wats. mat amaranth X

Amaranthus fimbriatus (Torr.) Benth. ex S. Wats. fringed amaranth X X O

Amaranthus palmeri S. Wats. carelessweed X X O O X O

Amaranthus powellii S. Wats. Powell’s amaranth X X

Froelichia arizonica Thornb. ex Standl. Arizona snakecotton X X X O

Gomphrena caespitosa Torr. tufted globe amaranth X X

Gomphrena nitida Rothrock pearly globe amaranth X X X O

Gomphrena sonorae Torr. Sonoran globe amaranth X X X O O O

Guilleminea densa (Humb. & Bonpl. ex Willd.) Moq. small matweed X X

Iresine heterophylla Standl. Standley’s bloodleaf X X O O O

Tidestromia lanuginosa (Nutt.) Standl. woolly tidestromia X O

Anacardiaceae Rhus aromatica Ait. fragrant sumac X

Rhus lancea L. African Sumac X

Rhus trilobata Nutt. skunkbush sumac O O O

Rhus trilobata var. pilosissima Engelm. pubescent squawbush X O

Rhus trilobata var. racemulosa (Greene) Barkl. skunkbush sumac X X
100

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Anacardiaceae Rhus virens var. choriophylla (Woot. & Standl.) L. Benson evergreen sumacXX

Toxicodendron radicans ssp. divaricatum (Greene) Gillis eastern poison ivyX

Toxicodendron rydbergii (Small ex Rydb.) Greene western poison ivyXOO

Apiaceae Bowlesia incana Ruiz & Pavón hoary bowlesiaXXXOOX

Daucus pusillus Michx. American wild carrotXXXOOOOOO

Lomatium nevadense (S. Wats.) Coult. & Rose Nevada biscuitrootXO

Lomatium nevadense (S. Wats.) Coult. & var. nevadense Nevada biscuitrootXX

Pseudocymopterus montanus (Gray) Coult. & Rose alpine false springparsleyOXXO

Spermolepis echinata (Nutt. ex DC.) Heller bristly scaleseedXXXOOOOO

Yabea microcarpa (Hook. & Arn.) K.-Pol. false carrotXXO

Apocynaceae Apocynum androsaemifolium L. spreading dogbaneXXO

Apocynum cannabinum L. IndianhempXXX

Haplophyton crooksii (L. Benson) L. Benson cockroachplantXXOOOOO

Macrosiphonia brachysiphon (Torr.) Gray Huachuca Mountain rocktrumpetX

Nerium oleander L. oleanderOX

Araliaceae Aralia humilis Cav. Arizona spikenardXXOO

Aristolochiaceae Aristolochia watsonii Woot. & Standl. Watson’s dutchman’s pipeOX

XOOOOO

Asclepiadaceae Asclepias angustifolia Schweig. Arizona milkweedXX

Asclepias asperula (Dcne.) Woods. ssp. asperula spider milkweedX

Asclepias glaucescens Kunth nodding milkweedXXO

Asclepias hypoleuca (Gray) Woods. mahogany milkweedXX

Asclepias linaria Cav. pineneedle milkweedOXXOOOO

Asclepias nyctaginifolia Gray Mojave milkweedXXXOO

Asclepias quinquedentata Gray slimpod milkweedXX

Asclepias tuberosa L. butterfly milkweedX

Asclepias tuberosa ssp. interior Woods. butterfly milkweedXO

Cynanchum arizonicum (Gray) Shinners Arizona swallow-wortXXXOOO

Funastrum crispum (Benth.) Schlechter wavyleaf twinevineX

Funastrum cynanchoides ssp. cynanchoides (Dcne.) Schlechter fringed twinevineXO

Funastrum cynanchoides ssp. heterophyllum (Vail) Kartesz, comb. 

nov. ined. Hartweg’s twinevine

XXXOOOO

Matelea arizonica (Gray) Shinners Arizona milkvineXXOOOO

Matelea parvifolia (Torr.) Woods. spearleafXXOX

Matelea producta (Torr.) Woods. Texas milkvineX

Aspleniaceae Asplenium trichomanes L. maidenhair spleenwortXX

Asteraceae Achillea millefolium L. common yarrow

X

Achillea millefolium var. occidentalis DC. western yarrowXO

Acourtia nana (Gray) Reveal & King dwarf desertpeonyXXO

OO

Acourtia thurberi (Gray) Reveal & King Thurber’s desertpeonyXXOOO

Acourtia wrightii (Gray) Reveal & King brownfootOXXOOOO

Adenophyllum porophylloides (Gray) Strother San Felipe dogweedOXXOO

Ageratina herbacea (Gray) King & H.E. Robins. fragrant snakerootOXX

Ageratina paupercula (Gray) King & H.E. Robins. Santa Rita snakerootXXXOO

Ageratina rothrockii (Gray) King & H.E. Robins. Rothrock’s snakerootXX
101

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Asteraceae Ambrosia ambrosioides (Cav.) Payne ambrosia leaf burr ragweedXXXOOOOOO

Ambrosia confertiflora DC. weakleaf burr ragweedXXXOOOOO

Ambrosia cordifolia (Gray) Payne Tucson burr ragweedOO

Ambrosia deltoidea (Torr.) Payne triangle burr ragweedXX

Ambrosia dumosa (Gray) Payne burrobushXXO

Ambrosia psilostachya DC. Cuman ragweedOO

Anaphalis DC. pearly everlastingO

Antennaria marginata Greene whitemargin pussytoesXO

Antennaria parvifolia Nutt. small-leaf pussytoesXO

Antheropeas lanosum (Gray) Rydb. white easterbonnetsXXOOX

Artemisia dracunculus L. tarragonXO

Artemisia dracunculus ssp. dracunculus L. wormwoodXX

Artemisia ludoviciana Nutt. white sagebrushXXOOOO

Artemisia ludoviciana ssp. albula (Woot.) Keck white sagebrushXOO

Artemisia ludoviciana ssp. sulcata (Rydb.) Keck white sagebrushX

Baccharis brachyphylla Gray shortleaf baccharisXXOOX

Baccharis pteronioides DC. yerba de pasmoXX

Baccharis salicifolia (Ruiz & Pavón) Pers. mule’s fatXXXOOOO

OO

Baccharis sarothroides Gray desertbroomXXOOOOOOO

Baccharis thesioides Kunth Arizona baccharisXXXOOOO

Bahia absinthifolia Benth. hairyseed bahiaOXOOO

Bahia absinthifolia var. dealbata (Gray) Gray Dealbata’s bahiaXOO

Bahia dissecta (Gray) Britt. ragleaf bahiaXXXOO

Baileya multiradiata Harvey & Gray ex Gray desert marigoldOXXOOO

Bebbia juncea (Benth.) Greene sweetbushXOOO

Bebbia juncea var. aspera Greene sweetbushX

Bidens aurea (Ait.) Sherff Arizona beggarticksXXXOOO

Bidens heterosperma Gray Rocky Mountain beggarticksXXXO

Bidens lemmonii Gray Lemmon’s beggarticksXXX

Bidens leptocephala Sherff fewflower beggarticksXXXO

Brickellia amplexicaulis B.L. Robins. earleaf brickellbushXXOOO

Brickellia baccharidea Gray resinleaf brickellbushXX

Brickellia betonicifolia Gray betonyleaf brickellbushXXXOOOO

Brickellia californica (Torr. & Gray) Gray California brickellbushXXXOOOOO

Brickellia coulteri Gray Coulter’s brickellbushXX

XOOOOX

Brickellia eupatorioides var. chlorolepis (Woot. & Standl.) B.L. Turnerfalse bonesetXXX

Brickellia grandiflora (Hook.) Nutt. tasselflower brickellbushOXXO

Brickellia pringlei Gray Pringle’s brickellbushXX

Brickellia rusbyi Gray stinking brickellbushXX

Brickellia venosa (Woot. & Standl.) B.L. Robins. veiny brickellbushXXOOOOO

Brickelliastrum fendleri (Gray) King & H.E. Robins. Fendler’s brickellbushXO

Calycoseris wrightii Gray white tackstemOXO

Carminatia tenuiflora DC. plumeweedXXXOOOO

Carphochaete bigelovii Gray Bigelow’s bristleheadXXXOO
102

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Asteraceae Centaurea melitensis L. Maltese star-thistle

XXXOXO

Chaenactis stevioides Hook. & Arn. Steve’s dustymaidenXX

Chaetopappa ericoides (Torr.) Nesom rose heathX

Cirsium neomexicanum Gray New Mexico thistleOXXOOOOX

Cirsium undulatum (Nutt.) Spreng. wavyleaf thistleOXXO

Cirsium wheeleri (Gray) Petrak Wheeler’s thistleXXO

Conyza bonariensis (L.) Cronq. asthmaweedXX

Conyza canadensis (L.) Cronq. Canadian horseweedOXOOOOO

Conyza canadensis (L.) Cronq. var. canadensis Canadian horseweedX

Coreocarpus arizonicus (Gray) Blake little lemonheadXXXOOOO

Cosmos parviflorus (Jacq.) Pers. southwestern cosmosOXXOO

Dimorphotheca sinuata DC. glandular cape marigoldOOX

Encelia farinosa Gray ex Torr. goldenhillsOXOOOOOOO

Encelia farinosa Gray ex Torr. var. farinosa goldenhillsX

Encelia frutescens (Gray) Gray var. frutescens button brittlebushXX

Ericameria cuneata (Gray) McClatchie cliff goldenbushX

Ericameria cuneata var. spathulata (Gray) Hall cliff goldenbush

X

Ericameria laricifolia (Gray) Shinners turpentine bushXXXOOOOOXO

Erigeron colomexicanus A. Nels. running fleabaneXXOOO

Erigeron divergens Torr. & Gray spreading fleabaneXXXOOOOXO

Erigeron flagellaris Gray trailing fleabaneXO

Erigeron neomexicanus Gray New Mexico fleabaneXXOOOO

Erigeron oreophilus Greenm. chaparral fleabaneXXXOOO

Erigeron speciosus var. macranthus (Nutt.) Cronq. aspen fleabaneXX

Eryngium L. eryngoX

Evax verna var. verna Raf. spring pygmycudweedXX

Filago arizonica Gray Arizona cottonroseXX

Filago californica Nutt. California cottonroseXXXOXO

Filago depressa Gray dwarf cottonroseXO

Fleischmannia pycnocephala (Less.) King & H.E. Robins. lavender thoroughwortXX

Galinsoga parviflora Cav. gallant-soldierXX

Gamochaeta purpurea (L.) Cabrera spoonleaf purple everlastingXXXOO

Gnaphalium palustre Nutt. western marsh cudweedO

Guardiola platyphylla Gray Apache plantXXXOOOO

Gutierrezia arizonica (Gray) M.A. Lane Arizona snakeweedXX

Gutierrezia microcephala (DC.) Gray threadleaf snakeweedXXXO

O

Gutierrezia sarothrae (Pursh) Britt. & Rusby broom snakeweedXOX

Gutierrezia serotina Greene late snakeweedXX

Gymnosperma glutinosum (Spreng.) Less. gumheadXXXOOOO

Helenium thurberi Gray Thurber’s sneezeweedXXX

Helianthella quinquenervis (Hook.) Gray fivenerve helianthellaXX

Helianthus annuus L. common sunflowerXX

Heliomeris longifolia var. annua (M.E. Jones) Yates longleaf false goldeneyeXXOOOOX

Heliomeris multiflora var. multiflora Nutt. showy goldeneyeO
103

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Asteraceae Heliomeris multiflora var. nevadensis (A. Nels.) Yates Nevada goldeneyeXXX

Heterosperma pinnatum Cav. wingpetalXXO

Heterotheca fulcrata (Greene) Shinners rockyscree false goldenasterX

Heterotheca fulcrata (Greene) Shinners var. fulcrata rockyscree false goldenasterX

Heterotheca subaxillaris (Lam.) Britt. & Rusby camphorweedXXXOOOXX

Hieracium carneum Greene Huachuca hawkweedXXXO

Hieracium fendleri Schultz-Bip. yellow hawkweedXOO

Hieracium fendleri var. discolor Gray yellow hawkweedX

Hieracium lemmonii Gray Lemmon’s hawkweedO

Hymenoclea monogyra Torr. & Gray ex Gray singlewhorl burrobrushOOOX

Hymenoclea salsola Torr. & Gray ex Gray burrobrushXXX

Hymenopappus mexicanus Gray Mexican woollywhiteXO

Hymenothrix wislizeni Gray TransPecos thimbleheadXX

Hymenothrix wrightii Gray Wright’s thimbleheadXXXOOOOO

Hymenoxys hoopesii (Gray) Bierner owl’s-clawsOXXO

Isocoma coronopifolia (Gray) Greene common goldenbushX

Isocoma tenuisecta Greene burroweedOXXOOOOOO

Koanophyllon solidaginifolium (Gray) King & H.E. Robins. shrubby thoroughwortXXXOO

OOO

Lactuca serriola L. prickly lettuceXXXOOOX

Laennecia coulteri (Gray) Nesom conyzaXXXO

Laennecia eriophylla (Gray) Nesom X

Laennecia schiedeana (Less.) Nesom pineland marshtailXXO

Laennecia sophiifolia (Kunth) Nesom leafy marshtailXXO

Lasianthaea podocephala (Gray) K. Becker San Pedro daisyXXXO

Lasthenia californica DC. ex Lindl. California goldfieldsXXOO

Leibnitzia lyrata (D. Don) Nesom Seeman’s sunbonnetsXX

Machaeranthera arida B.L. Turner & Horne arid tansyasterO

Machaeranthera asteroides var. asteroides (Torr.) Greene New Mexico tansyasterXX

Machaeranthera canescens var. incana (Lindl.) Gray hoary tansyasterXOO

Machaeranthera gracilis (Nutt.) Shinners slender goldenweedOXXOOOO

Machaeranthera pinnatifida (Hook.) Shinners lacy tansyasterXOO

Machaeranthera pinnatifida var. pinnatifida (Hook.) Shinners lacy tansyasterXXOO

Machaeranthera tagetina Greene mesa tansyasterXXXOOOOX

Malacothrix clevelandii Gray Cleveland’s desertdandelionXXXOO

Malacothrix fendleri Gray Fendler’s desertdandelionXO

Malacothrix glabrata (Gray ex D.C. Eat.) Gray smooth desertdandelionX

Malacothrix stebbinsii W.S. Davis & Raven Stebbins’ desertdandelionO

Melampodium longicorne Gray Arizona blackfootOO

Monoptilon bellioides (Gray) Hall Mojave desertstarO

Packera neomexicana var. neomexicana (Gray) W.A. Weber & A. 

Löve New Mexico groundsel

XXXOOO

Parthenice mollis Gray annual monsterwortXXXO

Parthenium incanum Kunth mariolaOXXOOO

Pectis cylindrica (Fern.) Rydb. Sonoran cinchweedOO
104

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Asteraceae Pectis filipes Harvey & Gray fivebract cinchweedXX

Pectis filipes var. subnuda Fern. fivebract cinchweedXO

Pectis longipes Gray longstalk cinchweedXO

Pectis papposa Harvey & Gray manybristle cinchweedX

Pectis papposa Harvey & Gray var. papposa manybristle cinchweedX

Pectis prostrata Cav. spreading cinchweedXX

Perityle coronopifolia Gray crowfoot rockdaisyXX

Perityle lemmonii (Gray) J.F. Macbr. Lemmon’s rockdaisyXXXOOO

Peucephyllum schottii Gray Schott’s pygmycedarO

Porophyllum gracile Benth. slender poreleafXXXOOOOO

Porophyllum ruderale (Jacq.) Cass. yerba porosaO

Porophyllum ruderale ssp. macrocephalum (DC.) R.R. Johnson yerba porosaXXXOO

Pseudognaphalium canescens ssp. canescens (DC.) W.A. Weber Wright’s cudweedOXXOOOOXO

Pseudognaphalium leucocephalum (Gray) A. Anderb. white cudweedXXXOOX

Pseudognaphalium macounii (Greene) Kartesz, comb. nov. ined. Macoun’s cudweedX

Pseudognaphalium pringlei (Gray) A. Anderb. Pringle’s cudweedX

Pseudognaphalium stramineum (Kunth) W.A. Weber cottonbatting plantXXXO

Pseudognaphalium viscosum (Kunth) W.A. Weber winged cudweedXO

Psilactis asteroides Gray New Mexico tansyasterXOO

Psilostrophe cooperi (Gray) Greene whitestem paperflowerOXXOOOOO

Rafinesquia californica Nutt. California plumseedXX

Rafinesquia neomexicana Gray New Mexico plumseedXXOOO

Rudbeckia laciniata L. cutleaf coneflowerXX

Sanvitalia abertii Gray Albert’s creeping zinniaXXOOOO

Senecio bigelovii Gray nodding ragwortXO

Senecio bigelovii Gray var. bigelovii nodding ragwortX

Senecio flaccidus var. douglasii (DC.) B.L. Turner & T.M. Barkl. Douglas’ ragwortXXO

Senecio flaccidus var. monoensis (Greene) B.L. Turner & T.M. Barkl. Mono ragwortXXO

Senecio lemmonii Gray Lemmon’s ragwortXXXOOOX

Senecio wootonii Greene Wooton’s ragwortXXO

Solidago canadensis var. scabra Torr. & Gray 

a Canada goldenrod

X

Solidago missouriensis Nutt. Missouri goldenrodXXXO

Solidago rugosa P. Mill. var. rugosa wrinkleleaf goldenrodX

Solidago velutina DC. threenerve goldenrodXXXOOO

Solidago wrightii Gray Wright’s goldenrodXXO

Solidago wrightii var. wrightii Gray Wright’s goldenrodX

Sonchus asper (L.) Hill spiny sowthistleXXXOOO

Sonchus oleraceus L. common sowthistleXXXOOOOO

Stephanomeria pauciflora (Torr.) A. Nels. brownplume wirelettuce

XXXOOOOOO

Stevia lemmonii (Gray) Gray Lemmon’s candyleafXX

Stevia plummerae Gray Plummer’s candyleafXX

Stevia serrata Cav. sawtooth candyleafXXO

Stylocline micropoides Gray woollyhead neststrawXXOX

Symphyotrichum falcatum var. commutatum (Torr. & Gray) Nesom white prairie asterX
105

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

AsteraceaeSymphyotrichum praealtum var. praealtum (Poir.) Nesom willowleaf asterX

Tagetes lemmonii Gray Lemmon’s marigoldXXXOOO

Tagetes micrantha Cav. licorice marigoldXXXOOO

Taraxacum laevigatum (Willd.) DC. rock dandelionXXX

Taraxacum officinale G.H. Weber ex Wiggers common dandelionO

Thymophylla pentachaeta var. belenidium (DC.) Strother fiveneedle pricklyleafX

Thymophylla pentachaeta var. pentachaeta (DC.) Small fiveneedle pricklyleafXXOOO

Trixis californica Kellogg American threefoldOXXOOOOOO

Uropappus lindleyi (DC.) Nutt. Lindley’s silverpuffsXXXOOOOO

Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray golden crownbeardX

Verbesina encelioides ssp. exauriculata (Robins. & Greenm.) J.R. 

Coleman golden crownbeard

X

Viguiera cordifolia Gray heartleaf goldeneyeXXXO

Viguiera deltoidea Gray Parish’s goldeneyeX

Viguiera dentata var. lancifolia Blake toothleaf goldeneyeXXOOOO

Xanthium strumarium L. rough cockleburrXXOOXO

Xanthium strumarium var. canadense (P. Mill.) Torr. & Gray Canada cockleburrXX

Zinnia acerosa (DC.) Gray desert zinniaXXXO

OOOOX

Berberidaceae Berberis wilcoxii Kearney Wilcox’s barberryXXXO

Betulaceae Alnus incana ssp. tenuifolia (Nutt.) Breitung 

a thinleaf alder

X

Alnus oblongifolia Torr. Arizona alderXXOO

Bignoniaceae Chilopsis linearis (Cav.) Sweet desert willowXX

Tecoma stans (L.) Juss. ex Kunth yellow trumpetbushXXOO

Bixaceae Amoreuxia palmatifida Moc. & Sessé ex DC. Mexican yellowshowXO

Boraginaceae Amsinckia menziesii var. intermedia (Fisch & C.A. Mey.) Ganders common fiddleneckXXXOOO

Amsinckia tessellata Gray bristly fiddleneckX

Cryptantha angustifolia (Torr.) Greene Panamint cryptanthaXXO

Cryptantha barbigera (Gray) Greene bearded cryptanthaXXXOOOO

Cryptantha micrantha (Torr.) I.M. Johnston redroot cryptanthaXXXOOOO

Cryptantha muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. pointed cryptanthaXO

Cryptantha muricata var. denticulata (Greene) I.M. Johnston pointed cryptanthaXO

Cryptantha nevadensis A. Nels. & Kennedy Nevada cryptanthaXXXO

Cryptantha pterocarya (Torr.) Greene wingnut cryptanthaOXOOX

Cryptantha pterocarya var. cycloptera (Greene) J.F. Macbr. wingnut cryptanthaX

Harpagonella palmeri Gray Palmer’s grapplinghookOXXOO

Lappula occidentalis var. occidentalis (S. Wats.) Greene flatspine stickseedXXOO

Lithospermum cobrense Greene smooththroat stoneseed

XXO

Lithospermum multiflorum Torr. ex Gray manyflowered stoneseedXXO

Macromeria viridiflora DC. giant-trumpetsXX

Pectocarya heterocarpa (I.M. Johnston) I.M. Johnston chuckwalla combseedXX

Pectocarya platycarpa (Munz & Johnston) Munz & Johnston broadfruit combseedOXXOOO

Pectocarya recurvata I.M. Johnston curvenut combseedOXXOOO

Pectocarya setosa Gray moth combseedO

Plagiobothrys arizonicus (Gray) Greene ex Gray Arizona popcornflowerXXXOX
106

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Boraginaceae Plagiobothrys collinus (Phil.) I.M. Johnston Cooper’s popcornflowerXXOO

