Providing the knowledge and technology 

required to improve the availability of native 

plantmaterialsfor restoring diverse native

plant communities across the GreatBasin.

GREAT BASIN NATIVE PLANT PROJECT

2014 PROGRESSREPORT

USDA FOREST SERVICE, ROCKY MOUNTAIN RESEARCH STATION

AND USDI BUREAU OF LAND MANAGEMENT, BOISE, ID

APRIL 2015

COOPERATORS

USDA Forest Service, Rocky Mountain Research Station

Grassland, Shrubland and Desert Ecosystem Research Program, Boise, ID, Provo, UT, and Albuquerque, NM

USDI Bureau of Land Management, Plant Conservation Program, Washington,DC

Boise State University, Boise, ID

Brigham Young University, Provo, UT

College of Western Idaho, Nampa, ID

Eastern Oregon Stewardship Services, Prineville, OR

Oregon State University Malheur Experiment Station, Ontario,OR

Private Seed Industry

Texas Tech University, Lubbock, TX

Truax Company, Inc., New Hope, MN

University of Idaho, Moscow, ID

University of Idaho Parma Research and Extension Center, Parma,ID

University of Nevada, Reno, NV

University of Nevada Cooperative Extension, Elko and Reno, NV

Utah State University, Logan, UT

USDA Agricultural Research Service, Bee Biology and Systematics Laboratory, Logan, UT

USDA Agricultural Research Service, Eastern Oregon Agriculture Research Center, Burns,OR

USDA Agricultural Research Service, Forage and Range Research Laboratory, Logan,UT

USDA Agricultural Research Service, Great Basin Rangelands Research Unit, Reno,NV

USDA Agricultural Research Service, Western Regional Plant Introductions Station, Pullman,WA

USDA Forest Service, National Seed Laboratory, Dry Branch, GA

USDA Forest Service, Pacific Northwest Research Station, Corvallis, OR

USDA Natural Resources Conservation Service, Aberdeen Plant Materials Center, Aberdeen,ID

USDI Bureau of Land Management, Morley Nelson Birds of Prey National Conservation Area, Boise,ID

US Geological Survey Forest and Rangeland Ecosystem Science Center, Boise,ID

Utah Division of Wildlife Resources, Great Basin Research Center, Ephraim,UT

Utah Crop Improvement Association, Logan, UT
i

Great Basin Native Plant Project 

2014 Progress Report

The Interagency Native Plant Materials Development Program outlined in the 2002 United States 

Department of Agriculture (USDA) and United States Department of Interior (USDI) Report to 

Congress encouraged use of native plant materials for rangeland rehabilitation and restoration 

where feasible. The Great Basin Native Plant Project is a cooperative project lead by the Bureau 

of Land Management (BLM) Plant Conservation Program and the United States Forest Service 

(USFS), Rocky Mountain Research Station that was initiated to provide information that will be 

useful to managers when making decisions about the selection of genetically appropriate 

materials and technologies for vegetation restoration. The Project is supported by the USDI 

Bureau of Land Management, Plant Conservation Program and administered by the USFS Rocky 

Mountain Research Station’s Grassland, Shrubland and Desert Ecosystem Research Program.

Research priorities are to:

 Increase the variety of native plant materials available for restoration in the Great Basin.

 Provide an understanding of species variability and potential response to climate change 

to improve seed transfer guidelines.

 Develop seeding technology and equipment for successful reestablishment of native plant 

communities.

 Transfer research results to land managers, private sector seed growers, and restoration 

contractors.

We thank our many collaborators for their dedication and their institutions for their in-kind 

contributions. The wide array of expertise represented by this group has made it possible to 

address the many challenges involved with this endeavor. We especially thank Alexis Malcomb 

for the many hours she spent carefully editing and compiling this report and other outreach 

materials and Nancy Shaw and Jeff Ott for help in editing, as well as Corey Gucker for help in 

managing our financial reporting. Special thanks also to our resident students and volunteers: 

Matt Fisk, Jameson Rigg, and Kristof Bihari for managing our field and laboratory research.