Plagiobothrys pringlei Greene Pringle’s popcornflowerXXX

Plagiobothrys tenellus (Nutt. ex Hook.) Gray Pacific popcornflowerXX

Tiquilia canescens (DC.) A. Richards. woody crinklematXXOX

Brassicaceae Arabis perennans S. Wats. perennial rockcressXXOOOXO

Brassica tournefortii Gouan Asian mustardXXX

Capsella bursa-pastoris (L.) Medik. shepherd’s purseXXOX

Descurainia pinnata (Walt.) Britt. western tansymustardXXXOOXO

Dimorphocarpa wislizeni (Engelm.) Rollins touristplantO

Draba cuneifolia Nutt. ex Torr. & Gray wedgeleaf drabaOXO

Draba cuneifolia var. integrifolia S. Wats. wedgeleaf drabaXX

Draba helleriana Greene Heller’s drabaO

Draba helleriana var. bifurcata C.L. Hitchc. Heller’s drabaX

Draba petrophila Greene var. petrophila Santa Rita Mountain drabaXO

Dryopetalon runcinatum Gray rockmustardXXO

Guillenia lasiophylla (Hook. & Arn.) Greene California mustardXXOX

Lepidium lasiocarpum Nutt. shaggyfruit pepperweedOXXOX

Lepidium thurberi Woot. Thurber’s pepperweedXXX

Lepidium virginicum L. Virginia pepperweed

XO

Lepidium virginicum var. medium (Greene) C.L. Hitchc. medium pepperweedXXOOO

Lesquerella gordonii (Gray) S. Wats. Gordon’s bladderpodXXOX

Pennellia longifolia (Benth.) Rollins longleaf mock thelypodyXX

Pennellia micrantha (Gray) Nieuwl. mountain mock thelypodyXXXO

Schoenocrambe linearifolia (Gray) Rollins slimleaf plainsmustardXXXOOOO

Sisymbrium irio L. London rocketXXXOOOXO

Streptanthus carinatus C. Wright ex Gray lyreleaf jewelflowerO

Streptanthus carinatus ssp. arizonicus (S. Wats.) Kruckeberg, 

Rodman & Worthington lyreleaf jewelflower

OX

Thelypodium Endl. thelypodyX

Thlaspi montanum var. fendleri (Gray) P. Holmgren Fendler’s pennycressXX

Thysanocarpus curvipes Hook. sand fringepodOXXOOOX

Cactaceae Carnegiea gigantea (Engelm.) Britt. & Rose saguaroXXOOOOOOO

Echinocereus coccineus Engelm. scarlet hedgehog cactusXOO

Echinocereus coccineus Engelm. var. coccineus scarlet hedgehog cactusX

Echinocereus fendleri (Engelm.) F. Seitz pinkflower hedgehog cactusXO

Echinocereus fendleri var. fasciculatus (Engelm. ex B.D. Jackson) 

N.P. Taylor pinkflower hedgehog cactus

OXOOOOOO

Echinocereus fendleri var. rectispinus (Peebles) L. Benson pinkflower hedgehog cactusO

Echinocereus pectinatus (Scheidw.) Engelm. rainbow cactusO

Echinocereus rigidissimus (Engelm.) Haage f. rainbow hedgehog cactusX

XOO

Echinocereus triglochidiatus Engelm. kingcup cactusO

Escobaria vivipara var. bisbeeana (Orcutt) D.R. Hunt Bisbee spinystarXOO

Escobaria vivipara var. vivipara (Nutt.) Buxbaum spinystarO

Ferocactus wislizeni (Engelm.) Britt. & Rose candy barrelcactusXXOOOOOOO
107

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Cactaceae Mammillaria grahamii Engelm. Graham’s nipple cactusOOO

Mammillaria grahamii var. grahamii Engelm. Graham’s nipple cactusOXOO

Mammillaria grahamii var. oliviae (Orcutt) L. Benson Graham’s nipple cactusXOOO

Mammillaria heyderi var. macdougalii (Rose) L. Benson Macdougal’s nipple cactusXOOO

Mammillaria viridiflora (Britt. & Rose) Bödecker greenflower nipple cactusXXO

Opuntia acanthocarpa Engelm. & Bigelow buckhorn chollaOO

Opuntia arbuscula Engelm. Arizona pencil chollaXOOO

Opuntia basilaris Engelm. & Bigelow beavertail pricklypearO

Opuntia bigelovii Engelm. teddybear chollaOXOOOOOO

Opuntia chlorotica Engelm. & Bigelow dollarjoint pricklypearXOOOOO

Opuntia engelmannii Salm-Dyck cactus appleOOOOOO

Opuntia engelmannii Salm-Dyck var. engelmannii cactus appleXO

Opuntia ficus-indica (L.) P. Mill. tuna cactusO

Opuntia fulgida Engelm. jumping chollaOXOO

Opuntia fulgida Engelm. var. fulgida jumping chollaXOOOO

Opuntia fulgida var. mamillata (Schott ex Engelm.) Coult. jumping chollaXOOOOO

Opuntia leptocaulis DC. Christmas cactus

OXOOO

Opuntia phaeacantha Engelm. tulip pricklypearOXOOOOOO

Opuntia phaeacantha var. laevis (Coult.) L. Benson tulip pricklypearXO

Opuntia phaeacantha var. major Engelm. Mojave pricklypearXOO

Opuntia spinosior (Engelm.) Toumey walkingstick cactusXXOOOO

Opuntia versicolor Engelm. ex Coult. staghorn chollaOXXOOOOOOO

Opuntia ×tetracantha Toumey (pro sp.) [acanthocarpa × leptocaulis]X

Peniocereus greggii var. transmontanus (Engelm.) Backeberg nightblooming cereusX

Campanulaceae Lobelia anatina F. Wimmer Apache lobeliaXX

Lobelia cardinalis L. cardinalflowerXX

Nemacladus glanduliferus Jepson glandular threadplantOXO

Nemacladus glanduliferus var. orientalis McVaugh glandular threadplantX

Triodanis holzingeri McVaugh Holzinger’s Venus’ looking-glassOXXOO

Triodanis perfoliata (L.) Nieuwl. clasping Venus’ looking-glassOXOXO

Triodanis perfoliata var. biflora (Ruiz & Pavón) Bradley clasping Venus’ looking-glassXXXOOO

Triodanis perfoliata var. perfoliata (L.) Nieuwl. clasping Venus’ looking-glassXXO

Capparaceae Polanisia dodecandra (L.) DC. redwhisker clammyweedOO

Polanisia dodecandra ssp. trachysperma (Torr. & Gray) Iltis sandyseed clammyweedXXO

Caprifoliaceae Lonicera arizonica Rehd. Arizona honeysuckleXXO

Lonicera interrupta Benth. chaparral honeysuckleXX

Sambucus nigra ssp. canadensis (L.) R. Bolli common elderberryXXO

Sambucus nigra ssp. cerulea (Raf.) R. Bolli blue elderberryXX

Symphoricarpos oreophilus Gray mountain snowberryOXXOO

Arenaria lanuginosa var. longipedunculata Duncan spreading sandwortXX

Arenaria lanuginosa ssp. saxosa (Gray) Maguire spreading sandwortXXO

Cerastium fontanum ssp. vulgare (Hartman) Greuter & Burdet big chickweedO

Cerastium gracile Dufour slender chickweedXXOO

Cerastium nutans Raf. nodding chickweedXO
108

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Caryophyllaceae Cerastium texanum Britt. Texas chickweed X O O O

Drymaria leptophylla (Cham. & Schlecht.) Fenzl ex Rohrb. canyon drymary X X O

Drymaria molluginea (Lag.) Didr. slimleaf drymary X O O O

Herniaria hirsuta ssp. cinerea (DC.) Coutinho hairy rupturewort X X

Loeflingia squarrosa Nutt. spreading pygmyleaf O X X O

Sagina decumbens ssp. occidentalis (S. Wats.) Crow western pearlwort X X

Silene antirrhina L. sleepy silene X X X O O O O O

Silene scouleri ssp. pringlei (S. Wats.) C.L. Hitchc. & Maguire simple campion X X

Stellaria nitens Nutt. shiny chickweed X X

Chenopodiaceae Atriplex canescens (Pursh) Nutt. fourwing saltbush X X O O O O O X

Atriplex elegans var. thornberi M.E. Jones wheelscale saltbush X X

Chenopodium berlandieri Moq. pitseed goosefoot X X O

Chenopodium fremontii S. Wats. Fremont’s goosefoot X X O

Chenopodium graveolens Willd. fetid goosefoot O X X O O

Chenopodium incanum (S. Wats.) Heller mealy goosefoot X

Chenopodium incanum var. elatum Crawford mealy goosefoot X

Chenopodium murale L. nettleleaf goosefoot X X

Chenopodium neomexicanum Standl. New Mexico goosefoot X O O X

Chenopodium neomexicanum var. palmeri (Standl.) T.W. Walters Palmer’s goosefoot X O

Chenopodium pratericola Rydb. desert goosefoot X

Salsola kali L. Russian thistle X

Salsola tragus L. prickly Russian thistle X O

Clusiaceae Hypericum formosum H.B.K. X X

Commelinaceae Commelina dianthifolia Delile birdbill dayflower X X X O O O

Commelina erecta L. whitemouth dayflower X O O O O

Commelina erecta var. angustifolia (Michx.) Fern. whitemouth dayflower X

Tradescantia occidentalis (Britt.) Smyth prairie spiderwort X X X O O O

Tradescantia occidentalis var. scopulorum (Rose) E.S. Anderson & 

Woods. prairie spiderwort X

Tradescantia pinetorum Greene pinewoods spiderwort X X X O

Convolvulaceae Convolvulus arvensis L. field bindweed X

Evolvulus alsinoides (L.) L. slender dwarf morning-glory X X O O O O

Evolvulus alsinoides var. angustifolius Torr. slender dwarf morning-glory X O

Evolvulus arizonicus Gray wild dwarf morning-glory X X X O O O O O

Evolvulus nuttallianus J.A. Schultes shaggy dwarf morning-glory X O

Ipomoea barbatisepala Gray canyon morning-glory X X X O O O O

Ipomoea coccinea L. redstar X O X O

Ipomoea costellata Torr. crestrib morning-glory X X X O O O O O

Ipomoea cristulata Hallier f. Transpecos morning-glory X O O

Ipomoea hederacea Jacq. ivyleaf morning-glory X O

Ipomoea plummerae Gray Huachuca Mountain morning-glory X X O

Ipomoea purpurea (L.) Roth tall morning-glory X X O X O

Ipomoea tenuiloba Torr. spiderleaf X

Ipomoea tenuiloba var. lemmonii (Gray) Yatskievych & Mason spiderleaf X X
109

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Convolvulaceae Ipomoea ternifolia var. leptotoma (Torr.) J.A. McDonald tripleleaf morning-gloryXXXOOOO

Jacquemontia pringlei Gray Pringle’s clustervineXXOOO

Cornaceae Cornus sericea ssp. sericea L. 

a redosier dogwood

X

Crassulaceae Crassula connata (Ruiz & Pavón) Berger sand pygmyweedOX

Crassula connata var. connata (Ruiz & Pavón) Berger sand pygmyweedXXOOO

Graptopetalum bartramii Rose Patagonia Mountain leatherpetalXXX

Graptopetalum rusbyi (Greene) Rose San Francisco River leatherpetalXO

Sedum cockerellii Britt. Cockerell’s stonecropXXXOO

Crossosomataceae Crossosoma bigelovii S. Wats. ragged rockflowerXXOOOOOO

Cucurbitaceae Apodanthera undulata Gray melon locoXX

Cucurbita digitata Gray fingerleaf gourdXXO

Cucurbita foetidissima Kunth Missouri gourdXX

Echinopepon wrightii (Gray) S. Wats. wild balsam appleXXXOOO

Marah gilensis Greene Gila manrootXXXOOX

Cupressaceae Cupressus arizonica Greene Arizona cypressX

Cupressus arizonica Greene ssp. arizonica Arizona cypressX

Juniperus coahuilensis (Martinez) Gaussen ex R.P. Adams redberry juniper

XXX

Juniperus deppeana Steud. alligator juniperOXXOOOOOO

Cyperaceae Bulbostylis capillaris (L.) Kunth ex C.B. Clarke densetuft hairsedgeXXXO

Bulbostylis funckii (Steud.) C.B. Clarke Funck’s hairsedgeOOO

Carex agrostoides Mackenzie grassleaf sedgeXO

Carex athrostachya Olney slenderbeak sedgeXX

Carex bonplandii Kunth Bonpland’s sedgeO

Carex chihuahuensis Mackenzie Chihuahuan sedgeXXOO

Carex foenea Willd. dryspike sedgeX

Carex geophila Mackenzie White Mountain sedgeXXO

Carex lativena S.D. & G.D. Jones broadvein sedgeX

Carex leucodonta Holm Huachuca Mountain sedgeXXXO

Carex meadii Dewey Mead’s sedgeX

Carex occidentalis Bailey western sedgeXXXO

Carex senta Boott swamp carexXXO

Carex squarrosa L. squarrose sedgeO

Carex subfusca W. Boott brown sedgeXXX

Carex thurberi Dewey Thurber’s sedgeX

Carex vallicola Dewey valley sedgeXX

Carex vallicola var. rusbyi (Mackenzie) F.J. Herm. Rusby’s sedgeX

Cyperus aggregatus (Willd.) Endl. inflatedscale flatsedgeXX

Cyperus dipsaceus Liebamann Wright’s flatsedgeXXOO

Cyperus esculentus L. chufa flatsedgeXXXOOOOX

Cyperus fendlerianus Boeckl. Fendler’s flatsedgeXXXOO

Cyperus mutisii (Kunth) Griseb. Mutis’ flatsedgeXXXOOOO

Cyperus pallidicolor (Kükenth.) G. Tucker pallid flatsedgeXXOO

Cyperus cf. parishii Britt. Parish’s flatsedgeO

Cyperus sphaerolepis Boeckl. Rusby’s flatsedgeXXXO
110

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Cyperaceae Cyperus squarrosus L. bearded flatsedgeXXXOOOOO

Cyperus strigosus L. strawcolored flatsedgeXX

Eleocharis montana (Kunth) Roemer & J.A. Schultes mountain spikerushXXX

Eleocharis montevidensis Kunth sand spikerushXXXOO

Fimbristylis annua (All.) Roemer & J.A. Schultes annual fimbryX

Lipocarpha micrantha (Vahl) G. Tucker smallflower halfchaff sedgeXXOO

Scirpus microcarpus J.& K. Presl panicled bulrushXX

Dryopteridaceae Cystopteris fragilis (L.) Bernh. brittle bladderfernX

Cystopteris reevesiana Lellinger Reeves’ bladderfernXX

Dryopteris filix-mas (L.) Schott male fernXXO

Woodsia cochisensis Windham Cochise cliff fernXOO

Woodsia mexicana Fée phanerophlebiaXXO

Woodsia oregana D.C. Eat. Oregon cliff fernO

Woodsia plummerae Lemmon Plummer’s cliff fernXXOO

Elatinaceae Elatine americana (Pursh) Arn. American waterwortX

Elatine brachysperma Gray shortseed waterwortXX

Ephedraceae Ephedra trifurca Torr. ex S. Wats. longleaf jointfirXXOOO

Equisetaceae Equisetum ×ferrissii Clute (pro sp.) ferris horsetailO

Ericaceae Arbutus arizonica (Gray) Sarg. Arizona madroneXXOO

Arctostaphylos pringlei Parry Pringle manzanita

XX

Arctostaphylos pungens Kunth pointleaf manzanitaOXXOOOO

Euphorbiaceae Acalypha neomexicana Muell.-Arg. New Mexico copperleafXXXOOOOXO

Argythamnia lanceolata (Benth.) Muell.-Arg. narrowleaf silverbushO

Argythamnia neomexicana Muell.-Arg. New Mexico silverbushXXXOOX

Bernardia incana Morton hoary myrtlecrotonX

Chamaesyce abramsiana (L.C. Wheeler) Koutnik Abrams’ sandmatXX

Chamaesyce albomarginata (Torr. & Gray) Small whitemargin sandmatXX

Chamaesyce arizonica (Engelm.) Arthur Arizona sandmatXX

Chamaesyce capitellata (Engelm.) Millsp. head sandmatXXXOOO

Chamaesyce dioica (Kunth) Millsp. royal sandmatXX

Chamaesyce florida (Engelm.) Millsp. Chiricahua Mountain sandmatXXOOOX

Chamaesyce gracillima (S. Wats.) Millsp. Mexican sandmatXXO

Chamaesyce hyssopifolia (L.) Small hyssopleaf sandmatXXXOOOOXO

Chamaesyce melanadenia (Torr.) Millsp. squaw sandmatXXXOOOO

Chamaesyce micromera (Boiss. ex Engelm.) Woot. & Standl. Sonoran sandmatXX

Chamaesyce pediculifera (Engelm.) Rose & Standl. Carrizo Mountain sandmatXXOOO

Chamaesyce polycarpa (Benth.) Millsp. ex Parish smallseed sandmatXXO

Chamaesyce prostrata (Ait.) Small prostrate sandmatX

Chamaesyce revoluta (Engelm.) Small threadstem sandmat

XXO

Chamaesyce setiloba (Engelm. ex Torr.) Millsp. ex Parish Yuma sandmatX

Croton pottsii (Klotzsch) Muell.-Arg. leatherweedXX

Croton pottsii var. pottsii (Klotzsch) Muell.-Arg. leatherweedX

Euphorbia brachycera Engelm. horned spurgeXX

Euphorbia chamaesula Boiss. mountain spurgeXX
111

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Euphorbiaceae Euphorbia cuphosperma (Engelm.) Boiss. X

Euphorbia cyathophora Murr. fire on the mountainX

Euphorbia dentata var. dentata Michx. toothed spurgeX

Euphorbia heterophylla L. Mexican fireplantXXXOOOOX

Euphorbia spathulata Lam. warty spurgeXX

Jatropha cardiophylla (Torr.) Muell.-Arg. sangre de cristoOXXOOOOOOO

Manihot angustiloba (Torr.) Muell.-Arg. desertmountain manihotXXO

Tragia nepetifolia Cav. catnip noseburnXXXOOOOXO

Tragia ramosa Torr. branched noseburnX

Fabaceae Acacia angustissima (P. Mill.) Kuntze prairie acaciaOXOOOOOOO

Acacia angustissima var. suffrutescens (Rose) Isely prairie acaciaXX

Acacia constricta Benth. whitethorn acaciaOXXOOOOO

Acacia greggii Gray catclaw acaciaOXXOOOOOOO

Acacia millefolia S. Wats. milfoil wattleXX

Amorpha californica Nutt. California false indigoXXX

Amorpha fruticosa L. desert false indigo

XXXOOOOOO

Astragalus allochrous Gray halfmoon milkvetchXXXOO

Astragalus arizonicus Gray Arizona milkvetchXXXO

Astragalus didymocarpus Hook. & Arn. dwarf white milkvetchX

Astragalus humistratus Gray groundcover milkvetchO

Astragalus nothoxys Gray sheep milkvetchXXOO

Astragalus nuttallianus DC. smallflowered milkvetchXXOOX

Astragalus nuttallianus var. austrinus (Small) Barneby smallflowered milkvetchX

Astragalus tephrodes Gray ashen milkvetchO

Calliandra eriophylla Benth. fairydusterOXXOOOOOOO

Calliandra humilis Benth. dwarf stickpeaXOO

Calliandra humilis Benth. var. humilis dwarf stickpeaX

Calliandra humilis var. reticulata (Gray) L. Benson dwarf stickpeaXXX

Chamaecrista nictitans (L.) Moench partridge peaOOO

Chamaecrista nictitans var. leptadenia (Greenm.) Gandhi & Hatch partridge peaXXX

Chamaecrista nictitans (L.) Moench ssp. nictitans partridge peaX

Clitoria mariana L. Atlantic pigeonwingsXX

Cologania angustifolia Kunth longleaf cologaniaXXOOO

Cologania lemmonii Gray Lemmon’s cologaniaXX

Cologania pallida Rose pale cologaniaO

Coursetia caribaea (Jacq.) Lavin anil falsoXO

Coursetia caribaea var. caribaea (Jacq.) Lavin anil falsoX

Coursetia glandulosa Gray rosary babybonnetsXXOOOO

Crotalaria pumila Ortega low rattleboxXXOOO

Crotalaria sagittalis L. arrowhead rattleboxXXO

Dalea albiflora Gray whiteflower prairie cloverXXXOOO

Dalea exigua Barneby Chihuahuan prairie cloverXX

Dalea filiformis Gray Sonoran prairie cloverXXXOOO

Dalea formosa Torr. featherplumeXXX
112

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Fabaceae Dalea lumholtzii B.L. Robins. & Fern. Lumholtz’s prairie cloverXXX

Dalea pogonathera Gray bearded prairie cloverXX

Dalea polygonoides Gray sixweeks prairie cloverXX

Dalea pringlei Gray Pringle’s prairie cloverXXXOOOOX

Dalea pulchra H.C. Gentry Santa Catalina prairie cloverXXXOOOO

Dalea versicolor Zucc. oakwoods prairie cloverX

Dalea versicolor var. sessilis (Gray) Barneby oakwoods prairie cloverXXOO

Dalea wrightii Gray Wright’s prairie cloverXX

Desmodium angustifolium (Kunth) DC. grassleaf ticktrefoilX

Desmodium arizonicum S. Wats. Arizona ticktrefoilXX

Desmodium batocaulon Gray San Pedro ticktrefoilXXOOOOO

Desmodium cinerascens Gray spiked ticktrefoilXXOO

Desmodium grahamii Gray Graham’s ticktrefoilXXOO

Desmodium gramineum Gray grassleaf ticktrefoilX

Desmodium neomexicanum Gray New Mexico ticktrefoilXXO

Desmodium procumbens (P. Mill.) A.S. Hitchc. western trailing ticktrefoilXXO

Desmodium procumbens var. exiguum (Gray) Schub. western trailing ticktrefoilXO

Desmodium psilocarpum Gray Santa Cruz Island ticktrefoilX

Desmodium rosei Schub. Rose’s ticktrefoilXXXO

OOO

Erythrina flabelliformis Kearney coralbeanXXOOO

Eysenhardtia orthocarpa (Gray) S. Wats. Tahitian kidneywoodXXXOOO

Galactia wrightii Gray Wright’s milkpeaXXXOOOOO

Indigofera sphaerocarpa Gray Sonoran indigoXX

Lathyrus graminifolius (S. Wats.) White grassleaf peaXXOOO

Lathyrus lanszwertii var. leucanthus (Rydb.) Dorn Nevada peaXXO

Lotus greenei Ottley ex Kearney & Peebles Greene’s bird’s-foot trefoilXXO

Lotus humistratus Greene foothill deervetchXXXOOOOO

Lotus plebeius (Brand) Barneby New Mexico bird’s-foot trefoilXXXOOOO

Lotus rigidus (Benth.) Greene shrubby deervetchXXOOO

Lotus strigosus (Nutt.) Greene strigose bird’s-foot trefoilO

Lotus strigosus var. tomentellus (Greene) Isely strigose bird’s-foot trefoilXX

Lotus wrightii (Gray) Greene Wright’s deervetchXXXO

Lupinus bicolor Lindl. miniature lupineO

Lupinus concinnus J.G. Agardh scarlet lupineOXOOOO

Lupinus concinnus ssp. orcuttii (S. Wats.) D. Dunn Orcutt’s lupineXO

Lupinus palmeri S. Wats. bluebonnet lupineOXXO

Lupinus sparsiflorus Benth. Mojave lupineOXOOO

Lupinus sparsiflorus ssp. mohavensis Dziekanowski & D. Dunn Mojave lupineXX

Lysiloma watsonii Rose littleleaf false tamarindOXXOOOOO

Macroptilium gibbosifolium (Ortega) A. Delgado variableleaf bushbeanXXOOOOO

Marina parryi (Torr. & Gray) Barneby Parry’s false prairie-cloverOXXOOO

Melilotus officinalis (L.) Lam. yellow sweetcloverX

Mimosa aculeaticarpa var. biuncifera (Benth.) Barneby catclaw mimosaXXXOOOOOOO

Mimosa grahamii Gray Graham’s mimosaXX
113

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Fabaceae Nissolia schottii (Torr.) Gray Schott’s yellowhoodXO

Parkinsonia florida (Benth. ex Gray) S. Wats. blue paloverdeOXXOOOOO

Parkinsonia microphylla Torr. yellow paloverdeOXXOOOOOO

Phaseolus acutifolius Gray tepary beanXXOOO

Phaseolus acutifolius var. tenuifolius Gray tepary beanXXOOO

Phaseolus angustissimus Gray slimleaf beanX

Phaseolus maculatus Scheele spotted beanXO

Phaseolus parvulus Greene Pinos Altos Mountain beanXXO

Phaseolus ritensis M.E. Jones Santa Rita Mountain beanX

Prosopis glandulosa Torr. honey mesquiteOO

Prosopis velutina Woot. velvet mesquiteOXXOOOOOOO

Rhynchosia senna Gillies ex Hook. Texas snoutbeanO

Rhynchosia senna var. texana (Torr. & Gray) M.C. Johnston Texas snoutbeanXXXO

Robinia neomexicana Gray New Mexico locustXXXOO

Senna bauhinioides (Gray) Irwin & Barneby twinleaf sennaXX

Senna covesii (Gray) Irwin & Barneby Coves’ cassiaOXXOOOO

Senna hirsuta (L.) Irwin & Barneby woolly senna

X

Senna hirsuta var. glaberrima (M.E. Jones) Irwin & Barneby woolly sennaXXO

Sphinctospermum constrictum (S. Wats.) Rose hourglass peaseedO

Tephrosia leiocarpa Gray smoothpod hoarypeaXXOO

Tephrosia tenella Gray red hoarypeaXXXOOO

Trifolium pinetorum Greene woods cloverXXO

Trifolium variegatum Nutt. whitetip cloverXXX

Vicia americana Muhl. ex Willd. American vetchXXO

Vicia americana Muhl. ex Willd. ssp. americana American vetchXO

Vicia leucophaea Greene 

a

 Mogollon Mountain vetchX

Vicia ludoviciana Nutt. Louisiana vetchXXOOOO

Vicia ludoviciana ssp. ludoviciana Nutt. Louisiana vetchOXX

Vicia pulchella Kunth sweetclover vetchXXO

Zornia gemella Vogel dos hoja zazabacoa de dos hojasXXX

Fagaceae Quercus arizonica Sarg. Arizona white oakXXXOOOOO

Quercus dunnii Kellogg Palmer oakXX

Quercus emoryi Torr. Emory oakXXXOOOOO

Quercus gambelii Nutt. Gambel oakXXO

Quercus hypoleucoides A. Camus silverleaf oakXXOOO

Quercus oblongifolia Torr. Mexican blue oakXXXO

OOOO

Quercus rugosa Née netleaf oakXXXO

Quercus toumeyi Sarg. Toumey oakXX

Quercus turbinella Greene Sonoran scrub oakOXXO

Fouquieriaceae Fouquieria splendens Engelm. ocotilloOXXOOOOOOO

Fumariaceae Corydalis aurea Willd. scrambled eggsXOOO

Corydalis curvisiliqua ssp. occidentalis (Engelm. ex Gray) W.A. 

Weber curvepod fumewort

XX

Garrya wrightii Torr. Wright’s silktasselOXOOOO
114

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Gentianaceae Centaurium calycosum (Buckl.) Fern. Arizona centauryXXXOO

Centaurium exaltatum (Griseb.) W. Wight ex Piper desert centauryX

Centaurium nudicaule (Engelm.) B.L. Robins. Santa Catalina Mountain centauryXXXO

Frasera speciosa Dougl. ex Griseb. elkweedXX

Gentiana affinis Griseb. pleated gentianXX

Gentianella microcalyx (J.G. Lemmon) J. Gillett Chiricahua dwarf gentianXXXO

Geraniaceae Erodium cicutarium (L.) L’Hér. ex Ait. redstem stork’s bill

XXOOXO

Erodium texanum Gray Texas stork’s billXO

Geranium caespitosum James pineywoods geraniumXXOO

Geranium carolinianum L. Carolina geraniumXXXOO

Geranium richardsonii Fisch. & Trautv. Richardson’s geraniumXXO

Hydrangeaceae Philadelphus argenteus Rydb. silver mock orangeXXX

Philadelphus argyrocalyx Woot. silvercup mock orangeX

Philadelphus microphyllus Gray littleleaf mock orangeO

Hydrophyllaceae Emmenanthe penduliflora Benth. whisperingbellsXXO

Eriodictyon angustifolium Nutt. narrowleaf yerba santaXXX

Eucrypta chrysanthemifolia (Benth.) Greene spotted hideseedXOX

Eucrypta chrysanthemifolia var. bipinnatifida (Torr.) Constance spotted hideseedXO

Eucrypta micrantha (Torr.) Heller dainty desert hideseedXXX

Nama demissum Gray purplemat

X

Nama dichotomum (Ruiz & Pavón) Choisy wishbone fiddleleafXX

Nama hispidum Gray bristly namaXXXO

Phacelia affinis Gray limestone phaceliaXXO

Phacelia bombycina Woot. & Standl. Mangas Spring phaceliaOXXO

Phacelia caerulea Greene skyblue phaceliaXXO

Phacelia crenulata Torr. ex S. Wats. cleftleaf wildheliotropeO

Phacelia cryptantha Greene hiddenflower phaceliaXXO

Phacelia distans Benth. distant phaceliaXXXOOO

Phacelia egena (Greene ex Brand) Greene ex J.T. Howell Kaweah River phaceliaXO

Phacelia ramosissima Dougl. ex Lehm. branching phaceliaXXO

Iridaceae Sisyrinchium arizonicum Rothrock Arizona blue-eyed grassO

Sisyrinchium cernuum (Bickn.) Kearney nodding blue-eyed grassXXXOOO

Sisyrinchium demissum Greene stiff blue-eyed grassXXO

Sisyrinchium longipes (Bickn.) Kearney & Peebles timberland blue-eyed grassXX

Juglandaceae Juglans major (Torr.) Heller Arizona walnutXXOOOO

Juncaceae Juncus acuminatus Michx. tapertip rushXXOO

Juncus balticus Willd. Baltic rushO

Juncus bufonius L. toad rushXXXOO

Juncus effusus L. common rushXXXO

Juncus effusus var. brunneus Engelm. lamp rushX

Juncus interior Wieg. inland rushX

XXO

Juncus marginatus Rostk. grassleaf rushXXOO

Juncus saximontanus A. Nels. Rocky Mountain rushXXX

Juncus tenuis Willd. poverty rushXOOO
115

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Juncaceae Juncus xiphioides E. Mey. irisleaf rushX

Luzula multiflora (Ehrh.) Lej. common woodrushXXX

Krameriaceae Krameria erecta Willd. ex J.A. Schultes littleleaf ratanyXXOO

Krameria grayi Rose & Painter white ratanyOO

Krameria lanceolata Torr. trailing krameriaXX

Lamiaceae Agastache breviflora (Gray) Epling TransPecos giant hyssopXX

Agastache pallidiflora (Heller) Rydb. Bill Williams Mountain giant hyssopO

Agastache wrightii (Greenm.) Woot. & Standl. Sonoran giant hyssopXX

Hedeoma dentata Torr. dentate false pennyroyalXXOO

Hedeoma hyssopifolia Gray aromatic false pennyroyalXXO

Hedeoma nana (Torr.) Briq. dwarf false pennyroyalX

Hedeoma nana (Torr.) Briq. ssp. nana dwarf false pennyroyalXOO

Hedeoma nanum (Torrey) Briq. OXO

Hyptis emoryi Torr. desert lavenderOXXOOOOOO

Marrubium vulgare L. horehoundXXOOX

Monarda citriodora Cerv. ex Lag. lemon beebalmO

Monarda citriodora ssp. austromontana (Epling) Scora lemon beebalmXXXO

Monarda fistulosa var. menthifolia (Graham) Fern. wild bergamotXX

Monardella odoratissima Benth. mountain monardellaX

Salvia arizonica Gray desert indigo sageXX

O

Salvia columbariae Benth. chiaOXXOOO

Salvia reflexa Hornem. lanceleaf sageXOO

Salvia subincisa Benth. sawtooth sageO

Stachys coccinea Ortega scarlet hedgenettleXXXOOOO

Trichostema arizonicum Gray Arizona bluecurlsXX

Liliaceae Allium bigelovii S. Wats. Bigelow’s onionO

Allium bisceptrum var. palmeri (S. Wats.) Cronq. aspen onionXX

Allium geyeri S. Wats. Geyer’s onionXXXO

Allium macropetalum Rydb. largeflower onionXX

Calochortus ambiguus (M.E. Jones) Ownbey doubting mariposa lilyXXOO

Calochortus kennedyi Porter desert mariposa lilyXO

Dasylirion wheeleri S. Wats. common sotolXXOOOOOOO

Dichelostemma capitatum (Benth.) Wood bluedicksO

Dichelostemma capitatum (Benth.) Wood ssp. capitatum bluedicksOXOOOO

Echeandia flavescens (J.A. & J.H. Schultes) Cruden Torrey’s craglilyXXXO

Maianthemum racemosum ssp. racemosum (L.) Link feathery false lily of the vallyXX

Maianthemum stellatum (L.) Link starry false lily of the vallyXXO

Nolina microcarpa S. Wats. sacahuistaOXXOOOOO

Nothoscordum texanum M.E. Jones Texas false garlicXX

Zephyranthes longifolia Hemsl. copper zephyrlilyX

XOO

Linaceae Linum lewisii Pursh prairie flaxXXX

Linum neomexicanum Greene New Mexico yellow flaxXXXO

Loasaceae Mentzelia affinis Greene yellowcometXOOX

Mentzelia albicaulis (Dougl. ex Hook.) Dougl. ex Torr. & Gray whitestem blazingstarXXOO
116

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Loasaceae Mentzelia asperula Woot. & Standl. Organ Mountain blazingstarXOOO

Mentzelia isolata H.C. Gentry isolated blazingstarO

Mentzelia jonesii (Urban & Gilg) H.J. Thompson & Roberts Jones’ blazingstarX

Mentzelia multiflora (Nutt.) Gray Adonis blazingstarX

Mentzelia nitens Greene shining blazingstarXX

Lythraceae Cuphea wrightii Gray Wright’s waxweedXXOO

Malpighiaceae Janusia gracilis Gray slender janusiaOXXOOOOOO

Malvaceae Abutilon abutiloides (Jacq.) Garcke ex Britt. & Wilson shrubby indian mallowOOOOOX

Abutilon berlandieri Gray ex S. Wats. Berlandier Indian mallowXX

Abutilon incanum (Link) Sweet pelotazoXXXOOOOX

Abutilon mollicomum (Willd.) Sweet Sonoran Indian mallowXXOOOO

Abutilon parishii S. Wats. Parish’s Indian mallowXXOOO

Abutilon parvulum Gray dwarf Indian mallowX

Abutilon reventum S. Wats. yellowflower Indian mallowXXXOX

Anoda abutiloides Gray Indian anodaXXXOOO

Anoda cristata (L.) Schlecht. crested anodaXXOOOO

Gossypium thurberi Todaro Thurber’s cottonXXXOOOOOO

Herissantia crispa (L.) Briz. bladdermallowXXOOOO

Hibiscus biseptus S. Wats. Arizona rosemallowXXOO

Hibiscus coulteri Harvey ex Gray desert rosemallowXXOOOOOO

Hibiscus denudatus Benth. palefaceOXOOO

Horsfordia newberryi (S. Wats.) Gray Newberry’s velvetmallowX

Malva parviflora L. cheeseweed mallowX

Rhynchosida physocalyx (Gray) Fryxell buffpetalXOO

Sida abutifolia P. Mill. spreading fanpetalsXXXX

Sida spinosa L. prickly fanpetalsX

Sphaeralcea ambigua Gray desert globemallowO

Sphaeralcea emoryi Torr. ex Gray Emory’s globemallowXXO

Sphaeralcea fendleri Gray Fendler’s globemallowXXXOOO

Sphaeralcea fendleri ssp. venusta Kearney thicket globemallowX

Sphaeralcea laxa Woot. & Standl. caliche globemallowXXXOOOO

Molluginaceae Mollugo cerviana (L.) Ser. threadstem carpetweed

XXO

Mollugo verticillata L. green carpetweedXXOO

Monotropaceae Pterospora andromedea Nutt. woodland pinedropsXX

Moraceae Morus microphylla Buckl. Texas mulberryXXOOO

O

Nyctaginaceae Allionia incarnata L. trailing windmillsOXXOOOOOO

Boerhavia coccinea P. Mill. scarlet spiderlingXXXXO

Boerhavia coulteri (Hook. f.) S. Wats. Coulter’s spiderlingXX

Boerhavia diffusa L. red spiderlingOOOO

Boerhavia erecta L. erect spiderlingXXO

Boerhavia gracillima Heimerl slimstalk spiderlingXOO

Boerhavia intermedia M.E. Jones fivewing spiderlingXXOX

Boerhavia purpurascens Gray purple spiderlingXX

Boerhavia scandens L. climbing wartclubOXXOOOOO
117

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Nyctaginaceae Boerhavia spicata Choisy creeping spiderlingXX

Boerhavia wrightii Gray largebract spiderlingX

Mirabilis albida (Walt.) Heimerl white four o’clockXXXOOO

Mirabilis coccinea (Torr.) Benth. & Hook. f. scarlet four o’clockXXXOOOO

Mirabilis comata (Small) Standl. hairy-tuft four o’clockX

Mirabilis glabra (S. Wats.) Standl. smooth four o’clockO

Mirabilis longiflora L. sweet four o’clockXXOOO

Mirabilis oxybaphoides (Gray) Gray smooth spreading four o’clockOO

Oleaceae Fraxinus anomala Torr. ex S. Wats. singleleaf ashXO

Fraxinus velutina Torr. velvet ashXXOOOOO

Menodora scabra Gray rough menodoraXXOOOO

Onagraceae Calylophus hartwegii (Benth.) Raven Hartweg’s sundropsO

Calylophus hartwegii ssp. pubescens (Gray) Towner & Raven Hartweg’s sundropsX

Camissonia californica (Nutt. ex Torr. & Gray) Raven California suncupXXOX

Camissonia chamaenerioides (Gray) Raven longcapsule suncupXXX

Epilobium canum ssp. latifolium (Hook.) Raven hummingbird trumpetXXXOOOO

Epilobium foliosum (Torr. & Gray) Suksdorf California willowherbXX

Gaura coccinea Nutt. ex Pursh scarlet beeblossomXXO

Gaura hexandra ssp. gracilis (Woot. & Standl.) Raven & Gregory harlequinbush

XXO

Gaura mollis James velvetweedX

Oenothera caespitosa Nutt. tufted evening-primroseXXXO

Oenothera elata ssp. hirsutissima (Gray ex S. Wats.) W. Dietr. Hooker’s evening-primroseXOO

Oenothera elata ssp. hookeri (Torr. & Gray) W. Dietr. & W.L. Wagner Hooker’s evening-primroseXXO

Oenothera laciniata Hill cutleaf evening-primroseXXO

Oenothera primiveris Gray desert evening-primroseXXXOO

Oenothera pubescens Willd. ex Spreng. South American evening-primroseX

Orchidaceae Corallorrhiza maculata (Raf.) Raf. summer coralrootX

Corallorrhiza maculata var. occidentalis (Lindl.) Ames summer coralrootX

Corallorrhiza striata Lindl. hooded coralrootX

Hexalectris spicata (Walt.) Barnh. spiked crested coralrootX

Malaxis ehrenbergii (Reichenb. f.) Kuntze Ehrenberg’s adder’s-mouth orchidO

Malaxis macrostachya (Lex.) Kuntze Chiricahua adder’s-mouth orchidXXO

Spiranthes parasitica A. Rich. & Gal. parasitic ladies’-tressesX

Orobanchaceae Orobanche cooperi (Gray) Heller desert broomrapeXX

Orobanche fasciculata Nutt. clustered broomrapeXO

Oxalidaceae Oxalis albicans ssp. pilosa (Nutt.) Eiten radishroot woodsorrelXXXOOOOO

Oxalis alpina (Rose) Rose ex R. Knuth alpine woodsorrelXXOOO

Oxalis decaphylla Kunth tenleaf woodsorrelX

Oxalis drummondii Gray Drummond’s woodsorrelO

Papaveraceae Argemone polyanthemos (Fedde) G.B. Ownbey crested pricklypoppyO

Eschscholzia californica ssp. mexicana (Greene) C. Clark California poppyXOO

Platystemon californicus Benth. creamcupsXOX

Parmeliaceae Usnea arizonica Mot. Arizona beard lichen

X

Passifloraceae Passiflora mexicana Juss. Mexican passionflowerXXXO
118

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Pedaliaceae Proboscidea althaeifolia (Benth.) Dcne. desert unicorn-plantX

Proboscidea parviflora (Woot.) Woot. & Standl. doubleclawXXOO

Phytolaccaceae Phytolacca americana L. American pokeweedOO

Phytolacca icosandra L. X

Rivina humilis L. rougeplantXXOOOO

Pinaceae Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. white firXO

Pinus arizonica Engelm. var. arizonica Arizona pineXO

Pinus cembroides Zucc. Mexican pinyonOXOOO

Pinus discolor D.K. Bailey & Hawksworth border pinyonOXXOO

Pinus edulis Engelm. twoneedle pinyonOO

Pinus leiophylla Schiede & Deppe Chihuahuan pineO

Pinus leiophylla var. chihuahuana (Engelm.) Shaw Chihuahuan pineXXOOO

Pinus ponderosa P.& C. Lawson ponderosa pineOO

Pinus ponderosa var. scopulorum Engelm. ponderosa pineXXO

Pinus strobiformis Engelm. southwestern white pineXXO

Pseudotsuga menziesii (Mirbel) Franco Douglas firO

Pseudotsuga menziesii var. glauca (Beissn.) Franco Rocky Mountain Douglas firX

Plantaginaceae Plantago ovata Forsk. desert IndianwheatXXOO

Plantago patagonica Jacq. woolly plantainXXXOOOX

O

Plantago virginica L. Virginia plantainXXXOOOOO

Platanaceae Platanus wrightii S. Wats. Arizona sycamoreXXOO

Plumbaginaceae Plumbago scandens L. doctorbushXXXOOOOX

Poaceae Aegopogon tenellus (DC.) Trin. fragilegrassXOOO

Agrostis elliottiana J.A. Schultes Elliott’s bentgrassX

Agrostis exarata Trin. spike bentgrassXX

Agrostis gigantea Roth redtopO

Agrostis scabra Willd. rough bentgrassXXXOOXO

Agrostis stolonifera L. creeping bentgrassXO

Alopecurus carolinianus Walt. Carolina foxtailXX

Andropogon L. bluestemO

Aristida adscensionis L. sixweeks threeawnXXXOOOOOOO

Aristida arizonica Vasey Arizona threeawnO

Aristida californica var. glabrata Vasey Santa Rita threeawnXX

Aristida havardii Vasey Havard’s threeawnO

Aristida purpurea Nutt. purple threeawnXXOO

Aristida purpurea var. longiseta (Steud.) Vasey Fendler threeawnO

Aristida purpurea var. nealleyi (Vasey) Allred blue threeawnOO

Aristida purpurea var. parishii (A.S. Hitchc.) Allred Parish’s threeawnXO

Aristida purpurea var. purpurea Nutt. purple threeawnXX

Aristida purpurea var. wrightii (Nash) Allred Wright’s threeawnXO

Aristida schiedeana var. orcuttiana (Vasey) Allred & Valdés-Reyna Orcutt’s threeawnXXXOO

Aristida ternipes Cav. spidergrassOXXOOO

Aristida ternipes var. gentilis (Henr.) Allred spidergrassXXXOO

Aristida ternipes Cav. var. ternipes spidergrassXXOOO
119

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Poaceae Avena fatua L. wild oat

OXXOOOXO

Avena sativa L. common oatX

Blepharoneuron tricholepis (Torr.) Nash pine dropseedXXXO

Bothriochloa barbinodis (Lag.) Herter cane bluestemXXXOOOOOOO

Bothriochloa ischaemum (L.) Keng yellow bluestemX

Bouteloua aristidoides (Kunth) Griseb. needle gramaXXOOOOO

Bouteloua barbata Lag. sixweeks gramaXXXOOO

Bouteloua chondrosioides (Kunth) Benth. ex S. Wats. sprucetop gramaOXXOO

Bouteloua curtipendula (Michx.) Torr. sideoats gramaXXXOOOOOOO

Bouteloua eludens Griffiths Santa Rita Mountain gramaOO

Bouteloua eriopoda (Torr.) Torr. black gramaXXOOX

Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths blue gramaXXO

Bouteloua hirsuta Lag. hairy gramaXXXOOOOOO

Bouteloua radicosa (Fourn.) Griffiths purple gramaXXXO

Bouteloua repens (Kunth) Scribn. & Merr. slender grama

XXXOOOOXO

Bouteloua rothrockii Vasey Rothrock’s gramaXXO

Bouteloua trifida Thurb. red gramaXX

Bromus anomalus Rupr. ex Fourn. nodding bromeX

Bromus arizonicus (Shear) Stebbins Arizona bromeXXX

Bromus carinatus Hook. & Arn. California bromeXXXOOXO

Bromus catharticus Vahl rescuegrassXX

Bromus ciliatus L. fringed bromeXXXO

Bromus ciliatus var. richardsonii (Link) Boivin fringed bromeX

Bromus rubens L. red bromeOXXOOOOOXOO

Bromus tectorum L. cheatgrassXXO

Cenchrus longispinus Walt. Burgrass 

b

Cenchrus spinifex Cav. coastal sandburO

Chloris crinita Lag. false Rhodes grassXXX

Chloris virgata Sw. feather fingergrassXXOOXO

Cortaderia selloana (J.A. & J.H. Schultes) Aschers. & Graebn. Uruguayan pampas grassOO