Francis Kilkenny 

USDA Forest Service

Rocky Mountain Research Station 

Boise, ID

ffkilkenny@fs.fed.us

Great Basin Native Plant Project

http://www.fs.fed.us/rm/boise/research/shrub/greatbasin.shtml

Citation

Kilkenny, Francis; Halford, Anne; Malcomb, Alexis. 2015. Great Basin Native Plant Project: 

2014 Progress Report. Boise, ID: U.S. Department of Agriculture, Rocky Mountain Research 

Station. 210 p.
ii

Results in this report should be considered preliminary in nature and should not be quoted or 

cited without the written consent of the Principal Investigator for the study.

The use of trade or firm names in this report is for reader information only and does not imply 

endorsement by the USDA or USDI of any product or service.

Pesticide Precautionary Statement: This publication reports some research involving pesticide 

use. It does not contain recommendations for their use, nor does it imply that the uses discussed 

here have been registered. All uses of pesticides must be registered by appropriate State and/or 

Federal agencies before they can be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals, desirable plants, and fish 

or other wildlife― if they are not handled or applied appropriately. Use all pesticidesselectively 

and carefully. Follow recommended practices for the disposal of surplus pesticides and pesticide 

containers.

The USDA and USDI are equal opportunity providers and employers.
iii

HIGHLIGHTS FOR 2014

GENETICS AND SEED ZONES

Genetic Diversity and Genecology of Prairie Junegrass (Koeleria macrantha) and 

Squirreltail (Elymus elymoides)

Francis Kilkenny and Brad St. Clair

 Common gardens were established in contrasting sites for prairie junegrass (Koeleria 

macrantha) and squirreltail (Elymus elymoides).

 We evaluated and compared growth, morphology, and reproductive traits for populations 

of each both species.

 General patterns that emerged from the preliminary analysis of prairie junegrass common 

gardens were largely congruent with large-scale ecoregions, suggesting that Omernick’s 

Level III ecoregions may be useful for seed zones. A preliminary seed zone map was 

developed.

 Analyses from squirretail common gardens suggest that squirreltail traits are highly 

plastic with phenological traits varying primarily by year, and size traits varying 

primarily by planting site.

Testing the Efficacy of Seed Zones for Re-establishment and Adaptationof 

Bluebunch Wheatgrass (Pseudoroegneriaspicata)

Holly Prendeville, Francis Kilkenny and Brad St. Clair

 The locations for 16 common gardens test sites in the Blue Mountains, Columbia Plateau, 

Snake River Plain, and Northern Basin and Range ecoregions were finalized and sites 

prepared for establishment.

 The common garden sites were planted in the fall of 2014 with 20 replicate plugs from 78 

populations, one cultivar and one selected germplasm of bluebunch wheatgrass 

(Pseudoroegneria spicata).

Conservation, Adaptation, and Seed Zones for Key Great BasinSpecies

Richard C. Johnson, Erin K. Espeland, Matt Horning, Elizabeth Leger, Mike Cashman, and Ken

Vance-Borland

 Seed zones for Sandberg bluegrass (Poa secunda) across the Inter-mountain West and the 

Great Basin have been mapped and published.

 Seed zones have been modeled for Thurber’s needlegrass (Achnatherum thurberianum) 

for the Great Basin and will be mapped and submitted for publication in 2015.

 Basin wildrye (Leymus cinereus) data collection on 112 source populations in common 

gardens at two sites and three years has been completed. Initial analysis indicates climate 

linked genetic variation and interactions with ploidy are important for adaptation. Seed 

zones will be developed and published.

 Data collection in common gardens is completed for bottlebrush squirreltail (Elymus 

elymoides) and is being analyzed at the Rocky Mountain Research Station for seed zone 

development.
iv

 A common garden study of sulphur-flower buckwheat (Eriogonum umbellatum) collected 

in the Intermountain West and Great Basin was established and data will be collected in 

2015 and 2016.

Ecological Genetics of Big Sagebrush (Artemisia tridentata): GeneticStructure 

and Climate Based Seed Zone Mapping

Bryce Richardson, Nancy Shaw, and Matthew J. Germino

 Analyses of growth and seed yield indicate subspecies tridentata and wyomingensis have 

highly similar adaptive responses to climate.

 Overall, clinal patterns of growth and seed yield appear to be shaped by the accumulation 

of summer heat and the seasonal timing of precipitation.

 Plants from lower basins in the western regions of big sagebrush have increased growth 

and seed yield under hot-dry summers.