Cottea pappophoroides Kunth cotta grassXXOO

Cynodon dactylon (L.) Pers. BermudagrassXXXOOOOOXO

Dactyloctenium aegyptium (L.) Willd. Egyptian grassX

Danthonia californica Boland. California oatgrassX

X

Dasyochloa pulchella (Kunth) Willd. ex Rydb. low woollygrassXXOOOOO

Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. 

acuminatum acuminatum

XX

Dichanthelium oligosanthes var. scribnerianum (Nash) Gould Scribner’s rosette grassXXXOO

Digitaria californica (Benth.) Henr. Arizona cottontopXXXOOOOO

Digitaria ciliaris (Retz.) Koel. southern crabgrassXX

Digitaria cognata (J.A. Schultes) Pilger Carolina crabgrassXX

Digitaria cognata (J.A. Schultes) Pilger var. cognata Carolina crabgrassO

Digitaria sanguinalis (L.) Scop. hairy crabgrassXXX

Echinochloa colona (L.) Link jungle riceXXXOOX
120

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Poaceae Echinochloa crus-galli (L.) Beauv. barnyardgrass

OO

Elymus arizonicus (Scribn. & J.G. Sm.) Gould Arizona wheatgrassXXO

Elymus elymoides (Raf.) Swezey squirreltailXXOOOOO

Elyonurus barbiculmus Hack. XXOOO

Enneapogon desvauxii Desv. ex Beauv. nineawn pappusgrassXXXOOOO

Eragrostis cilianensis (All.) Vign. ex Janchen stinkgrassXXOOOXX

Eragrostis curvula (Schrad.) Nees weeping lovegrassOOOXX

Eragrostis echinochloidea Stapf African lovegrassXXXX

Eragrostis intermedia A.S. Hitchc. plains lovegrassXXXOOOOOO

Eragrostis lehmanniana Nees Lehmann lovegrassXXXOOOOOXX

Eragrostis mexicana (Hornem.) Link Mexican lovegrassOXX

Eragrostis mexicana ssp. mexicana (Hornem.) Link Mexican lovegrassXOOO

Eragrostis pectinacea (Michx.) Nees ex Steud. tufted lovegrassXXO

Eragrostis pectinacea var. miserrima (Fourn.) J. Reeder desert lovegrassXOO

Eragrostis pectinacea (Michx.) Nees ex Steud. var. pectinacea tufted lovegrass

OOO

Eriochloa acuminata (J. Presl) Kunth tapertip cupgrassXO

Eriochloa acuminata var. acuminata (J. Presl) Kunth tapertip cupgrassXXOOO

Eriochloa aristata Vasey bearded cupgrassXXOOOX

Eriochloa lemmonii Vasey & Scribn. canyon cupgrassO

Festuca sororia Piper 

a

 ravine fescueX

Glyceria striata (Lam.) A.S. Hitchc. fowl mannagrassXX

Hesperostipa comata (Trin. & Rupr.) ssp. comata needle and threadO

Hesperostipa neomexicana (Thurb. ex Coult.) Barkworth New Mexico feathergrassO

Heteropogon contortus (L.) Beauv. ex Roemer & J.A. Schultes tangleheadXXXOOOOOOO

Heteropogon melanocarpus (Ell.) Ell. ex Benth. sweet tangleheadXXOOO

Hilaria belangeri (Steud.) Nash curly-mesquiteOXXOOO

Hordeum murinum ssp. glaucum (Steud.) Tzvelev smooth barleyXXXO

Hordeum murinum ssp. leporinum (Link) Arcang. leporinum barleyXOO

Hordeum pusillum Nutt. little barleyXXXOO

Hordeum vulgare L. 

a common barley

X

Koeleria macrantha (Ledeb.) J.A. Schultes prairie JunegrassXXXO

Lamarckia aurea (L.) Moench goldentop grassX

Leptochloa dubia (Kunth) Nees green sprangletopXXXOOOOXO

Leptochloa fusca ssp. fascicularis (Lam.) N. Snow bearded sprangletopXXO

Leptochloa panicea ssp. brachiata (Steudl.) N. Snow mucronate sprangeltopXOOO

Leptochloa panicea ssp. mucronata (Michx.) Nowack mucronate sprangeltopXO

Lycurus phleoides Kunth common wolfstailOO

Lycurus setosus (Nutt.) C.G. Reeder bristly wolfstailXXXOOOOOO

Melinis repens (Willd.) Zizka rose Natal grassOXXOOOOX

Muhlenbergia arizonica Scribn. Arizona muhlyOXXOOO

Muhlenbergia dumosa Scribn. ex Vasey bamboo muhlyXXOOOX

Muhlenbergia elongata Scribn. ex Beal sycamore muhlyXX

Muhlenbergia emersleyi Vasey bullgrassXXXOOOOOO

Muhlenbergia fragilis Swallen delicate muhlyXXXOO
121

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Poaceae Muhlenbergia longiligula A.S. Hitchc. longtongue muhlyXO

Muhlenbergia microsperma (DC.) Trin. littleseed muhlyXXOO

Muhlenbergia minutissima (Steud.) Swallen annual muhlyXXOOO

Muhlenbergia montana (Nutt.) A.S. Hitchc. mountain muhlyXX

Muhlenbergia pauciflora Buckl. New Mexico muhlyXXOOO

Muhlenbergia pectinata C.O. Goodding combtop muhlyO

Muhlenbergia porteri Scribn. ex Beal bush muhlyOXXOOOOOO

Muhlenbergia ramulosa (Kunth) Swallen green muhlyXX

Muhlenbergia rigens (Benth.) A.S. Hitchc. deergrassXXXOOOOO

Muhlenbergia sinuosa Swallen marshland muhlyXXXOOOO

Muhlenbergia tenuifolia (Kunth) Trin. slimflower muhlyXXOOO

Muhlenbergia texana Buckl. Texas muhlyXXO

Muhlenbergia virescens (Kunth) Kunth screwleaf muhlyXXXO

Nassella tenuissima (Trin.) Barkworth finestem tussockgrassO

Panicum bulbosum Kunth bulb panicgrassXXXOOOOO

Panicum capillare L. witchgrassXOO

Panicum hallii Vasey var. hallii Hall’s panicgrass

OO

Panicum hirticaule J. Presl Mexican panicgrassXXOOOO

Pappophorum vaginatum Buckl. whiplash pappusgrassXXO

Paspalum dilatatum Poir. dallisgrassO

Pennisetum ciliare (L.) Link buffelgrassOXOOOOXXO

Pennisetum setaceum (Forsk.) Chiov. crimson fountaingrassXOOOOOXO

Phalaris canariensis L. annual canarygrassO

Phalaris caroliniana Walt. Carolina canarygrassXXXOX

Phalaris minor Retz. littleseed canarygrassX

Phleum pratense L. timothyXX

Piptochaetium fimbriatum (Kunth) A.S. Hitchc. pinyon ricegrassXXOOOO

Piptochaetium pringlei (Beal) Parodi Pringle’s speargrassXXXO

Poa annua L. annual bluegrassXX

Poa bigelovii Vasey & Scribn. Bigelow’s bluegrassXXXOX

Poa fendleriana (Steud.) Vasey muttongrassXXO

Poa pratensis L. Kentucky bluegrassXXO

Poa secunda J. Presl Sandberg bluegrassX

Polypogon monspeliensis (L.) Desf. annual rabbitsfoot grassXXXOOOOXXO

Polypogon viridis (Gouan) Breistr. beardless rabbitsfoot grassXXXO

Schismus arabicus Nees Arabian schismus

XXO

Schismus barbatus (Loefl. ex L.) Thellung common Mediterranean grassOXXOOX

Schizachyrium cirratum (Hack.) Woot. & Standl. Texas bluestemXXXOOOO

Schizachyrium sanguineum (Retz.) Alston crimson bluestemXOO

Schizachyrium sanguineum var. hirtiflorum (Nees) Hatch crimson bluestemXXOO

Setaria grisebachii Fourn. Grisebach’s bristlegrassXXOOO

Setaria leucopila (Scribn. & Merr.) K. Schum. streambed bristlegrassXO

Setaria vulpiseta (Lam.) Roemer & J.A. Schultes plains bristlegrassXXXOOO

Sorghum halepense (L.) Pers. JohnsongrassXXOX
122

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Poaceae Sphenopholis obtusata (Michx.) Scribn. prairie wedgescaleXXXO

Sporobolus airoides (Torr.) Torr. alkali sacatonXX

Sporobolus contractus A.S. Hitchc. spike dropseedOXOOX

Sporobolus cryptandrus (Torr.) Gray sand dropseedXX

Sporobolus texanus Vasey Texas dropseedO

Sporobolus wrightii Munro ex Scribn. big sacatonOOOX

Stipa L. needlegrassO

Trachypogon spicatus (L.) Kuntze spiked crinkleawnXXXOOOO

Tridens muticus (Torr.) Nash slim tridensXXXOOX

Tridens muticus var. muticus (Torr.) Nash slim tridensX

Trisetum interruptum Buckl. prairie false oatXXX

Urochloa arizonica (Scribn. & Merr.) O. Morrone & F. Zuloaga Arizona signalgrassXXXOO

Vulpia microstachys (Nutt.) Munro small fescueXOX

Vulpia microstachys var. ciliata (Beal) Lonard & Gould Eastwood fescueXXO

Vulpia microstachys var. pauciflora (Scribn. ex Beal) Lonard & Gould Pacific fescueO

Vulpia myuros (L.) K.C. Gmel. rat-tail fescueO

Vulpia octoflora (Walt.) Rydb. sixweeks fescueXXXOXO

Vulpia octoflora var. hirtella (Piper) Henr. sixweeks fescueXO

Vulpia octoflora var. octoflora (Walt.) Rydb. sixweeks fescueXOO

Polemoniaceae Allophyllum gilioides (Benth.) A.& V. Grant dense false gilyflower

XX

Eriastrum diffusum (Gray) Mason miniature woollystarXXXOOOO

Eucrypta Nutt. hideseedX

Gilia flavocincta A. Nels. lesser yellowthroat giliaXOO

Gilia flavocincta ssp. australis (A.& V. Grant) Day & V. Grant lesser yellowthroat giliaO

Gilia mexicana A.& V. Grant El Paso giliaO

Gilia sinuata Dougl. ex Benth. rosy giliaXXOO

Gilia stellata Heller star giliaXX

Ipomopsis longiflora (Torr.) V. Grant flaxflowered ipomopsisX

Ipomopsis multiflora (Nutt.) V. Grant manyflowered ipomopsisOXXOOO

Linanthus aureus (Nutt.) Greene golden linanthusXXOOOO

Linanthus bigelovii (Gray) Greene Bigelow’s linanthusXXO

Linanthus nuttallii (Gray) Greene ex Milliken Nuttall’s linanthusXX

Phlox gracilis (Hook.) Greene slender phloxOX

Phlox gracilis ssp. gracilis (Hook.) Greene slender phloxXXO

Phlox tenuifolia E. Nels. Santa Catalina Mountain phloxXXOO

Polygalaceae Monnina wrightii Gray blue pygmyflowerXXXOOO

Polygala alba Nutt. white milkwortXXXOO

Polygala macradenia Gray glandleaf milkwortXXO

Polygala obscura Benth. velvetseed milkwortXXXOOO

Polygala scoparioides Chod. broom milkwortXX

Polygonaceae Chorizanthe brevicornu Torr. brittle spineflowerXO

Eriogonum abertianum Torr. Abert’s buckwheatXXXOX

Eriogonum deflexum Torr. flatcrown buckwheatX

Eriogonum deflexum Torr. var. deflexum flatcrown buckwheatX
123

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Polygonaceae Eriogonum palmerianum Reveal Palmer’s buckwheatX

Eriogonum pharnaceoides Torr. wirestem buckwheatX

Eriogonum pharnaceoides Torr. var. pharnaceoides wirestem buckwheatXO

Eriogonum polycladon Benth. sorrel buckwheatXOX

Eriogonum thurberi Torr. Thurber’s buckwheatO

Eriogonum trichopes Torr. little deserttrumpetX

Eriogonum wrightii Torr. ex Benth. bastardsageXXOOOO

Eriogonum wrightii var. wrightii Torr. ex Benth. bastardsageXXOOO

Polygonum aviculare L. prostrate knotweedXXO

Polygonum douglasii ssp. johnstonii (Munz) Hickman Johnston’s knotweedXX

Polygonum hydropiperoides Michx. swamp smartweedXO

Polygonum persicaria L. spotted ladysthumbXXX

Pterostegia drymarioides Fisch. & C.A. Mey. woodland pterostegiaXXOO

Rumex acetosella L. common sheep sorrelXX

Rumex crispus L. curly dockXXOOOX

Rumex hymenosepalus Torr. canaigre dockXXOOO

Polypodiaceae Polypodium hesperium Maxon western polypodyXX

Portulacaceae Calandrinia ciliata (Ruiz & Pavón) DC. fringed redmaidsXXOO

Cistanthe monandra (Nutt.) Hershkovitz common pussypawsXXO

Claytonia perfoliata Donn ex Willd. miner’s lettuce

OOO

Claytonia perfoliata ssp. perfoliata Donn ex Willd. miner’s lettuceX

Portulaca halimoides L. silkcotton purslaneXXO

Portulaca oleracea L. little hogweedXXOO

Portulaca suffrutescens Engelm. shrubby purslaneXXXOOOOOXO

Portulaca umbraticola Kunth wingpod purslaneXXX

Portulaca umbraticola Kunth ssp. umbraticola wingpod purslaneOO

Talinum aurantiacum Engelm. orange fameflowerXX

Talinum paniculatum (Jacq.) Gaertn. jewels of OparXXOOO

Talinum parviflorum Nutt. sunbrightXX

Primulaceae Anagallis minima (L.) Krause chaffweedXXO

Androsace occidentalis Pursh western rockjasmineXXXOOX

Androsace septentrionalis L. pygmyflower rockjasmineX

Androsace septentrionalis ssp. puberulenta (Rydb.) G.T. Robbins pygmyflower rockjasmineXX

Primula rusbyi Greene Rusby’s primroseXXO

Samolus vagans Greene Chiricahua Mountain brookweedXXX

Psilotaceae Psilotum nudum (L.) Beauv. whisk fernOO

Pteridaceae Adiantum capillus-veneris L. common maidenhairXOOO

Astrolepis cochisensis (Goodding) Benham & Windham Cochise scaly cloakfernXXOOO

Astrolepis cochisensis ssp. cochisensis (Goodding) Benham & 

Windham Cochise scaly cloakfern

XOO

Astrolepis integerrima (Hook.) Benham & Windham hybrid cloakfern

XX

Astrolepis sinuata (Lag. ex Sw.) Benham & Windham wavy scaly cloakfernXOOO

Astrolepis sinuata (Lag. ex Sw.) Benham & Windham ssp. sinuata wavy scaly cloakfernXXXOOOO

Bommeria hispida (Mett. ex Kuhn) Underwood copper fernXXXOOOOO
124

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Pteridaceae Cheilanthes bonariensis (Willd.) Proctor golden lipfernXXOOOOOO

Cheilanthes covillei Maxon Coville’s lipfernOOO

Cheilanthes eatonii Baker Eaton’s lipfernXXO

Cheilanthes feei T. Moore slender lipfernXX

Cheilanthes fendleri Hook. Fendler’s lipfernXXOO

Cheilanthes lindheimeri Hook. fairyswordsXXXOOOOOXO

Cheilanthes wootonii Maxon beaded lipfernXXXOOOO

Cheilanthes wrightii Hook. Wright’s lipfernXXXOOOOOOO

Cheilanthes yavapensis Reeves ex Windham graceful lipfernXO

Notholaena grayi Davenport Gray’s cloak fernXXO

Notholaena lemmonii D.C. Eat. Lemmon’s cloak fernXXXOO

Notholaena standleyi Maxon star cloak fernXXXOOOOOO

Pellaea truncata Goodding spiny cliffbrakeXXXOOOOOOO

Pellaea wrightiana Hook. Wright’s cliffbrake

XXXOOOOOO

Pentagramma triangularis (Kaulfuss) Yatskievych, Windham & 

Wollenweber goldback fern

X

Pentagramma triangularis ssp. maxonii (Weatherby) Yatskievych, 

Windham & Wollenweber Maxon’s goldback fern

XXOO

Pentagramma triangularis ssp. triangularis (Kaulfuss) Yatskievych, 

Windham & Wollenweber goldback fern

XO

Pteridium aquilinum (L.) Kuhn western brackenfernXX

Pteridium aquilinum var. pubescens Underwood hairy brackenfernOXO

Selaginella underwoodii Hieron. Underwood’s spikemossOO

Pyrolaceae Chimaphila maculata (L.) Pursh striped prince’s pineX

Ranunculaceae Anemone tuberosa Rydb. tuber anemoneXXOOOX

Aquilegia chrysantha Gray golden columbineXXOO

Aquilegia desertorum (M.E. Jones) Cockerell ex Heller desert columbineX

Aquilegia triternata Payson Chiricahua Mountain columbineXO

Clematis drummondii Torr. & Gray Drummond’s clematisXX

Clematis ligusticifolia Nutt. western white clematisXXOOOX

Delphinium parishii Gray ssp. parishii Parish’s larkspurXX

Delphinium scaposum Greene tall mountain larkspurXXOOOO

Myosurus cupulatus S. Wats. Arizona mousetailOXXO

Ranunculus arizonicus J.G. Lemmon ex Gray Arizona buttercupXX

Thalictrum fendleri Engelm. ex Gray Fendler’s meadow-rueOXO

Thalictrum fendleri var. wrightii (Gray) Trel. Wright’s meadow-rueX

Rhamnaceae Ceanothus fendleri Gray Fendler’s ceanothusOXX

O

Ceanothus greggii Gray desert ceanothusOXX

Ceanothus integerrimus Hook. & Arn. deerbrushOXX

Condalia correllii M.C. Johnston Correll’s snakewoodXXO

Condalia warnockii M.C. Johnston Warnock’s snakewoodXXOOO

Condalia warnockii var. kearneyana M.C. Johnston Kearney’s snakewoodXOO

Frangula betulifolia ssp. betulifolia (Greene) V. Grub. beechleaf frangulaXXXO

Frangula californica ssp. californica (Eschsch.) Gray California buckthornXXOO
125

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Rhamnaceae Frangula californica ssp. ursina (Greene) Kartesz & Gandhi California buckthornX

Rhamnus crocea Nutt. redberry buckthornOXO

Rhamnus ilicifolia Kellogg hollyleaf redberryXOOOO

Ziziphus obtusifolia (Hook. ex Torr. & Gray) Gray lotebushOOOOOO

Ziziphus obtusifolia var. canescens (Gray) M.C. Johnston lotebushXXO

Rosaceae Agrimonia striata Michx. 

a roadside agrimony

X

Cercocarpus montanus Raf. alderleaf mountain mahoganyO

Fragaria vesca ssp. bracteata (Heller) Staudt woodland strawberryXX

Holodiscus discolor (Pursh) Maxim. oceansprayX

Holodiscus dumosus (Nutt. ex Hook.) Heller rockspireaXXO

Potentilla glandulosa Lindl. sticky cinquefoilO

Potentilla subviscosa var. ramulosa (Rydb.) Kearney & Peebles Navajo cinquefoilXXO

Potentilla thurberi var. atrorubens (Rydb.) Kearney & Peebles scarlet cinquefoilXO

Prunus serotina var. rufula (Woot. & Standl.) McVaugh black cherryXXO

Prunus serotina var. virens (Woot. & Standl.) McVaugh black cherryXO

Prunus virginiana L. chokecherryO

Rosa woodsii var. ultramontana (S. Wats.) Jepson Woods’ roseX

Rosa woodsii Lindl. var. woodsii 

a Woods’ rose

X

Rubus arizonensis Focke Arizona dewberryX

Rubus neomexicanus Gray New Mexico raspberryXXXO

Vauquelinia californica (Torr.) Sarg. Arizona rosewoodXXXOOO

OOO

Vauquelinia californica (Torr.) Sarg. ssp. californica Arizona rosewoodX

Rubiaceae Bouvardia ternifolia (Cav.) Schlecht. firecrackerbushXXXOOOOO

Cephalanthus occidentalis L. common buttonbushXXXOOX

Diodia teres Walt. poorjoeXXOO

Diodia teres var. angustata Gray poorjoeX

Galium aparine L. stickywillyXXOOOXOO

Galium boreale L. northern bedstrawO

Galium fendleri Gray Fendler’s bedstrawXXO

Galium mexicanum Kunth Mexican bedstrawXO

Galium mexicanum ssp. asperrimum (Gray) Dempster Mexican bedstrawXXOOO

Galium microphyllum Gray bracted bedstrawXXXOOOOO

Galium proliferum Gray limestone bedstrawXXXOOOO

Galium wrightii Gray Wright’s bedstrawXXXOOOO

Hedyotis greenei (Gray) W.H. Lewis Greene’s starvioletXXO

Houstonia pusilla Schoepf tiny bluetX

Houstonia wrightii Gray pygmy bluetXXO

Rutaceae Ptelea trifoliata L. common hoptreeO

Ptelea trifoliata ssp. angustifolia (Benth.) V. Bailey common hoptreeXX

Ptelea trifoliata var. cognata (Greene) Kearney & Peebles pallid hoptree

X

Thamnosma texana (Gray) Torr. rue of the mountainsXXX

Salicaceae Populus fremontii S. Wats. Fremont cottonwoodOOO

Populus fremontii S. Wats. ssp. fremontii Fremont cottonwoodXXOO

Populus tremuloides Michx. quaking aspenX
126

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Salicaceae Salix bonplandiana Kunth Bonpland willowXXX

Salix exigua Nutt. narrowleaf willowXXXOOOOO

Salix gooddingii Ball Goodding’s willowXXXOOOOO

Salix irrorata Anderss. dewystem willowX

Salix scouleriana Barratt ex Hook. Scouler’s willowX

Salix taxifolia Kunth yewleaf willowXXOO

Santalaceae Comandra umbellata (L.) Nutt. bastard toadflaxO

Comandra umbellata ssp. pallida (A. DC.) Piehl pale bastard toadflaxOXXOO

Sapindaceae Dodonaea viscosa (L.) Jacq. Florida hopbushXXXOOOO

Sapindus saponaria L. wingleaf soapberryXO

Sapindus saponaria var. drummondii (Hook. & Arn.) L. Benson western soapberryXXXOOOOO

Sapotaceae Sideroxylon lanuginosum Michx. gum bullyXOO

Sideroxylon lanuginosum ssp. rigidum (Gray) T.D. Pennington gum bullyX

Saxfragiaceae Fendlera rupicola Gray cliff fendlerbushXXXOO

Heuchera parvifolia var. arizonica Nutt. ex Torr. & Gray littleleaf alumrootO

Heuchera rubescens var. versicolor (Greene) M.G. Stewart pink alumrootX

Heuchera sanguinea Engelm. coralbells

OX

XOOOOO

Saxifraga eriophora S. Wats. redfuzz saxifrage

XXX

Scrophulariaceae Brachystigma wrightii (Gray) Pennell Arizona desert foxglove

XXXOO

Scrophulariaceae Castilleja austromontana Standl. & Blumer Rincon Mountain Indian paintbrush

XXOO

Castilleja exserta (Heller) Chuang & Heckard exserted Indian paintbrush

X

Castilleja exserta ssp. exserta (Heller) Chuang & Heckard exserted Indian paintbrush

OX

Castilleja integra Gray wholeleaf Indian paintbrush

XX

Castilleja lanata Gray Sierra woolly Indian paintbrush

XX

Castilleja minor (Gray) Gray lesser Indian paintbrush

XXXO

Castilleja tenuiflora Benth. Santa Catalina Indian paintbrush

XXXOOOO

Maurandella antirrhiniflora (Humb. & Bonpl. ex Willd.) Rothm. roving sailor

XXOOOOX

Mecardonia procumbens (P. Mill.) Small baby jump-up

XXOOOO

Mimetanthe pilosa (Benth.) Greene false monkeyflower

XXXO

Mimulus floribundus Lindl. manyflowered monkeyflower

XXX

Mimulus guttatus DC. seep monkeyflower

XXXOOOOOOO

Mimulus rubellus Gray little redstem monkeyflower

XXXOO

Nuttallanthus texanus (Scheele) D.A. Sutton Texas toadflax

XXXOOOXO

Pedicularis centranthera Gray dwarf lousewort

XX

Penstemon barbatus (Cav.) Roth beardlip penstemon

XOO

Penstemon barbatus (Cav.) Roth ssp. barbatus beardlip penstemon

X

Penstemon barbatus ssp. torreyi (Benth.) Keck Torrey’s penstemon

XO

Penstemon linarioides Gray toadflax penstemon

O

Penstemon parryi (Gray) Gray Parry’s beardtongue

OXXOOOOOO

Penstemon pseudospectabilis M.E. Jones desert penstemon

X

Penstemon pseudospectabilis ssp. connatifolius (A. Nels.) Keck desert beardtongue

X

Sairocarpus nuttallianus (Benth. ex A. DC.) D.A. Sutton violet snapdragon

OXXOOX

Schistophragma intermedia (Gray) Pennell harlequin spiralseed

XOO

Scrophularia parviflora Woot. & Standl. pineland figwort

XXXO

Stemodia durantifolia (L.) Sw. whitewoolly twintip

XXXOOO

Verbascum thapsus L. common mullein

O

Veronica anagallis-aquatica L. water speedwell

XX
127

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Scrophulariaceae Veronica peregrina L. neckweed

XXOO

Veronica peregrina ssp. xalapensis (Kunth) Pennell hairy purslane speedwell

XO

Selaginellaceae Selaginella arizonica Maxon Arizona spikemoss

XXXOOO

Selaginella rupincola Underwood rockloving spikemoss

OXXOOOO

Simmondsiaceae Simmondsia chinensis (Link) Schneid. jojoba

XOO

Solanaceae Chamaesaracha coronopus (Dunal) Gray greenleaf five eyes

XX

Chamaesaracha sordida (Dunal) Gray hairy five eyes

X

Datura discolor Bernh. desert thorn-apple

XXO

Datura wrightii Regel sacred thorn-apple

XO

Lycium andersonii Gray water jacket

XO

Lycium berlandieri Dunal Berlandier’s wolfberry

OOOOOO

Lycium berlandieri var. parviflorum (Gray) Terracc. Berlandier’s wolfberry

X

Lycium exsertum Gray Arizona desert-thorn

XXOOOO

Lycium pallidum Miers pale desert-thorn

XO

Margaranthus solanaceus Schlecht. netted globecherry

XX

Nicotiana obtusifolia var. obtusifolia Mertens & Galeotti desert tobacco

XOOOO

Physalis crassifolia Benth. yellow nightshade groundcherry

XXOO

Physalis crassifolia var. versicolor (Rydb.) Waterfall yellow nightshade groundcherry

XOO

Physalis hederifolia Gray ivyleaf groundcherry

OXOOO

Physalis hederifolia var. fendleri (Gray) Cronq. Fendler’s groundcherry

XXO

Physalis hederifolia var. hederifolia Gray ivyleaf groundcherry

XX

Physalis latiphysa Waterfall broadleaf groundcherry

X

Physalis pubescens L. husk tomato

X

Quincula lobata (Torr.) Raf. Chinese lantern

X

Solanum americanum P. Mill. American black nightshade

O

Solanum douglasii Dunal greenspot nightshade

XXXOOO

Solanum elaeagnifolium Cav. silverleaf nightshade

XOO

Solanum fendleri Gray ex Torr. Fendler’s horsenettle

XXO

Solanum nigrescens Mart. & Gal. divine nightshade

X

Sterculiaceae Ayenia compacta Rose California ayenia

OXOOOO

Ayenia filiformis S. Wats. TransPecos ayenia

XXO

Ayenia microphylla Gray dense ayenia

OOO

Waltheria indica L. uhaloa

XX

Tamaricaceae Tamarix aralensis Bunge Russian tamarisk

XX

Tamarix ramosissima Ledeb. saltcedar

XXO

Typhaceae Typha domingensis Pers. southern cattail

XXOOOOO

Ulmaceae Celtis laevigata var. reticulata (Torr.) L. Benson netleaf hackberry

XXXOOOO

Celtis pallida Torr. spiny hackberry

OXXOOOOOO

Urticaceae Parietaria hespera Hinton rillita pellitory

XXXOOOX

Parietaria hespera Hinton var. hespera rillita pellitory

X

Valerianaceae Plectritis ciliosa (Greene) Jepson longspur seablush

X

Plectritis ciliosa ssp. insignis (Suksdorf) Morey longspur seablush

XOO

Valeriana arizonica Gray Arizona valerian

X

Verbenaceae Aloysia wrightii Heller ex Abrams Wright’s beebrush

OXXOOOOOO

Glandularia bipinnatifida (Nutt.) Nutt. Dakota mock vervain

XXOO

Glandularia bipinnatifida var. bipinnatifida (Nutt.) Nutt. Dakota mock vervain

XXOOO

Glandularia gooddingii (Briq.) Solbrig southwestern mock vervain

OXXOO

Tetraclea coulteri Gray Coulter’s wrinklefruit

XX
128

Family Scientific name Common name UAI UAH B&M R&D FIa FIb FI F&B GU H&G LTM FEM

Violaceae Viola adunca Sm. hookedspur violet X

Viola affinis Le Conte sand violet X

Violaceae Viola canadensis L. Canadian white violet X X O

Viola nephrophylla Greene northern bog violet X X

Arceuthobium vaginatum (Willd.) J. Presl pineland dwarf mistletoe X X O

Viscaceae Phoradendron californicum Nutt. mesquite mistletoe O X X O O O O O

Phoradendron capitellatum Torr. ex Trel. downy mistletoe X X O

Phoradendron coryae Trel. Cory’s mistletoe X X O O O O

Phoradendron juniperinum Engelm. ex Gray juniper mistletoe X X O O O

Phoradendron leucarpum (Raf.) Reveal & M.C. Johnston oak mistletoe X

Phoradendron macrophyllum (Engelm.) Cockerell Colorado Desert mistletoe O O

Phoradendron macrophyllum (Engelm.) Cockerell ssp. 

macrophyllum Colorado Desert mistletoe X

Phoradendron pauciflorum Torr. fir mistletoe X X

Phoradendron tomentosum (DC.) Engelm. ex Gray Christmas mistletoe X X

Phoradendron villosum (Nutt.) Nutt. Pacific mistletoe O

Vitaceae Cissus trifoliata (L.) L. sorrelvine X X O O

Parthenocissus quinquefolia (L.) Planch. Virginia creeper X

Parthenocissus vitacea (Knerr) A.S. Hitchc. woodbine X

Vitis arizonica Engelm. canyon grape X X O O O O

Zygophyllaceae Kallstroemia californica (S. Wats.) Vail California caltrop X X

Kallstroemia grandiflora Torr. ex Gray Arizona poppy X X O

Kallstroemia parviflora J.B.S. Norton warty caltrop X X

Larrea tridentata (Sessé & Moc. ex DC.) Coville creosote bush X X O O O

Tribulus terrestris L. puncturevine X X X

a Cited as extirpated by Bowers and McLaughlin (1987) and found in the UA Herbarium. All specimens were collected by J.C. Blumer and have not been observed since then. We exclude them from 

the number of species found in the park. 

b Found along the Rincon Creek Trail (Danielle Foster, pers. comm.) 
129

Appendix B. List of amphibian and reptile species observed or documented at Saguaro National Park, Rincon Mountain District by UA inventory personnel (total 

number of observations; 2001-2002) or by other survey efforts or lists. Lowe and Holm (1991; L&H), Murray (1996; MU), Goode et al. (1998; GO), Bonine and Schwalbe 

(2003; B&S). Total number of observations for UA effort should not be used as a measure of relative abundance because these data have not been scaled by survey effort 

or area. Species in bold-faced type are non-native. See Appendices E and F for additional information on voucher specimens and photographs from UA inventory and other 

collections. 

Voucher 

Specimen 

(S),

Photo (P)

Order

 Family Scientific name Common name Intensive Extensive Road Incidental L&H MU Go B&S ESA BLM USFS AZ

Caudata

 Ambystomatidae Ambystoma tigrinum tiger salamanderb

Anura

 Pelobatidae Scaphiopus couchii Couch’s spadefoot 25 45 2 P, S X X

Spea multiplicata Mexican spadefoot 1 X

 Bufonidae Bufo alvarius Sonoran Desert toad 11 82 194 17 P, S X X X X

Bufo punctatus red-spotted toad 41 275 71 P, S X X X X

Bufo cognatus Great Plains toad 1 X X X

 Hylidae Hyla arenicolor canyon treefrog 2 168 80 P, S X X X

 Ranidae Rana yavapaiensis lowland leopard frog 100 37 P, S X X X X X X

Rana catesbeiana American bullfrog P, S X

Testudines

 Kinosternidae Kinosternon sonoriense Sonoran mud turtle 26 31 P, S X X X X

 Emydidae Terrapene ornata western box turtle Pc X

 Testudinidae Gopherus agassizii sonoran desert tortoise 1 14 13 P, S X X X X X X

Squamata

 Gekkonidae Coleonyx variegatus western banded gecko 1 11 29 4 P, S X X X

Hemidactylus turcicus Mediterranean house gecko Pc

, Sd

 Crotaphytidae Crotaphytus collaris eastern collared lizard 2 4 23 P, S X X X

Gambelia wislizenii long-nosed leopard lizard X X

 Phrynosomatidae Holbrookia maculata lesser earless lizard 3 5 P, S X

Cophosaurus texanus greater earless lizard 5 35 3 75 P, S X X X X

Callisaurus draconoides zebra-tailed lizard 61 47 P, S X X X

Sceloporus magister desert spiny lizard 22 89 15 P, S X X X

Sceloporus clarkii Clark’s spiny lizard 91 164 70 P, S X X X X

Sceloporus undulatus eastern fence lizard 39 113 79 P, S X

Uta stansburiana common side-blotched lizard 5 102 1 23 P, S X X X

Urosaurus ornatus ornate tree lizard 166 441 2 141 P, S X X X X

Phrynosoma hernandesi greater short-horned lizard 10 1 P, S X

 Phrynosomatidae Phrynosoma solare regal horned lizard 3 8 11 P, S X X X X

 Scincidae Eumeces obsoletus Great Plains skink 1 P X

 Teiidae Cnemidophorus burti canyon spotted whiptail 7 P, S X X X X X

UA Survey type Species list or study Conservation designationa
130

Voucher 

Specimen 

(S),

Photo (P)

Order

 Family Scientific name Common name Intensive Extensive Road Incidental L&H MU Go B&S ESA BLM USFS AZ

Squamata

 Teiidae Cnemidophorus sonorae Sonoran spotted whiptail 28 122 124 P, S X X X X

Cnemidophorus flagellicaudus Gila spotted whiptail 13 19 33 P, S X X

Cnemidophorus tigris western whiptail (tiger whiptail) 8 32 2 45 P X X X

 Anguidae Elgaria kingii Madrean alligator lizard 2 4 P, S X X

 Helodermatidae Heloderma suspectum Gila monster 12 6 25 P, S X X X

 Leptotyphlopidae Leptotyphlops humilis western blind snake Pc

, Sd X

 Colubridae Diadophis punctatus ring-necked snake 1 P X

Phyllorhynchus browni saddled leaf-nosed snake S X

Masticophis flagellum coachwhip 1 3 2 10 P, S X X X

Masticophis bilineatus Sonoran whipsnake 5 6 10 P, S X X X X

Salvadora hexalepis western patch-nosed snake 1 1 1 1 P, S X X

Salvadora grahamiae mountain patch-nosed snake 1 1 P, S X

Pituophis catenifer gopher snake 3 3 P, S X X

Arizona elegans glossy snake X

Lampropeltis getula common kingsnake 1 1 P, S X X X

Lampropeltis pyromelana Sonoran mountain kingsnake 2 1 P X

Rhinocheilus lecontei long-nosed snake 3 6 2 P X

Thamnophis cyrtopsis black-necked garter snake 5 65 38 P X X X X

Sonora semiannulata western ground snake 2 P, S X

Chilomeniscus cinctus variable sandsnake Pe

, S X X

Tantilla hobartsmithi southwestern black-headed snake S X X

Trimorphodon biscutatus western lyre snake Sd X X

Hypsiglena torquata night snake 6 1 P, S X X

 Elapidae Micruroides euryxanthus Sonoran coral snake 1 1 P, S X

 Viperidae Crotalus atrox western diamond-backed rattlesnake 1 48 6 17 P X X X X

Crotalus molossus black-tailed rattlesnake 1 13 1 12 P, S X X X

Crotalus tigris tiger rattlesnake 1 15 8 10 P, S X X X

Crotalus viridis western rattlesnake 2 11 16 P, S X

Crotalus scutulatus Mojave rattlesnake 1 S X

a

 ESA = Species of Concern, Endangered Species Act, U.S. Fish and Wildlife Service (in HDMS 2004); BLM = Bureau of Land Management, “sensitive” species; USFS = U. S. Forest Service, 

sensitive species; AZ = Arizona Game and Fish, “Wildlife of Special Concern”. Data from HDMS (2004).

b Observed by Danielle Foster near Rincon Creek in 2001. 

c Don Swann has a photograph from the park in his collection.

d Voucher specimen collected by Don Swann and not yet accessioned into the UA herpetology collection (D. Swann, pers. comm.). e Photograph by Matt Goode (1997) along the Loop Drive (photograph now accessioned in the I&M office in Tucson). 

UA survey type Species list or study Conservation designationa
131

Appendix C. List of bird species observed at Saguaro National Park, Rincon Mountain District by UA inventory personnel (2001-2003) or by other survey efforts or 

lists. Marshall (1956; MA), Monson and Smith (1985; M&S), Freiderici (1998; FR), Boal and Mannan (1996; B&M), Short (1996; SH), Powell (1999; P99), and Powell (2004; 

P04). See text for descriptions of UA survey types. Underlined species (scientific names) are neotropical migrants (Rappole 1995) and species in bold-faced type are non-native. 

Underlined “X” or number in UA incidental column indicates evidence of breeding was observed during that study (see Table 5.8 for breeding observations by UA personnel). 