Factors Affecting Initial Establishment of Big Sagebrush Outplants: Land 

Treatments, Seed Sources, and ClimateEffects

Matthew J. Germino, Martha Brabec, and Bryce Richardson

 Land treatments such as mowing, herbicide application, and drill seeding affect the 

abundance of herbs that sagebrush seedlings interact (and apparently compete) with, and 

they influence initial survival of sagebrush outplants.

 Growth and initial survival of sagebrush seedlings differed among subspecies, cytotypes, 

and populations, and the variation was most related to differences in response to freezing, 

generally supporting the use of minimum temperatures in provisional seed zones.

 Paradoxically, mountain big sagebrush consistently expressed less ability to withstand 

freezing than the low-elevation subspecies, suggesting that further inquiry is needed to 

understand broad-scale climate adaptation of big sagebrush.

Species and Population-level Variation in Germination Strategies of Cold Desert 

Forbs

Elizabeth A. Leger and Sara Barga

 We identified the dominant germination strategies of ten Great Basin annual and 

perennial herbaceous species.

 A majority of species had no preference for after-ripening temperature.

 Species exhibited differences in their preference for cold stratification.

 Five species were identified as facultative winter germinators by their ability to germinate 

at cold (2oC) temperatures, including Agoseris grandiflora, Blepharipappus scaber, 

Collinsia parviflora, Cryptantha pterocarya.

 We found that Agoseris grandiflora, Collinsia parviflora, Cryptantha pterocarya, and

Microsteris gracilis were able to germinate under a broad range of conditions.

 In contrast, Gilia inconspicua, Mentzelia albicaulis, and Phacelia hastata possessed high 

levels of dormancy.

 All species displayed population level differences in germination strategies.

Dynamics of cold hardiness accumulation and loss in Sulphur-flower buckwheat 

(Eriogonum umbellatum)
v

Anthony S. Davis, Matthew R. Fisk, and Kent G. Apostol

 October 2013 - October 2014 field collections and sample processing completed for 

sulphur-flower buckwheat (Eriogonum umbellatum) cold hardiness study.

 Initial results indicate cold hardiness variation across sulphur-flower buckwheatspecies 

range and annual seasonality.

 Sulphur-flower buckwheat cold hardiness data analysis models in development.

Community Structure of Arbuscular Mycorrhizal Fungi Colonizing Wyoming Big 

Sagebrush Seedlings

Marcelo D. Serpe and Bill E. Davidson

 Pre-inoculation of Wyoming big sagebrush seedlings with native arbuscular 

mycorrhizae (AMF) increased colonization of the roots that developed after 

transplanting, but did not affect the AMF taxa colonizing the seedlings.

 Sagebrush seedlings were colonized by a diversity of AMF taxa. However, four taxa 

were dominant. Similar results were obtained for non-inoculated and inoculated 

seedlings as well as for seedlings harvested in different seasons or from roots collected 

at different depths.

Smoke-induced Germination of Great Basin Native Forbs

Robert Cox

 Extensive literature reviews for all focal species have been conducted and species have 

been prioritized by germination requirements and potential for smoke response

 19 species have been tested for smoke response in germination, with results being 

analyzed now.

Herbicide Impacts on Forb Performance in Degraded Sagebrush Steppe 

Ecosystems

Marie-Anne De Graff and Aislinn Johns

 Two greenhouse experiments were conducted in which we (1) applied a gradient of 

imazapic concentrations, and (2) varied the timing of application relative to seeding of 

Astragalus filipes, Achillia millefolium, and Sphaeralcea grossulariifolia.

 Soils used were collected in degraded burned and unburned sagebrush ecosystems (0-10 

cm depth) at the Morley Nelson Birds of Prey National Conservation Area.

PLANT MATERIALS AND CULTURALPRACTICES

Increasing the Availability and Integration of Great Basin Native PlantProject 

Tools for BLM Restoration Practitioners

Anne Halford

 Apply science and research to improve the identification and protection of resistant and 

resilient sagebrush-steppe landscapes and the development of biocontrols and other tools 

for cheatgrass control to improve capability for long-term restoration of sagebrush-steppe 

ecosystems.
vi

 To the extent practicable, utilize locally-adapted seeds and native plant materials 

appropriate to the location, conditions, and management objectives for vegetation 

management and restoration activities, including strategic sourcing for acquiring,storing, 

and utilizing genetically appropriate seeds and other plant materials native to the 

sagebrush-steppe ecosystem.

Developing Protocols for Maximizing Establishment of Great BasinLegume 

Species

Douglas A. Johnson and B. Shaun Bushman

 Recent greenhouse studies showed substantial progress in obtaining high germination 

rates of basalt milkvetch (Astragalus filipes).

 Seeded plots were planted at three sites in Oregon: Oregon State University (OSU) 

Ontario Research Farm, BLM site near Clarno, and a USFS field near Prineville. Three 

plantings have been attempted at each site, comprising a fall and spring planting. Fall 

2011/spring 2012 showed the benefits of non-scarified seed planted in early spring. Fall 

2013/spring 2014 resulted in no seedling establishment because of extremely low spring 

and summer precipitation; however, basalt milkvetch germinated and emerged the 

following year and plants from the previous year survived. Fall 2014 was planted in 

October and the spring 2015 will be planted in March.

 Seed collections were made of ecotypes of Utah trefoil (Lotus utahensis) in southern 

Utah, Nevada, and Utah. These collections will be evaluated in common gardens, and a 

DNA marker technique will be used to evaluate genetic diversity among collections.

Native Forb Increase and Research at the Great Basin Research Center

Kevin Gunnell and Jason Stettler

 Cushion Buckwheat (Eriogonum ovalifolium) common gardens to analyze:seed 

production, G1 production seed, and drought tolerance.

 Identify a practical and effective method to treat iron deficiency chlorosis (IDC) for 

improved establishment in Lupine species.

 Increase seed from wildland collections to support further research and increase 

distribution of native seed to commercial growers.

 Develop and test methods to increase germination and establishment in argronomic 

settings.

 Determine the effects of N-sulate® fabric on germination and establishment of native 

forb species in a wildland setting.

 Evaluate native forb establishment in Wyoming big sagebrush communities using four 

different pieces of planting equipment: Truax drill, Lawson aerator, Dixie pipe harrow, 

and Ely chain.

Plant Material Work at the Provo Shrub Sciences Lab

Scott Jensen

 Added 72 new database location records representing 21 species.

 Made 45 wildland seed collections representing three species.

 Distributed 24 sources of two species for seed increase.

 Distributed 38 sources of eight species for commercial seed production.

 Study installation evaluating emergence rates of 20 native forb species.
vii

 Study installation comparing emergence rates of Basin wildrye sources.

Aberdeen Plant Materials Center Report ofActivities

Derek Tilley

 Release of Amethyst Germplasm Hoary Tansyaster.

 Progress towards release of Wyeth buckwheat.

 Development of NRCS Plant Guides.

Bee Pollination and Breeding Biology Studies

James H. Cane

 In the aftermath of a huge wildfire in sage steppe, we sampled native bee faunas at 

scattered remnant stands of sunflowers growing in ditches along gravel roads thatserved 

as fire lines.

 Comparable quantitative samples of bees were taken at co-flowering sunflowers outside 

the burn. Most all of the bee species were non-social and soil nesting.

 The bee faunas largely survived the fire intact (but for one surface-nesting species). 

Females even continued to collect pollen for their nests.

 The fire had passed the sample sites at night, a time when female bees are in their nest 

(and thus safe underground).

 Another sample of bees and bloom in a year-old fire found much the same result at 

blooming Penstemon and Balsamorhiza that has also survived the fire to bloom the 

following year.

Seed Production of Great Basin Native Forbs

Clinton C. Shock, Erik B. Feibert, Joel Felix, Nancy Shaw, and Francis Kilkenny

 Plant establishment report.

 Trial of post emergence herbicides.

 Irrigation trials for the first set of species that we have worked on for 9 years.

 Irrigation trials on a second set of species that we have worked on for 5 years.

 Irrigation trials on a third set of species that we have worked on for 2 years.

SEED INCREASE

Coordination of Great Basin Native Plant Project Plant MaterialsDevelopment, 

Seed Increase, and Use

Berta Youtie

 Preparation and planting of bluebunch wheatgrass (Pseudoroegneria spicata) test sites in 

recently burned areas within seed zones to test the efficacy of using species-specific 

provisional seed zones and seed transfer guidelines in native plant restoration efforts after 

fires.