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Anseriformes

 Anatidae Anas platyrhynchos mallard 4 1

Galliformes

 Phasianidae Meleagris gallopavo wild turkey 1 X

 Odontophoridae Callipepla squamata scaled quail 1 X

Callipepla gambelii Gambel’s quail 475 89 12 X X X X X X

Cyrtonyx montezumae Montezuma quail 13 28 X X

Ciconiiformes

 Cathartidae Coragyps atratus black vulture X

Cathartes aura turkey vulture 76 26 X X X X X X X

Falconiformes

 Accipitridae Pandeon haliaetus osprey X

 Circus cyaneus northern harrier X X

Accipiter striatus sharp-shinned hawk 1 2 X S

Accipiter cooperii Cooper’s hawk 21 2 9 X X X X X X X

Accipiter gentilis northern goshawk 5 14 X SC S WSC

Asturina nitida gray hawk 4 1 X SC S WSC

Buteogallus anthracinus common black-hawk 1 X X S WSC P

Parabuteo unicinctus Harris’s hawk 1 X X X

Buteo swainsoni Swainson’s hawk X

Buteo albonotatus zone-tailed hawk 9 18 X X X X

Buteo jamaicensis red-tailed hawk 26 5 19 X X X X X X X

Buteo regalis ferruginous hawk X X SC WSC

Aquila chrysaetos golden eagle 4 2 X X

 Falconidae Falco sparverius American kestrel 14 4 4 X X X X X X

Falco peregrinus peregrine falconb 4 5 X X X SC WSC BCC

Falco mexicanus prairie falcon 6 X X X

Falco columbarius Merlinc

Charadriiformes

 Charadriidae Charadrius vociferus killdeer X

 Scolopacidae Actitis macularia spotted sandpiper X

Columbiformes

 Columbidae Columba livia rock pigeon 4

 Patagioenas fasciata band-tailed pigeon 15 12 X X X

 Zenaida asiatica white-winged dove 872 5 X X X X X X X

Zenaida macroura mourning dove 651 12 20 X X X X X

 Columbina inca Inca dove X X X
132

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Columbiformes

 Columbidae Columbina passerina common ground-dove 13 4 X X

Cuculiformes

 Cuculidae Coccyzus americanus 

occidentalis yellow-billed cuckoo 1 1 X X C S WSC P BCC

Geococcyx californianus greater roadrunner 10 13 X X X X X X X

Strigiformes

 Tytonidae Tyto alba barn owl 1 X X X

 Strigidae Otus flammeolus flammulated owl 7 4 X X

Megascops kennicottii western screech-owl 49 3 X X X X

Megascops trichopsis whiskered screech-owl 1 13 6 X X X

Bubo virginianus great horned owl 20 2 14 25 X X X X X X

Glaucidium gnoma northern pygmy-owl 2 1 X X

Glaucidium brasilianum 

cactorum cactus ferruginous pygmy-owld

Athene cunicularia 

hypugaea burrowing owl X SC

Micrathene whitneyi elf owl 116 16 X X X X X BCC

Strix occidentalis lucida Mexican spotted owl 2 4 X X LT S WSC

Asio otus long-eared owl 1 X

Caprimulgiformes

 Caprimulgidae Chordeiles acutipennis lesser nighthawk 2 3 3 X X X X X

Phalaenoptilus nuttallii common poorwill 6 31 20 X X X X X X

Caprimulgus vociferus whip-poor-will 4 26 11 X X X

Apodiformes

 Apodidae Chaetura vauxi Vaux’s swift X

Aeronautes saxatalis white-throated swift 44 15 X X X X X

 Trochilidae Cynanthus latirostris broad-billed hummingbird 36 11 X X X X X

Eugenes fulgens magnificent hummingbird 4 4 X X

Archilochus alexandri black-chinned hummingbird 67 12 X X X X X X X

Calypte anna Anna’s hummingbird 22 1 19 X X

Calypte costae Costa’s hummingbird 18 1 X X X X P

Stellula calliope calliope hummingbird X

Selasphorus platycercus broad-tailed hummingbird 46 10 X X X

Selasphorus rufus rufous hummingbird 4 1 X

Trogoniformes

 Trogonidae Trogon elegans elegant trogon 4 1 WSC

Coraciiformes

 Alcedinidae Ceryle alcyon belted kingfisher 1 2 X WSC

 Picidae Melanerpes formicivorus acorn woodpecker 62 21 X X X

 Melanerpes uropygialis Gila woodpecker 469 69 7 X X X X X X BCC

Sphyrapicus thyroideus Williamson’s sapsucker X
133

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Coraciiformes

 Picidae Sphyrapicus varius yellow-bellied sapsucker X

Sphyrapicus nuchalis red-naped sapsucker 1 3

Picoides scalaris ladder-backed woodpecker 133 29 3 X X X X X X X

Picoides villosus hairy woodpecker 35 12 X X X

Picoides arizonae Arizona woodpecker 12 13 X X

Colaptes auratus northern flicker 53 13 6 X X X X X X

Colaptes chrysoides gilded flicker 63 15 5 X X X X P BCC

Passeriformes

 Tyrannidae Camptostoma imberbe northern beardless-tyrannulet 22 3 X X X X

Contopus cooperi olive-sided flycatcher 1 X SC

Contopus pertinax greater pewee 35 7 X X X

Contopus sordidulus western wood-pewee 105 73 X X X

Empidonax traillii willow flycatcher 2 X WSC

Empidonax hammondii Hammond’s flycatcher 3 X

Empidonax wrightii gray flycatcher 6 1 X X X

Empidonax oberholseri dusky flycatcher 1 X X

Empidonax difficilis pacific-slope flycatcher 1

Empidonax fulvifrons buff-breasted flycatcher 4 Xi SC WSC

Empidonax occidentalis cordilleran flycatcher 60 7 X

Sayornis nigricans black phoebe 14 1 14 X X X X X

Sayornis saya Say’s phoebe 2 5 X X X X

Pyrocephalus rubinus vermilion flycatcher 25 4 X X X X X

Myiarchus tuberculifer dusky-capped flycatcher 63 12 X X X

Myiarchus cinerascens ash-throated flycatcher 462 1 26 X X X X X X X

Myiarchus tyrannulus brown-crested flycatcher 297 16 X X X X X X

Myiodynastes luteiventris sulphur-bellied flycatcher 8 6

Tyrannus vociferans Cassin’s kingbird 48 30 X X X X X

Tyrannus verticalis western kingbird 15 2 X X X

 Laniidae Lanius ludovicianus loggerhead shrike 2 2 8 X X X X SC S

 Vireonidae Vireo bellii Bell’s vireo 194 28 X X X X X X S BCC

Vireo vicinior gray vireo 1 X X

 Vireonidae Vireo plumbeus plumbeous vireo 66 17 X X X

Vireo huttoni Hutton’s vireo 54 33 X X X X X

Vireo gilvus warbling vireo 13 7 X X X X X

 Corvidae Cyanocitta stelleri Steller’s jay 62 4 X X X

 Aphelocoma californica western scrub-jay 33 23 12 X X X X

Aphelocoma ultramarina Mexican jay 207 6 46 X X

 Gymnorhinus 

cyanocephalus pinyon jay 1

Nucifraga columbiana Clark’s nutcracker X

Corvus corax common raven 71 14 19 X X X X X X X
134

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Passeriformes

 Hirundinidae Progne subis purple martin 151 16 X X X X X X P

Tachycineta bicolor tree swallow X

Tachycineta thalassina violet-green swallow 74 4 23 X X X X X

Stelgidopteryx serripennis northern rough-winged swallow 2 X

Petrochelidon pyrrhonota cliff swallow X

 Paridae Poecile gambeli mountain chickadee 60 12 X X X

Baeolophus wollweberi bridled titmouse 50 22 45 X X X X X

Baeolphus ridgwayi juniper titmouse 2 1 X

 Remizidae Auriparus flaviceps verdin 359 53 6 X X X X X X

 Aegithalidae Psaltriparus minimus bushtit 105 33 72 X X X X

 Sittidae Sitta canadensis red-breasted nuthatch 7 X

Sitta carolinensis white-breasted nuthatch 87 1 18 X X X

Sitta pygmaea pygmy nuthatch 24 5 X X X

 Certhiidae Certhia americana brown creeper 18 4 X X X

 Troglodytidae Campylorhynchus 

brunneicapillus cactus wren 408 90 3 X X X X X X

Salpinctes obsoletus rock wren 54 7 24 X X X X X X X

Catherpes mexicanus canyon wren 159 2 17 X X X X X X X

Thryomanes bewickii Bewick’s wren 474 59 83 X X X X X X X

Troglodytes aedon house wren 51 8 14 X X X X X

 Regulidae Regulus calendula ruby-crowned kinglet 3 56 1 X X X X

Polioptila caerulea blue-gray gnatcatcher 71 28 X X X X

Polioptila melanura black-tailed gnatcatcher 78 16 9 X X X X X X

 Turdidae Sialia mexicana western bluebird 26 41 20 X X X X X X

Sialia currucoides mountain bluebird X

Sialia sialia eastern bluebird j

 Myadestes townsendi Townsend’s solitaire 17 1 X X X

Catharus guttatus hermit thrush 64 2 17 X X X X X

 Turdidae Turdus migratorius American robin 48 5 4 X X X

 Mimidae Mimus polyglottos northern mockingbird 104 3 5 X X X X X X

Oreoscoptes montanus sage thrasher X

Toxostoma bendirei Bendire’s thrasher X X

Toxostoma curvirostre curve-billed thrasher 207 61 14 X X X X X X

Toxostoma crissale crissal thrasher 11 9 9 X X X X X X X BCC

 Sturnidae Sturnus vulgaris European starling 3 X X X X X

 Bombycillidae Bombycilla cedrorum cedar waxwing 4 22 5 X

 Ptilogonatidae Phainopepla nitens phainopepla 99 10 10 X X X X X X

 Peucedramidae Peucedramus taeniatus olive warbler 23 3 X X X

 Parulidae Vermivora celata orange-crowned warbler 8 1 X

Vermivora ruficapilla Nashville warbler 2 X

Vermivora virginiae Virginia’s warbler 41 24 X X X
135

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Passeriformes

 Parulidae Vermivora luciae Lucy’s warbler 316 7 X X X X X X P

Dendroica petechia yellow warbler 27 X X X

Dendroica coronata yellow-rumped warbler 48 3 10 X X X X X

Dendroica nigrescens black-throated gray warbler 145 31 X X X

Dendroica townsendi Townsend’s warbler 16 4 X X

Dendroica occidentalis hermit warbler 1 X

Dendroica graciae Grace’s warbler 86 16 X X X X

Oporornis tolmiei MacGillivray’s warbler 4 X

Wilsonia pusilla Wilson’s warbler 31 7 X X X

Cardellina rubrifrons red-faced warbler 45 14 X X X

Myioborus pictus painted redstart 16 15 X X X

Icteria virens yellow-breasted chat 1 X

 Thraupidae Piranga flava hepatic tanager 61 35 X X X

Piranga rubra summer tanager 42 4 X X X X X

Piranga ludoviciana western tanager 116 29 X X X X X X

 Emberizidae Pipilo chlorurus green-tailed towhee 33 48 5 X X

Pipilo maculatus spotted towhee 265 28 62 X X X X

Pipilo fuscus canyon towhee 188 79 5 X X X X X

Pipilo aberti Abert’s towhee 55 8 19 X X X X X X

Aimophila carpalis rufous-winged sparrow 74 78 27 X X X X X P BCC

Aimophila cassinii Cassin’s sparrow 1 X

 Aimophila ruficeps rufous-crowned sparrow 235 25 37 X X X X X

Spizella passerina chipping sparrow 29 228 47 X X X X X

Spizella breweri Brewer’s sparrow 31 42 81 X X X X

Spizella atrogularis black-chinned sparrow 68 3 43 X X

Passerculus sandwichensis Savannah sparrow X X

Pooecetes gramineus vesper sparrow 1 4 1 X

Chondestes grammacus lark sparrow 13 30 X X X X X

Amphispiza bilineata black-throated sparrow 209 73 13 X X X X X

Amphispiza belli sage sparrow X

Calamospiza melanocorys lark bunting X X

Passerella iliaca fox sparrow X

 Melospiza melodia song sparrow 1 X

Melospiza lincolnii Lincoln’s sparrow 2 6 5 X X X

Zonotrichia albicollis white-throated sparrow X

 Zonotrichia leucophrys white-crowned sparrow 9 26 7 X X X X

Zonotrichia atricapilla golden-crowned sparrow X

Junco hyemalis dark-eyed junco 27 2 X X X X

Junco phaeonotus yellow-eyed junco 127 30 X X X

 Cardinalidae Cardinalis cardinalis northern cardinal 229 26 9 X X X X X X
136

UA survey type Survey or species lists Conservation designationa

Order

 Family Scientific name Common name VCP

line 

transect

NocturnalIncidental MA M&S FR B&M SH P99 P04 ESA USFS AZGF APF USFWS

Passeriformes

 Cardinalidae Cardinalis sinuatus pyrrhuloxia 31 6 9 X X X X X X

Pheucticus ludovicianus rose-breasted grosbeak X

Pheucticus 

melanocephalus black-headed grosbeak 138 26 X X X X X

Passerina caerulea blue grosbeak 49 6 X X X

Passerina amoena lazuli bunting 13 X X X X

Passerina cyanea indigo bunting 1 5

Passerina versicolor varied bunting 37 12 X X X X

 Icteridae Agelaius phoeniceus red-winged blackbird X

Sturnella magna eastern meadowlark 5 1

Sturnella neglecta western meadowlark X

Euphagus cyanocephalus Brewer’s blackbird X

Quiscalus mexicanus great-tailed grackle 1

Molothrus aeneus bronzed cowbird 1 X X X X X X

Molothrus ater brown-headed cowbird 202 11 X X X X X

Icterus cucullatus hooded oriole 9 5 X X X X X

Icterus bullockii Bullock’s oriole 13 8 X

Icterus parisorum Scott’s oriole 106 22 X X X X X

 Fringillidae Carpodacus cassinii Cassin’s finch X

Carpodacus mexicanus house finch 424 71 16 X X X X X X X

Loxia curvirostra red crossbill 12 X

Carduelis pinus pine siskin 16 5 1 X X

Carduelis psaltria lesser goldfinch 151 27 21 X X X X X X X

 Carduelis lawrencei Lawrence’s goldfinch 1 X

Coccothraustes vespertinus evening grosbeak X X

 Passeridae Passer domesticus house sparrow 2 X X X

a

 ESA = Endangered Species Act; U.S. Fish and Wildlife Service; “SC” = “Species of Concern”; “C” = Candidate for listing, “LT” = Listed as Threatened (HDMS 2004). USFS = U.S.D.A. Forest 

Service, “Sensitive species” (HDMS 2004). WSCA = Arizona Game and Fish Department, “Wildlife of Special Concern” (HDMS 2004). APF = Arizona Partners in Flight, “Priority species”; (Latta et al. 

1999). U.S. Fish and Wildlife Service, “Species of conservation concern” (HDMS 2004). b Known to breed in the park (Bailey 1994).

c Found by Jeff Kartheiser.

d

 Confirmed in Box Canyon on 12 October 1995 by Andy and Tani Hubbard. 

e

 Marshall did not observe this species, but reports that Herbert Brown collected a specimen at Manning Camp on 18 August 1911. The current location of specimen is unknown. f

 Seen by Dan Herrington near Madrona Ranger Station, May 1998.
137

Appendix D. Number of observations of mammal species by University of Arizona and Saguaro National Park Inventory personnel by survey type, Saguaro National 

Park, Rincon Mountain District, 2001 and 2002 (small mammals, bats, and observations of all taxa) and 1999-2005 (infrared-triggered photography). Numbers of 

observations are not scaled by search effort and should not be used for comparison among species or survey types. See Appendices E and F for additional information on 

voucher specimens and photographs. Legacy data from: Sumner (1951; SU), Duncan (1990; DU), Davis and Sidner (1992; D&S), Sidner and Davis (1994; S&D), Bucci (2001; 

BU), Sidner (2003; SI) Swann (2003; SW). Survey type = small mammal trapping (SMT), bat netting (BN), infrared-triggered photography (ITP), and incidental observations (IO). 

Species in bold-faced type are non-native. 

Survey type

Documentation type Survey or species lists Conservation designation

Order 

 Family Scientific name Common name SMT BN ITP IO

PhotographSpecimen SU DU D&S S&D BU SI SWU.S.

FWSa BLMb

U.S.

FSc

AZ 

G&Fd

Didelphimorphia 

 Didelphidae Didelphis virginiana Virginia opossum 2 1 X

Insectivora

 Soricidae Notiosorex crawfordi Crawford’s desert shrew X X

Notiosorex cockrumi Cockrum’s desert shrew 5

Chiroptera

 Phyllostomidae Choeronycteris mexicana Mexican long-tongued bat 1 X X SC S WSC

Leptonycteris curasoae yerbabuenae southern long-nosed bat 1 1 X LE S WSC

 Vespertilionidae Myotis yumanensis Yuma myotis X X SC

Myotis auriculus southwestern myotis 4 2 X X

Myotis velifer cave myotis 4 X X SC S

Myotis thysanodes fringed myotis 2 2 X X SC S

Myotis volans long-legged myotis 3 1 X X SC S

Myotis californicus California myotis 8 3 3 X X X

Myotis ciliolabrum western small-footed myotis X X SC S

Lasionycteris noctivagans silver-haired bat 4 1 X X

Pipistrellus hesperus western pipistrelle X X

Eptesicus fuscus big brown bat 55 1 X X X

Lasiurus blossevillii western red bat 1 2 WSC

Lasiurus cinereus hoary bat 7 1 X X X

Corynorhinus townsendii pallescens Townsend’s big-eared bat 1 X X SC

Antrozous pallidus pallid bat 1 X X

 Molossidae Tadarida brasiliensis Brazilian free-tailed bat 16 1 X X X

Nyctinomops femorosaccus pocketed free-tailed bat 2 2 1 X S

Carnivora 

 Ursidae Ursus americanus American black bear 34 6 1 X

 Procyonidae Procyon lotor northern raccoon 7 1 X X

Nasua narica white-nosed coati 30 2 2 X X

Bassariscus astutus ringtail 229 1 1 X X X

 Mustelidae Taxidea taxus American badger 1

 Mephitidae Spilogale gracilis western spotted skunk 6 1

 Mephitidae Mephitis mephitis striped skunk 165 7 1 X X

Mephitis macroura hooded skunk 185 1 1 X

Conepatus mesoleucus white-backed hog-nosed skunk 27 1 1 X

 Canidae Canis familiaris feral dog 4 1

Canis latrans coyote 120 4 1 X X X X

Urocyon cinereoargenteus common gray fox 1029 11 1 X X X
138

Survey type

Documentation type Survey or species lists Conservation designation

Order 

 Family Scientific name Common name SMT BN ITP IO

PhotographSpecimen SU DU D&S S&D BU SI SWU.S.

FWSa BLMb

U.S.

FSc

AZ 

G&Fd

 Felidae Felis catus feral cat 1

Puma concolor azteca mountian lion 75 9 1 1 X X

Lynx rufus bobcat 57 3 2 X X X X

Rodentia 

 Sciuridae Spermophilus variegatus rock squirrel 1 15 9 1 X X X

Spermophilus tereticaudus round-tailed ground squirrel 1 1 X

Ammospermophilus harrisii Harris’ antelope squirrel 7 4 1 X X

Neotamias dorsalis cliff chipmunk 39 16 X X X

Sciurus aberti Abert’s squirrel 1 9 7 1 X

Sciurus arizonensis Arizona gray squirrel 3 1 1 X SC

 Geomyidae Thomomys bottae Botta’s pocket gopher 1 1 X X X

 Heteromyidae Perognathus amplus Arizona pocket mouse X X

Chaetodipus penicillatus Sonoran Desert pocket mouse 42 3 X X X

Chaetodipus intermedius rock pocket mouse 115 1 X X X SC

Chaetodipus baileyi Bailey’s pocket mouse 13 X X X X

Dipodomys merriami Merriam’s kangaroo rat 9 X X

 Muridae Reithrodontomys megalotis western harvest mouse 4 1 X SC

Reithrodontomys fulvescens fulvous harvest mouse 4 2

Peromyscus eremicus cactus mouse 35 4 X X X X SC S

Peromyscus boylii brush mouse 165 1 6 X X

Onychomys torridus southern grasshopper mousee

Neotoma albigula western white-throated woodrat 75 14 2 X X X X

Neotoma mexicana Mexican woodrat 25 X SC

Sigmodon ochrognathus yellow-nosed cotton rat 12 X X SC

Sigmodon arizonae Arizona cotton rat 4 1 X X

Lagomorpha

 Leporidae Lepus alleni antelope jackrabbit 7 2 1 X X

Lepus californicus black-tailed jackrabbit 10 1 1 X X X

Sylvilagus floridanus eastern cottontail 2 X

Sylvilagus audubonii desert cottontail 21 1 X X X

Artiodactyla

 Bovidae Bos taurus domestic cattle 3 1 1 X

 Tayassuidae Pecari tajacu collared peccary 980 17 1 3 X X X X

 Cervidae Odocoileus hemionus mule deer 37 3 1 X X X X

Odocoileus virginianus white-tailed deer 202 40 1 X X X

a

 LE = “Listed Endangered”, SC = “Species of Concern”; U.S. Fish and Wildlife Service (HDMS 2004); * Eastern cottontail not confirmed; see text. 

b

 “Sensitive species”; Bureau of Land Management (HDMS 2004).

c

 “Sensitive species”; U.S. Forest Service (HDMS 2004).

d “Wildlife of special concern”; Arizona Game and Fish Department (HDMS 2004).

e

 Confirmed by roadkilled animal. See text for more information. 

f

 Observed in the mid 1990’s, but not since. May be extirpated. See text for more information. 
139

Appendix E. Vertebrate specimen and photograph vouchers collected by University of Arizona or park personnel, 

Saguaro National Park, Rincon Mountain District, 1997–2002. All specimen vouchers are located in the University of 

Arizona (AZ) collections. Unless otherwise indicated, all photographic vouchers are located in the I&M office in Tucson. 

Voucher 

type Taxon Species Collector/photographer

Collection 

 date

 AZ 

collection # Specimen type

Specimen Amphibian red-spotted toad Don E. Swann 07/29/99 54002 whole

American bullfrog Dan M. Bell 08/18/97 whole

Sonoran mud turtle Kevin E. Bonine 07/04/02 54632 whole

Don E. Swann 12/22/01 54001 whole

J. Moorbeck whole

Reptile desert tortoise Don E. Swann 07/30/99 whole

Don E. Swann 09/30/97 54658 whole

Kevin E. Bonine 07/30/02 whole

lesser earless lizard Mike D. Wall 05/24/01 53122 whole

zebra-tailed lizard Don E. Swann 01/11/02 54011 whole

desert spiny lizard Jay Loughlin 05/02/01 54010 whole

canyon spotted whiptail Dale S. Turner 03/23/01 53686 whole

coachwhip Chris K. Kirkpatrick 07/07/01 53640 whole

mountain patch-nosed snake Dave B. Prival 05/15/01 53089 whole

gopher snake Dale S. Turner 03/23/01 53684 whole

common kingsnake Don E. Swann 07/12/02 54005 whole

variable sandsnake Don E. Swann 02/19/02 54004 whole

southwestern black-headed snake Don E. Swann 07/12/99 54006 whole

western diamond-backed rattlesnake James E. Borgmeyer 05/02/01 53646 whole

Mojave rattlesnake Brian F. Powell 04/04/01 52449 whole

Mammal unknown desert shrew Ronnie Sidner 09/24/01 26913 Skull and Skeleton

Ronnie Sidner 09/13/01 26911 Skull and Skeleton

Ronnie Sidner 09/13/01 26915 Skull and Skeleton

Ronnie Sidner 09/13/01 26910 Skull and Skeleton

Neil D. Perry 07/24/01 Skull

southern long-nosed bat Ronnie Sidner 05/13/01 Lost by museum?

California myotis Ronnie Sidner 05/05/02 26854 Skin and Skull

Ronnie Sidner 09/23/01 26946 Skin and Skull

Ronnie Sidner 09/18/01 26855 Skin and Skull

pocketed free-tailed bat Ronnie Sidner 09/30/02 26856 Skin and Skull

ringtail Neil D. Perry 07/16/02 26769 Skull

common gray fox Jason A. Schmidt 05/01/01 26774 Skull

mountain lion Ronnie Sidner 06/02/01 26756 Skull

Botta’s pocket gopher Neil D. Perry 10/10/01 27040 Skull

Sonoran Desert pocket mouse Neil D. Perry 04/15/02 26916 Skull

Neil D. Perry 04/17/02 26868 Skin and Skull

Neil D. Perry 04/17/02 26888 Skin and Skull

rock pocket mouse Neil D. Perry 10/31/02 26921 Skull

western harvest mouse Neil D. Perry 07/23/01 26827 Skin and Skull

fulvous harvest mouse Neil D. Perry 04/11/02 26895 Skin and Skull

Neil D. Perry 04/16/02 26887 Skin and Skull

cactus mouse Neil D. Perry 04/15/02 26852 Skin and Skull

Neil D. Perry 05/14/01 27039 Skin and Skull

Neil D. Perry 04/11/02 26894 Skin and Skull

Neil D. Perry 04/11/02 26893 Skin and Skull

brush mouse Jason A. Schmidt 06/10/01 26837 Skin and Skull

Neil D. Perry 06/10/01 26889 Skin and Skull

Neil D. Perry 06/06/01 26826 Skin and Skull

Ronnie Sidner 09/13/01 26901 Skull

Specimen Mammal western white-throated woodrat Neil D. Perry 05/15/01 26857 Skin and Skull

Arizona cotton rat Neil D. Perry 05/13/01 26833 Skin and Skull

collared peccary Jason A. Schmidt 05/09/01 26760 Skull

Neil D. Perry 06/07/01 26772 Skull

Photograph Amphibian Couch’s spadefoot Dave B. Prival 08/14/01

Sonoran desert toad Dave B. Prival 07/09/01

red-spotted toad Dave B. Prival 04/29/01
140

Voucher 

type Taxon Species Collector/photographer

Collection 

 date

 AZ 

collection # Specimen type

Photograph Amphibian canyon treefrog Dave B. Prival 07/27/01

lowland leopard frog Dave B. Prival 06/25/01

Dave B. Prival 04/29/01

Reptile Sonoran mud turtle Dave B. Prival 05/01/01

desert tortoise Dave B. Prival 07/29/01

western banded gecko Dave B. Prival 06/28/01

eastern collared lizard Dave B. Prival 04/29/01

lesser earless lizard Dave B. Prival 04/30/01

greater earless lizard Dave B. Prival 04/29/01

zebra-tailed lizard Dave B. Prival 10/05/01

desert spiny lizard Dave B. Prival 04/04/01

Clark’s spiny lizard Mike D. Wall 06/27/01

eastern fence lizard Dave B. Prival 05/09/01

common side-blotched lizard Dave B. Prival 10/05/01

ornate tree lizard Dave B. Prival 04/29/01

greater short-horned lizard Dave B. Prival 05/12/01

pygmy short-horned lizard Neil D. Perry 08/19/01

regal horned lizard Dave B. Prival 06/27/01

Great Plains skink Dave B. Prival 06/25/01

canyon spotted whiptail Dan M. Bell 09/04/99

Sonoran spotted whiptail Dave B. Prival 06/28/01

Gila spotted whiptail Cecil R. Schwalbe 10/19/99

Dave B. Prival 06/28/01

Dave B. Prival 07/25/01

western whiptail (tiger whiptail) Dave B. Prival 10/05/01

Madrean alligator lizard Dave B. Prival 08/09/01

Gila monster Dave B. Prival 05/01/01

western blind snake Matt J. Goode 08/08/97

coachwhip Don E. Swann 10/01/01

Sonoran whipsnake Dave B. Prival 05/09/01

western patch-nosed snake Dave B. Prival 08/14/01

mountain patch-nosed snake Dave B. Prival 05/15/01

gopher snake Dave B. Prival 05/24/01

common kingsnake Dave B. Prival 06/26/01

Sonoran mountain kingsnake Dave B. Prival 05/11/01

long-nosed snake Dave B. Prival 05/02/01

black-necked garter snake Dave B. Prival 05/08/01

western ground snake Dave B. Prival 08/09/01

night snake Mike D. Wall 05/01/01

Sonoran coral snake Dave B. Prival 07/27/01

western diamond-backed rattlesnake Dave B. Prival 07/09/01

black-tailed rattlesnake Dave B. Prival 04/30/01

tiger rattlesnake Dave B. Prival 05/10/01

western rattlesnake Dave B. Prival 06/27/01

Mammala Virginia opossum Don Swann 12/08/99

southwestern myotis Ronnie Sidner 09/23/01

PhotographMammal fringed myotis Ronnie Sidner 05/19/01

long-legged myotis Ronnie Sidner 05/21/01

California myotis Ronnie Sidner 09/22/01

silver-haired bat Ronnie Sidner 05/20/01

big brown bat Ronnie Sidner 09/22/01

hoary bat Ronnie Sidner 05/17/01

Brazilian free-tailed bat Ronnie Sidner 05/20/01

pocketed free-tailed bat Ronnie Sidner 05/17/01

American black bear Don E. Swann 11/05/04

northern raccoon Don E. Swann 03/06/03

white-nosed coati Don E. Swann 10/21/02

ringtail Don E. Swann 08/12/99

western spotted skunk Don E. Swann 12/17/02

striped skunk Don E. Swann 09/21/99
141

Voucher 

type Taxon Species Collector/photographer

Collection 

 date

 AZ 

collection # Specimen type

PhotographMammal hooded skunk Don E. Swann 12/13/04

white-backed hog-nosed skunk Don E. Swann 11/21/02

feral dog Don E. Swann 03/00/05

coyote Don E. Swann 12/00/02

common gray fox Don E. Swann 12/19/99

feral cat Don E. Swann 02/23/05

mountain lion Neil D. Perry 08/19/01

bobcat Neil D. Perry 07/19/01

rock squirrel Neil D. Perry 06/17/01

round-tailed ground squirrel Don E. Swann 11/02/00

Harris’ antelope squirrel Don E. Swann 04/08/00

Abert’s squirrel Don E. Swann 08/12/00

Arizona gray squirrel Don E. Swann 12/10/04

antelope jackrabbit Don E. Swann 08/21/01

black-tailed jackrabbit Don E. Swann 08/15/01

eastern cottontail Don E. Swann 09/30/03

desert cottontail Don E. Swann 11/14/00

domestic cattle Don E. Swann 02/14/00

collared peccary Don E. Swann 11/28/00

mule deer Don E. Swann 11/04/01

white-tailed deer Don E. Swann 04/28/04

a Photographs taken by Don E. Swann are located at the park headquarters.
142

Appendix F. List of existing voucher specimens collected prior to this inventory effort. See Table 1.1 for list of 

collections queried for these data. 