 Increase of native seed collections for the Great Basin Native Plant Project (GBNPP) in 

Eastern Oregon and Northern Nevada.
viii

SPECIES INTERACTIONS

Use of Native Annual Forbs and Early- Seral Species in Seeding Mixturesfor 

Improved Success in Great Basin Restoration

Keirith A. Snyder and Shauna M. Uselman

 Seedling emergence and early survival of early seral native species was generally 

greater than that of late seral native species when seeded with cheatgrass or 

medusahead.

 Native grasses were more successful than native forbs and shrubs in terms of seedling 

emergence and early survival, especially in a coarse-textured soil.

 Emergence and survival of medusahead was consistently very high on divergent soil 

types, supporting the need for preventive measures against its further spread in the 

Intermountain West.

 Early seral natives show promise for improving success of rangeland 

restoration/rehabilitation reseeding efforts in the Great Basin.

 When the early seral native forb bristly fiddleneck (Amsinckia tessellata) was able to 

establish, its substantial biomass and reproductive output suppressed the biomass and 

reproductive output of the exotic annual grass medusahead.

Restoration of Native Understory Plants in Degraded SagebrushSteppe 

Ecosystems

David A. Pyke and Kari Veblen

 Seeds and seedlings were monitored twice at three loamy Wyoming Big Sagebrush 

project sites, one each in the Snake River Plains, Owyhee Plateau, and Great Salt Lake 

MLRAs. Monitoring will be repeated in spring/summer 2015.

 Preliminary results suggest higher success of seedling plantings than seedings.

SCIENCE DELIVERY

Science Delivery for the Great Basin Native Plant Project

Corey Gucker

 Maintaining contact with Great Basin Native Plant Program (GBNPP) cooperators 

through regular correspondence, event alerts, and communication of project due dates.

 Updating and producing hard copy posters and brochures for Great Basin events and 

meetings.

 Regularly updating the GBNPP website with meeting announcements, new publications, 

and updated cooperator information.

 Helping to develop the agenda and secure presenters for the annual GBNPP meeting.

 Tracking and reporting GBNPP accomplishments.

CRESTED WHEATGRASSDIVERSIFICATION
ix

Evaluating Strategies for Increasing Plant Diversity in CrestedWheatgrass 

Seedings

Kent McAdoo and John Swanson

 Project objectives were to evaluate various treatments for reducing crested wheatgrassin 

near-monoculture rangelands and determine the effect of crested wheatgrass control 

methods on establishment of seeded native species.

 Mechanical treatment (disking) was less successful at reducing crested wheatgrassthan 

was herbicide.

 Herbicide, but not mechanical treatment, increased the density of seeded native speciesin 

both trials.

 Glyphosate treatments, but not disking or other herbicides, increased the density of 

seeded species in two separate trial studies by effectively reducing the competitive cover 

of crested wheatgrass.

The Ecology of Great Basin Native Forb Establishment

Jeremy James

 Simultaneously seeding crested wheatgrass and forbs does not decrease forb 

establishment.

 Field germination of native forbs can vary 20-fold across sites within a year but even in 

the best germination scenarios more than 98% of germinated seeds die before they 

establish.

 Seedbed favorability for germination and emergence varied across the nine study sites

but this variation did not significantly influence forb recruitment.

 Improving our ability to increase forb and forb diversity in restoration may be facilitated 

by identifying traits and populations that have greater germination and emergence 

probabilities or by applying seed treatments that accelerate germination or minimize 

exposure of seeds to abiotic stresses.

Presence of Native Vegetation in Crested Wheatgrass Seedings and the Influenceof 

Crested Wheatgrass on Native Vegetation

Kirk Davies and Aleta Nafus

 We measured basal cover, density, species richness and diversity on 121 sites that 

spanned a 54230 km2 area in southeastern Oregon.

 Plant community composition of crested wheatgrass stands was quite variable. Some 

seedings were a monoculture of crested wheatgrass while others contained a more diverse 

assemblage of native species.

 Environmental factors explained a range of functional group variability ranging from 

24% of annual forb density to 51% of annual grass density.

 Soil texture appeared to be the most important environmental characteristic explaining 

functional group cover and density 10-50 years post seeding and was included in the best 

regression models for all plant functional groups.

 Native vegetation was, for all functional groups, positively correlated with soils lower in 

sand content.

 Our results suggest that environmental differences explain a large amount of the 

variability of native vegetation in crested wheatgrass stands and this information can
x

likely be invaluable in assessing the effects of seeding crested wheatgrass and the 

potential for native vegetation to co-occupy seeded sites with crested wheatgrass.

RESTORATION STRATEGIES ANDEQUIPMENT

Seeding Native Species Following Wildfire in Wyoming Big Sagebrush (Artemisia 

tridentata ssp. wyomingensis): Effects of a New Fire Incident

Jeff Ott and Nancy Shaw

 An operational-scale experiment was carried out to test equipment and strategies for 

seeding native plants following wildfire on Wyoming big sagebrush (Artemisia tridentata 

ssp. wyomingensis) sites in the northern Great Basin. One of the study sites (Saylor 

Creek) was hit by a second wildfire two years after the first, creating an opportunity to 

examine resilience of native seedings to burning. Because the fire did not burn the entire 

study area, it was possible to compare burned and unburned sections and thereby evaluate 

fire impacts on plant species cover and density.

 Seeded species began to establish in seeded treatments the first year then increased 

dramatically in cover by the second year. By the third year, following the new fire, 

seeded species cover decreased to levels intermediate between the first and second years. 

Non-seeded species generally followed a similar trajectory, although cheatgrass (Bromus 

tectorum) did not decrease in non-seeded treatments where it faced less competition from 

perennials.

 Seeded forbs and shrubs fell into three groups based on density data: those that initially 

had high densities, but declined in density by the third year (Achillea millefollium, 

Penstemon speciosus, Artemisia tridentata ssp. wyomingensis), those with lower initial 

densities followed by an increase (Sphaeralcea munroana, Ericameria nauseosa), and 

Astragalus filipes, which had low initial density followed by a decrease.

 To assess whether changes in cover and density were due to the fire as opposed to other 

factors, we tested for deviations from expected ratios in burned and unburned sections of 

the study area. Results suggest that some species were negatively affected by the fire 

(Artemisia tridentata, Ericameria nauseosa, Achillea millefollium, Poa secunda), others 

were positively affected (Penstemon speciosus, Achnatherum hymenoides) and others 

minimally affected (Pseudoroegneria spicata, Elymus elymoides, Sphaeralcea munroana, 

Astragalus filipes).

A New Biological Control Strategy for Cheatgrass and Ventenata dubia Based on 

Fungal Endophytes

George Newcombe

 Manuscript accepted pending revision (PLOS ONE) on the ecology of Sordaria fimicola

and its effects on Bromus tectorum.

 Manuscripts being prepared on: the effects of Fusarium endophytes isolated from 

cheatgrass (Bromus tectorum) and ventenata (Ventenata dubia) on these two species and 

native bluebunch wheatgrass (Pseudoroegneria spicata); and the effects of Sordaria

fimicola (CID 323) on B. tectorum in field experiments in MPG Ranch, MT.

 Collaborated and shared work at a Biological Invasion Working Group of the Mountain 

Social Ecological Observatory Network (MtnSEON) meeting, at which were discussed
xi

the processes, patterns, and mechanisms of Ventenata dubia invasion in complex western 

landscapes.

 Collaborated and shared work at two Blue Mountains Working Group of MtnSEON 

meetings, focused on social-ecological systems (including annual grass invasion) 

research centered on the USFS Starkey Experimental Forest and Range.

 Conducted disease surveys in Latah County Idaho. While diseases were present in many 

neighboring grasses, no signs or symptoms of disease were found on invasive grass 

species ventenata.

 Conducted greenhouse and lab experiments on susceptibility of V. dubia to the wheat 

pathogen Fusarium culmorum, and tested the hypothesis that B. tectorum and V. dubia 

endophytic Fusarium strains should protect V. dubia from the effects of disease caused 

by F. culmorum.

 Collected specimens of non-native ventenata across northern Idaho, western Montana, 

and northeast Oregon. Samples currently in first stages of metagenomics analyses to look 

at fungal communities living within ventenata grass tissues.