Taxon Common name Collectiona Collection number

Collection 

date Collector

Amphibian Couch’s spadefoot NPS 562 1964 B. A Lund

570 1977 W. F. Steenbergh

Sonoran desert toad FWMSH 1981 Tim Jones

red-spotted toad NPS 559, 565 to 569 1964 B. A. Lund

canyon treefrog NPS 555, 570, 611 1965-1968 B. A. Lund, H. Coss

lowland leopard frog UA 43205 1979

Reptile western banded gecko UA 1126 1960

eastern collared lizard UA 47101 1986

lesser earless lizard NPS 557 1966 B. A. Lund

greater earless lizard NPS 550, 551 1966 B. A. Lund

zebra-tailed lizard NPS 552 1966 B. A. Lund

eastern fence lizard USNM 042548, 048547

desert spiny lizard UA 10105 1963

Clark’s spiny lizard UA 2258 1960

common side-blotched lizard NHMLAC 98183

OMNH 30000 1959

UA 45649 1984

ornate tree lizard UA 3759 1960

pygmy short-horned lizard USNM 039311, 039312, 048549 1911 H. Brown

regal horned lizard UA 32291 1968

Sonoran spotted whiptail UA 4809 1968

Gila spotted whiptail UA 54480 1998

Madrean alligator lizard UA 7249 1959

Gila monster AU 1994 1959 G. Folkerts

saddled leaf-nosed snake UA 30838 1969

Sonoran whipsnake NPS 579, 581 1963 R. Fabel, R. Lutz, S. 

Ferguson

western patch-nosed snake AU 1931 1959 R. Faber

gopher snake NPS 1280 1977 K. Black

western ground snake UA 26361 1959

variable sandsnake UA 35166 1972

western lyre snake UA 26954 1959

night snake MPM 18366 1997 R. M. Blaney

Sonoran coral snake NPS 584

black-tailed rattlesnake NPS 588

tiger rattlesnake UM 130211 1969 R. W. Van Devender

AU 1964 1959 G. Folkerts

UA 43288 1979

Arizona black rattlesnake NPS 578, 592, 599 1967 H. Coss

Mojave rattlesnake AU 1963 1959 G. Folkerts

Bird Cooper’s hawk UA 4008, 12936, 14888 1911 H. Brown

red-tailed hawk UA 4027, 14771 1911 H. Brown

zone-tailed hawk UA 4034 H. Brown

Gambel’s quail UA 14911

Montezuma quail NPS 4712

band-tailed pigeon UA 3534, 3377 1911 H. Brown

white-winged dove UA 9934

flammulated owl UA 14877

great-horned owl UA 6153

elf owl Yale 7473, 7474 1958 D. H. Parsons

UA 14146, 14761, 16540

western screech-owl UA 13925, 14891

common poorwill Yale 7475 to 7478 1958 D. H. Parsons

black-chinned hummingbird UA 14893

Anna’s hummingbird UA 3200 1911 H. Brown

rufous hummingbird UA 9944

broad-tailed hummingbird UA 3495 1911 H. Brown

magnificent hummingbird UA 3496 1911 H. Brown
143

Taxon Common name Collectiona Collection number

Collection 

date Collector

Bird northern flicker UA 1502, 14892 1911 H. Brown

acorn woodpecker UA 1841 to 1845 1911 H. Brown

Gila woodpecker UA 7482, 13857, 14889, 14894

red-naped sapsucker UA 14894

ladder-backed woodpecker UA 2823

Stellar’s jay UA 2773, 2774 1911 H. Brown

brown-crested flycatcher UA 14145

dusky flycatcher UA 11306

Pacific-slope flycatcher UA 2659, 14874 1911 H. Brown

buff-breasted flycatcher UA 1871 1911 H. Brown

greater pewee UA 1874 to 1879 1911 H. Brown

western wood-pewee UA 1880, 1881 1911 H. Brown

violet-green swallow UA 1889 1911 H. Brown

purple martin UA 14880

house wren UA 14876

cactus wren UA 14144

curve-billed thrasher Yale 6474 1932 A. Walker

UA 14155

bridled titmouse UA 2788 1911 H. Brown

bushtit UA 2798, 3878 1911 H. Brown

white-breasted nuthatch UA 1913, 10823, 10824 1911 H. Brown

pygmy nuthatch UA 1915 to 1925, 2921, 2922, 

14873 1911 H. Brown

brown creeper UA 3876, 14879 1911 H. Brown

olive warbler UA 1988 to 1991 1911 H. Brown

Virginia’s warbler UA 3146 1911 H. Brown

yellow-rumped warbler UA 2000 1911 H. Brown

hermit warbler UA 2006, 3196 to 3198 1911 H. Brown

Grace’s warbler UA 2010 to 2012, 3245 1911 H. Brown

MacGillivray’s warbler UA 10256

painted redstart 1831 1911 H. Brown

UA 3318, 3705 1960 R. R. Johnson

Bell’s vireo UA 3121 1986 H. Brown

solitary vireo UA 1981 1911 H. Brown

warbling vireo UA 14875

spotted towhee UA 1678, 2488 1911 H. Brown

canyon towhee UA 2178 1958 J. T. Marshall

yellow-eyed junco UA 1622 to 1626, 2304 to 2308 1911 H. Brown

Brewer’s sparrow UA 10194, 10195

black-throated sparrow UA 14351, 14890

white-crowned sparrow UA 14897

western bluebird UA 1954, 1955, 3037, 3038, 3040, 

3041 1911 H. Brown

western tanager UA 14895

UA 14896

hepatic tanager UA 1697, 1698 1911 H. Brown

Mammal desert shrew UA 26017 1988 R.M. Sidner

Mexican long-tongued Bat UA 7906, 7955 1960 B. Hayward

UA 26677, 3651-3660 1999 J. Walner

southern long-nosed bat UA 7748, 7749 1960 P. J. Gould, B. J. Hayward 

et. al.

UA 14491, 14495 1966 R. J. Baker

UA 16011, 16115-16117, 17013 1967 J. T. Mascarello

Yuma myotis UA 25518 1986 R. M. Sidner

cave myotis UA 7750-7754 1960 P. J. Gould, B. J. Hayward 

et.al. 

southwestern myotis SDMNH 10086 1932 T. W. Sefton, L. M. Huey

UA 25350 1985 R. Davis

UA 25519, 22521, 25525 1986 R. M. Sidner

fringed myotis UA 15333-15335, 15361 1966 B. A. Lund

UA 25349, 25352 1985 R. Davis 
144

Taxon Common name Collectiona Collection number

Collection 

date Collector

UA 25522, 25524 1986 R. M. Sidner 

long-legged myotis SDMNH 10084 1932 L. M. Huey, J. W. Sefton

UA 25351 1985 R. Davis 

UA 25515-22517 1986 R. M. Sidner

UA 25526 1985 R. Davis

California myotis UA 25520 1986 R. Sidner

western small-footed myotis UA 25523 1986 R. M. Sidner 

silver-haired bat UA 25514 1986 R. M. Sidner

SDMNH 10076 1932 L. M. Huey, J. W. Sefton 

SDMNH 10105 1932 T. W. Sefton, L. M. Huey

UA 810-811 1911 H. Brown 

Townsend’s big-eared bat UA 16113, 16114 1967 J. T. Mascarello

UA 16746-16748, 16875, 16876, 

16974 1967 G. Clay Mitchell

desert cottontail UIMNH 23360, 26220, 26221 1946 W. & L. Goodpaster 

NPS 61 1958

black-tailed jackrabbit UA 7091 1959 G. V. R. Bradshaw

UA 12000 1964 J. H. Nelson

antelope jackrabbit UA 12007 1964 G. L. Hathaway

UIMNH 26244 1946 W. W. Goodpaster

cliff chipmunk SDMNH 10067 1932 L. M. Huey

SDMNH 10071 1932 L. M. Huey, S.G. Harter

SDMNH 10089, 10090, 10118-10120 1932 L. M. Huey, L.H. Cook

SDMNH 10094-10102, 10124, 10125 1932 T. W. Sefton, L. M. Huey

SDMNH 10132, 10143, 10144 1932 L. M. Huey

UA 879 1911 H. Brown

UA 16464-16466 1966 L. Christianson

UA 25346 1984 R. Davis 

NPS 543, 544 1966 Mulhern

Harris’s antelope squirrel UA 2817 R. E. Dingman

UA 2944, 2945 1963 J. L. Patton

UIMNH 18325 1958 I. A. Nadr

UIMNH 23983, 23984 1946 W.W. Goodpaster

NPS 60, 287 1940, 1960

rock squirrel SDMNH 10087, 10129 1932 L. M. Huey, J. W.Sefton

SDMNH 10116 1932 S. G. Harter, L. M. Huey

SDMNH 10145 1932 L.M. Huey

UIMNH 24013 1946 W.W. Goodpaster

round-tailed ground squirrel UA 2809-2816 1963 R.E. Dingman

UIMNH 23976-23979 1946 W.W. Goodpaster

Arizona gray squirrel SDMNH 10079, 10086, 10092, 10130 1932 L.M. Huey, J.W.Sefton

Botta’s pocket gopher SDMNH

10062-10083, 10085, 10088, 

10106-10110, 10117, 10127, 

10128, 10133-10151

1932 L.M. Huey

UA 875 1911 H. Brown 

UIMNH 24281-24284 1946 W. & L. Goodpaster 

silky pocket mouse UA

3004, 7098, 7119, 7120, 3003, 

3005, 3006, 7096-7102, 7121, 

7122

1959 G. V. R. Bradshaw

UA 12222-12232 1963 D. Wright

UA 16751 1966 J. L. Patton

UIMNH 4812-4813 1946 W. L. Goodpaster

UIMNH 24374, 24375, 24382, 18373, 

18374 1946 W. W. Goodpaster 

Bailey’s pocket mouse UA 7095, 24604, 24605, 25347 1959 G. V. R. Bradshaw

UA 25725-25727 1984 D. Johnson 

UIMNH 18375-18378 1958 D. F. Hoffmeister

UIMNH 24411, 24412, 24425-24427 1946 W. W. Goodpaster

rock pocket mouse NPS 292, 293, 294 1963 Dengler

kangaroo rat UA 2093 1959 E. L. Cockrum

bannertail kangaroo rat UIMNH 24329-24331 1946 W. W. Goodpaster
145

Taxon Common name Collectiona Collection number

Collection 

date Collector

Merriam’s kangaroo rat UA 7094 1959 G. V. R. Bradshaw

UIMNH 18465-18469 1958 J. S. Hall

UIMNH 24342, 24343 1946 W. W. Goodpaster

cactus mouse UA 24882, 26006, 26008 1984 R. Davis 

UIMNH 18659, 24564, 24565 1958 I. A. Nadr

deer mouse UA 1388, 1389 1954 W. Collins 

brush mouse SDMNH 10103, 10104, 10112-10115, 

10121, 10122, 10126, 10131 1932 L. M. Huey 

UA 1387, 1390-1392 1954 W. Collins 

UA 26939-26943 1985 R. Davis 

UIMNH 24549 1932 A. Walker 

southern grasshopper mouse UA 7092 1959 G. Bradshaw

UIMNH 18672, 25566-25568 1958 J. S. Hall

NPS 297 1983

Arizona cotton rat UA 25191 1983 R. Davis 

yellow-nosed cotton rat UA 25192-25194, 26011-26016 1984 R. Davis 

western white-throated woodrat UA 7093 1959 G.V.R. Bradshaw

UA 25655 1974 R. Dickson 

UIMNH 18700, 18709, 18710, 25844-

25850 1958 J. S. Hall

Mexican woodrat SDMNH 10111, 10123 1932 L.M. Huey 

UA 16461-16463 1966 L. Christianson

UA 26938 1985 R. Davis 

NPS 542 1966 Mulhern

common gray fox UA 25992 R. Davis 

NPS 306, 307, 1283 1962

North American porcupine SNP 536 1963 B. Lund 

American badger UA 17900 1968 Wade

jaguar UA 588,589 1932 H. Wilson 

mule deer SDMNH 10091 1932 L.M. Huey, L.H.Cook

collared peccary NPS 303, 305

a

 AU = Auburn University Museum; FWMSH = Fort Worth Museum of Science and History; MPM = Milwaukee Public Museum; NHMLAC = 

Natural History Museum, Los Angeles County; NPS = National Park Service Western Archaeologial Conservation Center; OMNH = Oklahoma 

Museum of Natural History; SDMNH = San Diego Museum of Natural History; SNP = Saguaro National Park; UA = University of Arizona 

Collections; UIMNH = University of Ilinois (Champaign-Urbana) Museum of Natural History; UM = University of Michigan; USMN = U.S. 

National Museum; Yale = Yale University, Peabody Museum of Natural History. 

b Observed outside but near the park. 
146

Riparian Sonoran Desertscrub

Species

Total

transects 

observed 

Lower 

Rincon 

Creek

Upper 

Rincon 

Creek

Box

Canyon

Loma

Verde

Wash 112 115 121 130 138 139

mallard 1 0.05

wild turkey 1

scaled quail 1

Gambel’s quail 13 2.03 0.51 1.22 1.09 0.69 1.81 0.63 0.25 0.44 0.13

Montezuma quail 6 0.06

turkey vulture 12 0.20 0.16 0.33 0.61 0.06 0.19 0.06 0.06

sharp-shinned hawk 1 0.01

Cooper’s hawk 4 0.14 0.12 0.02 0.04

northern goshawk 2

gray hawk 1 0.05

zone-tailed hawk 3 0.02 0.02

red-tailed hawk 7 0.14 0.06

golden eagle 3 0.13

American kestrel 7 0.03 0.08 0.02 0.06 0.06 0.06

peregrine falcon 3

rock pigeon 1 0.05

band-tailed pigeon 5 0.06

white-winged dove 18 1.62 1.10 2.49 2.87 1.63 2.19 1.63 2.25 2.31 1.19

mourning dove 16 1.47 1.08 1.73 3.57 1.19 2.25 1.56 2.00 1.69 2.06

common ground-dove 2 0.05 0.14

yellow-billed cuckoo 1 0.06

greater roadrunner 4 0.02 0.09

whiskered screech-owl 1

great horned owl 6 0.10 0.17 0.06 0.06

northern pygmy-owl 2

lesser nighthawk 1 0.02

common poorwill 2

whip-poor-will 2

white-throated swift 14 0.01 0.14 0.06 0.13 0.19 0.25 0.50 0.06

broad-billed hummingbird 3 0.09 0.18 0.24

magnificent hummingbird 2

black-chinned hummingbird 13 0.20 0.37 0.16 0.09 0.13 0.06 0.06 0.19

Anna’s hummingbird 8 0.13 0.06

Costa’s hummingbird 7 0.01 0.08 0.13 0.13 0.06

broad-tailed hummingbird 10 0.01 0.04 0.03

rufous hummingbird 2 0.02 0.06

elegant trogon 1

belted kingfisher 1 0.01

acorn woodpecker 9

gila woodpecker 12 1.30 0.37 1.56 1.74 1.69 2.31 0.31 1.13 0.75 2.13

ladder-backed woodpecker 14 0.44 0.24 0.33 0.48 0.25 0.19 0.50 0.19 0.13 0.38

hairy woodpecker 4

Arizona woodpecker 5

northern flicker 10 0.02 0.02

gilded flicker 10 0.06 0.04 0.19 0.13 0.19 0.44 0.44 0.19 0.06 0.81

northern beardless-tyrannulet 4 0.02 0.12 0.19 0.06

greater pewee 5

western wood-pewee 11 0.03 0.02 0.06

Hammond’s flycatcher 2 0.13

gray flycatcher 3 0.05 0.06 0.13

dusky flycatcher 1 0.06

Appendix G. Mean frequency of detection of birds, by community type and transect, recorded during repeatvisit VCP surveys, Saguaro National Park, Rincon Mountain District, 2001 and 2002. “Total transects” indicates 

in how many transects (all commununities; maximum value is 23) we recorded at least one individual during surveys. 

Frequency of detections includes all birds recorded including flyovers, birds seen >75 m from stations, and birds 

recorded outside of 8-minute-count periods. 
147

Riparian Sonoran Desertscrub

Species

Total

transects 

observed 

Lower 

Rincon 

Creek

Upper 

Rincon 

Creek

Box

Canyon

Loma

Verde

Wash 112 115 121 130 138 139

pacific-slope flycatcher 1 0.02

cordilleran flycatcher 7 0.01

black phoebe 5 0.02 0.14 0.02

vermilion flycatcher 2 0.28 0.02

dusky-capped flycatcher 9 0.13

ash-throated flycatcher 21 0.64 0.55 0.49 0.65 1.63 1.56 1.06 1.44 0.63 1.81

brown-crested flycatcher 13 1.24 0.49 0.87 0.39 0.50 0.06 0.56 0.63 0.75 0.63

sulphur-bellied flycatcher 1

Cassin’s kingbird 6 0.31 0.02 0.25 0.19

western kingbird 4 0.10 0.13 0.06 0.13

Bell’s vireo 6 0.51 1.35 0.94 0.09 0.13 0.19

plumbeous vireo 7

Hutton’s vireo 11 0.06

warbling vireo 5 0.02 0.02 0.04 0.06

Steller’s jay 4

western scrub-jay 10 0.08 0.13 0.19 0.06

Mexican jay 10

common raven 19 0.13 0.06 0.06 0.26 0.13 0.06 0.06 0.19 0.13

purple martin 4 1.02 0.49 0.35 0.06

violet-green swallow 11 0.05 0.06 0.03 0.06

mountain chickadee 3

bridled titmouse 10 0.07

verdin 11 0.87 0.76 1.62 1.30 0.88 1.00 0.06 0.50 0.69 0.69

bushtit 9

red-breasted nuthatch 1

white-breasted nuthatch 8

pygmy nuthatch 3

brown creeper 3

cactus wren 13 0.68 0.75 1.14 0.74 2.38 1.19 1.06 1.44 1.00 1.31

rock wren 13 0.02 0.13 0.19 0.63 0.19

canyon wren 17 0.16 0.56 0.31 0.56 0.31 0.38

Bewick’s wren 17 1.01 0.59 0.46 0.57 0.19 0.06

house wren 7 0.01 0.02

ruby-crowned kinglet 3 0.01 0.02

blue-gray gnatcatcher 14 0.03 0.02 0.02

black-tailed gnatcatcher 9 0.03 0.10 0.33 0.17 0.38 0.56 0.06 0.31 0.56

western bluebird 8 0.06

hermit thrush 5

American robin 5

northern mockingbird 13 0.02 0.04 0.05 0.19 0.63 0.25 0.13 0.13

curve-billed thrasher 11 0.43 0.33 0.92 0.87 1.00 1.00 0.19 0.06 0.44 0.44

Crissal thrasher 2 0.08 0.09

European starling 1 0.03

cedar waxwing 1 0.05

phainopepla 10 0.22 0.16 0.11 0.22 0.19 0.13 0.31 0.13

olive warbler 5 0.02

orange-crowned warbler 6 0.01 0.06 0.06 0.06

Nashville warbler 1 0.04

Virginia’s warbler 9 0.06

Lucy’s warbler 5 1.69 0.84 1.00 1.61 0.56

yellow warbler 3 0.22 0.12 0.03

yellow-rumped warbler 8 0.02 0.04

black-throated gray warbler 13 0.05 0.06 0.06

Townsend’s warbler 3 0.13

Grace’s warbler 6

Macgillivray’s warbler 2 0.01 0.03

Wilson’s warbler 10 0.10 0.12 0.05 0.17 0.13 0.06

red-faced warbler 4
148

Riparian Sonoran Desertscrub

Species

Total

transects 

observed 

Lower 

Rincon 

Creek

Upper 

Rincon 

Creek

Box

Canyon

Loma

Verde

Wash 112 115 121 130 138 139

painted redstart 3

hepatic tanager 9

summer tanager 3 0.30 0.27 0.02

western tanager 12 0.05 0.02 0.04 0.06

green-tailed towhee 8 0.02 0.02 0.30 0.22 0.13 0.06 0.06

spotted towhee 12 0.02

canyon towhee 13 0.22 0.37 0.44 0.52 0.81 0.44 1.13 0.63 1.00 0.50

Abert’s towhee 5 0.23 0.22 0.25 0.30 0.06

rufous-winged sparrow 8 0.36 0.16 0.21 0.26 0.06 0.06 0.06 0.13

rufous-crowned sparrow 19 0.01 0.02 0.03 0.31 0.31 1.19 1.44 0.50 0.81

chipping sparrow 8 0.03 0.20 0.08 0.26 0.06 0.06 0.06

Brewer’s sparrow 5 0.02 0.20 0.08 0.43 0.25

black-chinned sparrow 8 0.06 0.06

Lincoln’s sparrow 1 0.02

vesper sparrow 1 0.04

lark sparrow 5 0.09 0.04 0.02 0.04 0.06

black-throated sparrow 11 0.20 0.45 0.25 0.26 1.13 1.31 0.38 1.19 1.31 2.50

song sparrow 1 0.02

white-crowned sparrow 5 0.09 0.06 0.19 0.13

yellow-eyed junco 5

northern cardinal 10 0.79 0.41 1.03 0.39 0.19 0.44 0.38 1.19 0.06

pyrrhuloxia 7 0.10 0.08 0.26 0.06 0.13 0.13 0.13

black-headed grosbeak 16 0.02 0.06 0.08 0.04 0.06

blue grosbeak 5 0.25 0.33 0.02 0.06 0.06

lazuli bunting 4 0.03 0.16 0.02 0.06

indigo bunting 1 0.01

varied bunting 5 0.02 0.18 0.33 0.04 0.13

bronzed cowbird 1 0.06

brown-headed cowbird 18 0.41 0.33 0.78 0.22 0.44 0.13 0.38 0.56 0.31

hooded oriole 4 0.03 0.03 0.13 0.06

Bullock’s oriole 5 0.05 0.02 0.02 0.19 0.25

Scott’s oriole 17 0.06 0.02 0.04 0.38 0.13 0.75 0.25 0.44 0.31

house finch 14 0.53 0.20 2.24 0.52 2.00 1.94 1.38 0.75 0.19 0.94

pine siskin 2

lesser goldfinch 14 0.67 0.45 0.54 0.17 0.19 0.13 0.50 0.06

Appendix G continued. 