 A manuscript is being prepared on the native range of Ventenata dubia. This invader is 

commonly called ‘North Africa grass’ but its current Eurasian range does not include 

populations in North Africa.
xii

TABLE OF CONTENTS

GENETICS AND SEED ZONES

FRANCIS KILKENNY

BRAD STCLAIR

Genetic Diversity and Genecology of Prairie Junegrass 

(Koeleria macrantha) and Squirreltail (Elymus elymoides)

1

HOLLY PRENDEVILLE

FRANCIS KILKENNY

BRAD ST. CLAIR

Testing the Efficacy of Seed Zones for Re-establishment and 

Adaptation of Bluebunch Wheatgrass (Pseudoroegneria 

spicata)

10

RICHARD C.JOHNSON

ERIN K. ESPELAND

MATT HORNING

ELIZABETH LEGER

MIKE CASHMAN

KEN VANCE-BORLAND

Conservation, Adaptation, and Seed Zones for Key Great 

Basin Species

14

BRYCE RICHARDSON

NANCY SHAW

MATTHEW J. GERMINO

Ecological Genetics of Big Sagebrush (Artemisia tridentata): 

Genetic Structure and Climate-based Seed Zone Mapping

23

MATTHEW J. GERMINO

MARTHA BRABEC

BRYCE RICHARDSON

Factors Affecting Initial Establishment of Big Sagebrush 

Outplants: Land Treatments, Seed Sources, and Climate 

Effects

29

ELIZABETH A. LEGER

SARA BARGA

Species and Population-level Variation in Germination 

Strategies of Cold Desert Forbs

37

ANTHONY S. DAVIS

MATTHEW R. FISK

KENT G. APOSTOL

Dynamics of Cold Hardiness and Loss in Sulphur-flower 

Buckwheat (Eriogonum umbellatum)

42

MARCELO D. SERPE

BILL E. DAVIDSON

Community Structure of Arbuscular Mycorrhizal Fungi 

Colonizing Wyoming Big Sagebrush Seedlings

48

ROBERTCOX Smoke-induced Germination of Great Basin Native Forbs 59

MARIE-ANNE DE GRAAFF

AISLINN JOHNS

Herbicide Impacts on Forb Performance in Degraded 

Sagebrush Steppe Ecosystems

60
xiii

PLANT MATERIALS AND CULTURALPRACTICES

ANNE HALFORD Increasing the Availability and Integration of Great Basin 

Native Plant Project Tools for BLM Restoration 

Practitioners

69

DOUGLAS A.JOHNSON

B. SHAUN BUSHMAN

Developing Protocols for Maximizing Establishment of 

Great Basin Legume Species

74

KEVIN GUNNELL

JASON STETTLER

Native Forb Increase and Research at the Great Basin 

Research Center

77

SCOTT JENSEN Plant Material Work at the Provo Shrub Sciences Laboratory 86

DEREK TILLEY Aberdeen Plant Materials Center Report of Activities 90

JAMES H. CANE Bee Pollination and Breeding Biology Studies 92

CLINTON C. SHOCK

ERIK B. FEIBERT

JOEL FELIX

NANCY SHAW

FRANCIS KILKENNY

Seed Production of Great Basin Native Forbs 95

SEED INCREASE

BERTA YOUTIE Coordination of Great Basin Native Plant Project Seed 

Increase and Eastern Oregon/Northern Nevada Seed 

Collections

145

SPECIES INTERACTIONS

KEIRITH A. SNYDER

SHAUNA M. USELMAN

Use of Native Annual Forbs and Early-seral Species in 

Seeding Mixtures for Improved Success in Great Basin 

Restoration

149

DAVID A. PYKE

KARI VEBLEN

Restoration of Native Understory Plants in Degraded 

Sagebrush Steppe

155
xiv

SCIENCEDELIVERY

COREY GUCKER Science Delivery for the Great Basin Native Plant Project 157

CRESTED WHEATGRASSDIVERSIFICATION

KENT MCADOO

JOHN SWANSON

Evaluating Strategies for Increasing Plant Diversity in 

Crested Wheatgrass Seedings

158

JEREMY JAMES The Ecology of Great Basin Native Forb Establishment 168

KIRK DAVIES

ALETA NAFUS

Recruitment of Native Vegetation into Crested Wheatgrass 

Seedings and the Influence of Crested Wheatgrass on Native 

Vegetation

175

RESTORATION STRATEGIES AND EQUIPMENT

JEFF OTT

NANCY SHAW

Seeding Native Species Following Wildfire in Wyoming Big 

Sagebrush (Artemisia tridentata ssp. wyomingensis): Effects 

of a New Fire Incident

181

GEORGE NEWCOMBE

DAVID GRIFFITH

SARA ASHIGLAR

A New Biological Control Strategy for Cheatgrass and

Ventenata dubia Based on Fungal Endophytes

191
1

Project Title Genetic Diversity and Genecology of Prairie Junegrass

(Koeleria macrantha) and Squirreltail (Elymus 

elymoides)