Oak Savannah Pine-oak Woodland Conifer Forest

Species 101 106 189 111 107 125 155 120 128

Happy Valley

Saddle 113 191

Rincon 

Peak

wild turkey 0.03

scaled quail 0.06

Gambel’s quail 2.81 0.94 0.06

Montezuma quail 0.19 0.13 0.06 0.19 0.06

turkey vulture 0.25 0.06 0.13 0.06

northern goshawk 0.11 0.04

zone-tailed hawk 0.31

red-tailed hawk 0.06 0.13 0.06 0.13 0.06

golden eagle 0.06 0.06

American kestrel 0.13

peregrine falcon 0.06 0.06 0.06

band-tailed pigeon 0.13 0.06 0.06 0.08

white-winged dove 3.19 2.13 2.94 0.69 1.38 0.06 0.03 0.04

mourning dove 1.94 1.00 0.44 0.19 0.31 0.31

greater roadrunner 0.13 0.06

whiskered screech-owl 0.06

great horned owl 0.13 0.06
149

Oak Savannah Pine-oak Woodland Conifer Forest

Species 101 106 189 111 107 125 155 120 128

Happy Valley

Saddle 113 191

Rincon 

Peak

northern pygmy-owl 0.06 0.06

common poorwill 0.31 0.03

whip-poor-will 0.13 0.06

white-throated swift 0.19 0.19 0.06 0.06 0.06 0.08

magnificent hummingbird 0.13 0.08

black-chinned hummingbird 0.13 0.13 0.06 0.06 0.13

Anna’s hummingbird 0.19 0.06 0.19 0.13 0.06 0.17

Costa’s hummingbird 0.06 0.06

broad-tailed hummingbird 0.13 0.25 1.06 0.08 0.50 0.06 0.08

rufous hummingbird

elegant trogon 0.11

acorn woodpecker 0.06 0.19 0.13 0.13 0.19 0.19 1.11 0.06 0.04

gila woodpecker 0.13 0.13

ladder-backed woodpecker 0.44 0.31 0.56 0.06

hairy woodpecker 0.11 0.25 0.13 0.38

Arizona woodpecker 0.19 0.13 0.13 0.06 0.11

northern flicker 0.19 0.06 0.13 0.06 0.11 0.38 0.88 0.46

gilded flicker

greater pewee 0.06 0.31 0.56 0.19 0.17

western wood-pewee 0.38 0.13 0.69 0.38 0.69 1.08 0.25 0.38

Hammond’s flycatcher 0.06

gray flycatcher

cordilleran flycatcher 0.06 0.06 0.11 0.94 0.44 0.50

black phoebe 0.06 0.13

dusky-capped flycatcher 0.13 0.06 0.50 0.25 0.38 0.25 0.64 0.17

ash-throated flycatcher 1.75 0.50 1.25 1.44 0.94 1.69 0.94 1.56 0.56 0.39 0.08

brown-crested flycatcher 0.13 0.19 0.06

sulphur-bellied flycatcher 0.22

Cassin’s kingbird 0.19 0.50

western kingbird

plumbeous vireo 0.06 0.06 0.25 0.58 0.44 0.38 0.46

Hutton’s vireo 0.06 0.06 0.13 0.31 0.63 0.81 0.19 0.13 0.28 0.17

warbling vireo 0.13

Steller’s jay 0.13 0.88 1.06 0.42

western scrub-jay 0.25 0.31 0.25 0.06 0.13 0.06

Mexican jay 0.31 1.44 0.75 0.06 2.31 1.25 1.50 0.69 1.06 1.11

common raven 0.06 0.13 0.19 0.06 0.06 0.44 0.31 0.06 0.31 0.33

violet-green swallow 0.25 0.25 0.19 0.42 0.06 1.13 0.54

mountain chickadee 0.81 0.44 1.13

bridled titmouse 0.19 0.63 0.13 0.19 0.06 0.25 0.38 0.19 0.31

verdin 0.19

bushtit 0.06 1.13 0.31 0.56 0.06 0.56 1.88 0.39 0.33

red-breasted nuthatch 0.08

white-breasted nuthatch 0.06 0.31 0.06 0.25 0.56 0.75 0.13 0.75

pygmy nuthatch 0.06 0.44 0.13

brown creeper 0.13 0.19 0.29

cactus wren 0.75 0.25 0.94

rock wren 0.06 0.56 0.31 0.38 0.13 0.25 0.25 0.25

canyon wren 0.19 0.88 0.69 0.94 0.38 0.69 0.31 0.63 0.17 0.25 0.04

Bewick’s wren 1.44 1.31 1.38 1.19 1.75 1.69 1.94 2.06 1.56 1.50 0.13

house wren 0.06 0.06 0.81 0.56 0.08

ruby-crowned kinglet 0.04

blue-gray gnatcatcher 0.25 0.31 0.31 0.13 0.25 0.31 0.06 0.69 0.94 0.17 0.08

black-tailed gnatcatcher

western bluebird 0.06 0.13 0.06 0.06 0.38 0.56 0.04

hermit thrush 0.13 0.19 0.63 0.56 1.21

American robin 0.06 0.03 1.06 1.00 0.17

northern mockingbird 2.63 0.19 1.44 0.13 0.13

curve-billed thrasher 0.06
150

Oak Savannah Pine-oak Woodland Conifer Forest

Species 101 106 189 111 107 125 155 120 128

Happy Valley

Saddle 113 191

Rincon 

Peak

phainopepla 1.19 0.63

olive warbler 0.03 0.19 0.06 0.46

orange-crowned warbler 0.08 0.04

Virginia’s warbler 0.06 0.31 0.63 0.13 0.19 0.25 0.03 0.21

yellow-rumped warbler 0.19 0.25 0.03 0.31 0.63 0.33

black-throated gray warbler 0.94 0.31 0.06 1.00 0.75 1.13 0.94 1.00 0.47 0.54

Townsend’s warbler 0.31 0.25

Grace’s warbler 0.06 0.25 1.00 0.81 0.19 0.63

Wilson’s warbler 0.13 0.06 0.06 0.04

red-faced warbler 0.13 0.06 1.13 0.50

painted redstart 0.13 0.06 0.33

hepatic tanager 0.13 0.13 0.06 0.31 0.06 0.25 0.88 0.50 0.33

western tanager 0.25 0.13 0.13 0.13 0.42 0.81 0.81 1.04

green-tailed towhee 0.13

spotted towhee 0.06 1.31 0.06 1.88 2.13 1.63 2.13 1.63 0.83 1.13 0.42

canyon towhee 0.50 0.38 0.31

rufous-winged sparrow

rufous-crowned sparrow 1.38 1.94 0.69 2.00 0.81 0.56 0.25 0.69 0.44 0.08

chipping sparrow 0.06

black-chinned sparrow 0.06 1.31 0.06 0.81 0.88 0.94

black-throated sparrow 0.31

white-crowned sparrow 0.06

yellow-eyed junco 0.13 0.11 1.69 1.94 1.38

northern cardinal 0.06

pyrrhuloxia

black-headed grosbeak 0.38 1.13 0.06 0.19 0.25 0.69 0.06 1.44 0.50 0.50 0.46

brown-headed cowbird 0.63 0.25 0.38 0.06 0.38 0.31 0.31 0.06 0.22

Bullock’s oriole

Scott’s oriole 0.50 0.31 0.75 1.06 0.13 0.31 0.13 0.11

house finch 0.69 0.31 0.13 1.25

pine siskin 0.94 0.04

lesser goldfinch 0.13 0.13 0.06 0.06 0.19 0.14
151

Appendix H. Mean density (number of stems/hectare) of large trees and potential cavity-bearing plants at non-random, 

repeat-visit VCP stations, Saguaro National Park, Rincon Mountain District, 2001 and 2002. See Appendix A for common 

names.

Acacia Celtis

Transect

StationconstrictagreggiiCarnegia

gigantea

pallidareticulataParkinsonia

microphyllum

Fouquieria

splendens

Fraxinus

velutina

Platanus

wrightii

Populus

fremontii

Prosopis

velutina

Salix

gooddingii

Lower Rincon 

Creek 1 80.5 16.6 11.9 7.5 85.2 80.5

2 26.5 12.7 9.5 15.2 25.8

3 7.6 2.4 4.9 2.8 14.5 2.4

4 37.3 29.4 23.5 117.5

5 24.3 8.1 1.3 8.1 9.2 0.6 53.6

6 15.4 15.4 1.6 0.8 4.3 0.3 58.3

7 0.9 7.8 6.4 0.3 0.4 50.5 3.9

8 22.0 7.3 1.1 15.6 2.3 1.1 44.4

Upper Rincon 

Creek 1 9.8 29.5 6.7 42.3 10.5 8.3 20.8

2 16.1 29.8 32.3 16.1 38.5 2.5 67.1

3 18.7 22.1 28.0 24.3 38.2

4 16.0 0.9 26.7 8.3 1.2 0.2 48.0

5 17.3 3.9 34.6 8.8 1.1 0.7 28.8 0.4

6 4.6 1.5 18.6 9.4 0.8 23.2

7 7.1 9.8 6.6 5.5 1.6 22.9 6.6

Box Canyon 1 4.9 18.3 14.6 14.6 4.9 5.4

2 59.2 11.3 23.7 23.7 23.7 41.7

3 17.4 6.3 17.4 17.4 17.4 8.7 31.1

4 60.1 12.5 20.0 10.0 30.1 12.7 22.1

5 13.3 1.0 7.3 7.3 12.0 29.9

6 9.8 10.9 15.1 9.8 29.4 0.8 1.6 49.4

7 49.3 15.2 12.3 6.1 77.0

Upper Loma 

Verde Wash 1 20.0 150.3 1.4 15.8

2 6.5 45.5 6.5 27.6

3 37.0 27.8 5.9 9.2 32.6

4 78.0 9.5 11.1 33.4 89.2

5 12.1 10.2 6.1 48.8

Transect Station

Arctostaphylos

sp. a

Juniperus

deppeana

Pinus

cembroides

Pinus

leiophylla

Pinus

ponderosa

Pseudotsuga

menziesii

Quercus 

sp.b

Quercus

gambelii

Rhamnus

crocea

Happy Valley 

Saddle 1 342.3 185.9 720.6 294.2

2 93.4 100.2 53.5 137.5 157.1 46.7

3 76.6 197.5 31.1 15.5 40.6

4 66.6 42.7 5.0 123.0 351.9 85.5

5 50.4 145.7 1.2 105.1 269

6 24.3 272.4 8.5 24.3 129

Rincon Peak 1 5.1 34.5 107.3 76.6 344.6

2 21.9 34.9 4.5 131.3 150.6 139.5

3 15.5 19.4 162.8 42.6

4 216.3 61.9

a A. pringlei, and A. pungens. b Q. arizonica, Q. emoryi, Q. hypoleucoides, and Q. rugosa.
152

Appendix I. Details of small-mammal trapping effort, Saguaro National Park, Rincon Mountain District, 

2001 and 2002. 

Elevation 

stratum Plot group Plot type Visit

Number of 

traps set

Sprung but 

empty traps

Number 

of animals 

captured

Number 

of animals 

recaptured

Number of 

trap nights

Low 112 Random 1 126 0 5 1 123.0

2 225 12 43 17 189.0

139 Non-random 1 30 0 0 0 30.0

Random 1 189 0 1 0 188.5

2 225 4 6 0 220.0

Lower Rincon Creek Non-random 1 225 65 66 20 149.0

Upper Rincon Creek Non-random 1 120 15 15 3 104.5

Middle 101 Non-random 1 60 0 4 0 58.0

Random 1 189 0 19 3 178.0

2 225 1 27 10 206.0

111 Non-random 1 75 0 7 1 71.0

Random 1 105 0 2 0 104.0

121 Random 1 150 0 20 1 139.5

2 150 7 41 20 116.0

189 Random 1 225 0 10 5 217.5

2 150 5 20 10 132.5

Douglas Springs Non-random 1 400 36 49 44 335.5

Grass Shack Non-random 1 146 1 21 7 131.5

Juniper Basin Non-random 1 535 33 18 8 505.5

High 113 Random 1 189 1 15 7 177.5

2 225 3 48 10 194.5

128 Random 1 126 3 14 3 116.0

2 200 4 9 3 192.0

191 Random 1 189 1 0 0 188.5

Italian Spring Non-random 1 300 3 57 23 258.5

Manning Camp Non-random 1 15 0 5 1 15.0

Spud Rock Non-random 1 135 3 14 0 112.0

Spud Rock Spring Non-random 1 160 1 36 12 135.5

Appendix J. Summary of field effort for bats, Saguaro National Park, Rincon Mountain 

District, 2001 and 2002. See text for explanation of net hours calculations. 

Type of 

investigation Stratum Location Year Month/day

Total time 

(hours)

total net 

length (m)

Net 

hours

Roost NA Box Canyon Crevice 2001 05/13 NA

Tanque Verde Ridge 2002 05/23 NA

Helen’s Dome 2001 05/22 NA

Netting Low Chimenea Creek 2001 05/14 6.0 18 108.0

Lower Rincon Creek 2001 05/17 8.7 21 182.0

09/16 4.3 5 21.7

09/28 2.9 5 14.6

2002 08/13 6.0 37 222.0

09/30 4.3 19 82.3

09/18 4.7 15 70.0

Middle Deer Creek 2002 05/04 9.5 10 95.0

Wild Horse Canyon 2001 04/30 8.4 10 84.2

05/01 8.3 10 82.5

05/02 8.5 10 85.0

High Devil’s Bathtub 2001 09/23 3.0 9 27.0

Manning Camp Pond 2001 05/19 1.8 9 15.8

05/20 6.5 18 117.0

05/21 6.0 18 108.0

09/22 3.1 9 27.8

09/24 2.8 9 24.8
153

Random or 

Non-random

Camera 

number

Number of

camera nights

Number of 

photographs

Number of 

individuals

Number of

species

Elevation 

(m)

Non-random 1 99 91 139 7 940

2 38 28 32 5 919

3 12 6 7 1 920

4 5 16 16 3 846

5 12 16 17 1 860

6 6 7 7 1 849

7 12 12 12 1 866

8 21 27 28 3 943

9 12 20 22 6 844

10 20 9 9 5 968

11 15 23 23 6 863

12 40 27 27 5 935

13 15 31 34 3 957

14 52 68 89 3 982

15 16 12 12 4 948

16 10 10 10 4 980

17 46 33 38 4 997

18 16 2 2 1 954

19 14 18 18 3 1049

20 27 18 24 4 1000

21 113 58 78 8 999

22 51 32 34 5 987

23 19 56 59 4 974

24 21 8 8 4 967

25 5 8 9 2 1082

26 34 47 48 9 973

27 37 45 54 6 960

28 52 23 23 5 954

29 12 4 5 2 967

30 28 37 50 4 965

31 12 16 20 3 958

32 121 79 94 8 958

33 37 21 22 2 960

34 7 7 7 2 973

35 14 17 17 6 1384

36 35 41 41 7 1364

37 9 8 13 2 1143

38 27 25 32 4 1112

39 88 62 78 7 1025

40 47 23 27 9 1029

41 9 10 11 5 1018

42 25 10 10 6 1043

43 30 6 6 4 1034

44 25 19 19 6 1021

45 162 61 71 11 1057

46 211 215 265 10 1061

47 16 8 8 3 1067

48 61 59 99 8 1079

49 12 7 7 2 1023

50 12 12 14 3 1030

51 27 10 12 2 1019

52 15 6 7 3 1868

53 44 6 6 1 1791

54 95 13 13 3 1605

55 12 17 17 7 1603

56 13 4 5 2 2173

56 13 7 7 3 1109

Appendix K. Details of infrared-triggered camera effort and results, Saguaro National Park, Rincon 

Mountain District, 1999-2005. Survey effort summarized in Table 6.2. 
154

Random or 

Non-random

Camera 

number

Number of

camera nights

Number of 

photographs

Number of 

individuals

Number of

species

Elevation 

(m)

Non-random 58 43 51 62 8 1028

59 125 26 31 7 1082

60 4 2 2 2 2325

61 12 20 21 7 1039

62 55 16 16 3 2384

63 37 22 28 3 2421

64 32 7 8 3 2573

65 10 3 3 2 2591

66 52 25 25 8 2199

67 49 19 20 9 1862

68 12 2 2 1 1867

69 15 13 13 6 1869

70 13 18 18 3 963

71 17 24 25 3 975

72 40 14 14 6 958

73 59 42 42 9 968

74 42 13 13 3 960

Random 101C 11 6 6 3 1349

101D 11 2 2 2 1372

102C 12 11 15 3 1517

102D 4 2 2 1 1656

102R 12 27 27 2 2085

103C 25 23 23 3 1606

103D 2 12 12 3 1609

103R 9 12 12 3 958

106C 6 11 11 2 1739

106D 14 9 9 1 1592

106R 21 11 11 4 1654

109C 16 17 17 2 1197

109D 2 5 5 1 1164

109R 12 5 5 2 1302

10C 9 8 8 3 1707

10D 7 2 2 2 1778

10R 8 8 8 1 1821

110C 24 10 10 4 1076

110D 24 16 16 3 1085

110R 24 2 2 2 1069

111C 25 16 16 3 1266

111D 25 2 2 1 1249

112C 21 33 33 5 1022

112D 16 3 3 2 1021

112R 16 18 18 4 1002

115C 5 2 2 0 898

115D 16 9 9 3 890

115R 15 7 7 2 930

118C 16 42 47 4 947

118D 17 4 9 3 938

118R 17 23 45 4 966

119C 12 8 24 4 866

119D 11 4 4 2 865

119R 18 10 10 4 869

120D 13 17 17 1 1609

120R 9 2 2 1 1706

121C 1 1 1 1 1442

121D 19 15 17 3 1351

121R 2 5 6 3 1414

122C 17 18 19 1 1215

122R 3 2 2 1 1265

126C 18 8 18 4 1415

126D 41 43 50 5 1283

Random 126R 6 6 21 3 1386
155

Random or 

Non-random

Camera 

number

Number of

camera nights

Number of 

photographs

Number of 

individuals

Number of

species

Elevation 

(m)

Random 130C 2 38 70 3 1071

130D 14 13 13 3 1095

130R 14 10 10 1 1107

134C 7 16 16 4 1289

134D 16 10 10 4 1739

138C 8 6 6 1 1021

138D 12 4 4 2 1024

138R 12 4 5 1 1048

139C 16 24 24 7 974

139R 15 8 8 3 1008

143C 15 20 21 2 1019

143D 19 15 15 2 873

143R 15 22 22 3 970

144C 10 6 6 3 829

144D 9 5 5 2 827

144R 10 5 5 2 828

148C 18 6 6 1 1536

148D 12 6 6 2 1585

148R 24 10 10 2 1615

15C 14 17 17 3 1553

15D 1 1 1 1 1561

15R 10 10 10 2 1564

16C 3 4 4 1 1502

16D 13 6 6 2 1463

16R 5 9 9 3 1507

18C 14 7 7 3 1380

18D 19 31 31 3 1387

195C 3 14 14 4 981

195D 21 27 31 2 985

195R 24 28 29 4 1005

1C 1 2 2 1 1772

1D 17 23 23 2 1739

1R 15 7 9 1 1914

2C 15 9 12 2 1869

2D 10 18 23 2 1804

2R 17 13 15 2 1838

3C 7 3 3 2 1480

3D 11 9 10 2 1512

3R 3 5 5 1 1519

9R 14 9 9 5 1777

Q-9D 1 1 1 1 2609

Q-12R 7 6 6 4 2417

Q-16R 1 1 1 1 2426

Q-16D 12 12 12 2 2341

Q-18R 12 5 5 1 2279

Q-18D 3 2 2 1 2237

Q-19R 11 3 3 1 2090

Q-19D 4 5 5 2 2154

Q-1R 5 5 5 1 2213

Q-1D 15 10 10 2 2195

Q-20R 12 5 5 2 2348

Q-20D 1 1 1 1 2300

Q-20C 20 7 7 2 2295

Q-25R 4 3 4 1 2160

Q-25D 4 3 3 1 2152

Q-7R 12 3 3 3 2367

Q-9R 18 12 12 4 2568

	 	 	 	 	 	 	 	
156