Project Agreement No. 13-IA-11221632-161

Principal Investigators and Contact Information

Francis Kilkenny, Research Biologist 

USDA Forest Service

Rocky Mountain Research Station 

322 SW Front Street Suite 401

Boise, ID 83702-7373

208.373.4376, FAX 208.373.4391

ffkilkenny@fs.fed.us

Brad St. Clair, Research Geneticist 

USDA Forest Service

Pacific Northwest Research Station 

3200 SW Jefferson Way

Corvallis, OR 97331- 4401

541.750.7294, FAX 541.750.7329

bstclair@fs.fed.us

Project Description

Prairie junegrass

Prairie junegrass (Koeleria macrantha) is a highly variable, moderately long-lived, coolseason 

perennial bunchgrass that grows 15 to 60 cm (0.5 to 2 ft) tall. It is adapted to a wide variety of 

climates and soils and is an important component of many native plant communities. Prairie 

junegrass is often used in seed mixes for restoration of native prairie, savanna, coastal scrub, 

chaparral, and open forest habitats across much of North America. Good drought tolerance and 

fibrous roots also make it beneficial for revegetation and erosion control on mined lands, over 

septic systems, in construction areas, on burned sites, and in other disturbed areas. There is a 

need for greater genetic knowledge of prairie junegrass to ensure adapted populations are used 

for restoration and revegetation projects. Most information will be generated from two common 

garden studies conducted on separate and contrasting environments in Oregon.

Objectives

1) Use common gardens to determine the magnitude and patterns of genetic variation among 

prairie junegrass populations from a wide range of source environments in Oregon, 

Washington, and Idaho.

2) Relate genetic variation to environmental variation at collection locations.

3) Develop seed transfer guidelines.
2

Methods

Seed was collected from endemic stands of prairie junegrass during the summers of 2003 to 

2006. Populations came from Oregon, Washington, and adjacent areas of Idaho, primarily east of 

the Cascade Mountains. Two families (maternal parents) were sampled from each of 114 

populations, while only a single family was sampled from 12 populations, for a total of 240 

families from 126 populations. Common garden studies were established in 2008 at two 

contrasting sites in Oregon: the USDA Natural Resources Conservation, Plant Materials Center 

in Corvallis, Oregon, and Oregon State University’s Agricultural Experiment Station at Powell 

Butte, Oregon. A variety of traits were measured in 2009 and 2010 assess plant growth, 

morphology, phenology, and fecundity. Analyses have been done to evaluate differences among 

test sites, years, populations, families within populations, and interactions between these factors. 

Analyses have also investigated the relationship between population variation and climatic 

variation at source locations. Maps of genetic variation across the landscape have been produced, 

and preliminary seed zones have been delineated based on the results.

Results

Traits at Corvallis were significantly different from traits at Powell Butte. Plants at Corvallis 

were larger and had earlier phenology in both 2009 and 2010. Plants at Powell Butte flowered 

much more profusely than plants at Corvallis in 2009, but Corvallis plants had a higher 

inflorescence number in 2010. Variation among populations for size traits was much greater 

when plants were grown at Corvallis in both years, and plants grown at Powell Butte generally 

had higher variation among populations in phenology traits for both years. The percentage of the 

total phenotypic variation (populations, families, and residual) found within populations was 

high for most traits, especially crown width, leaf width, leaf ratio, and heading and bloom dates.

The population × site interaction was significant for most traits. Correlations of population 

means between test sites were higher in 2009 than in 2010, and correlations between years within 

sites were generally high. Populations that had wide crowns, more flowers, wide leaves, upright 

habit, and later phenology at one site were generally the same as those at the othersite.

Traits that had large population variation and were correlated with climates at the source 

locations were biomass, crown width, inflorescence number, leaf width, leaf ratio, heading date, 

and bloom date. As might be expected, leaf width was strongly correlated with leaf ratio

(r = - 0.73), and bloom date was strongly correlated with heading date (r = 0.93) though only 

moderately correlated with maturation date (r = 0.46). These traits are generally correlated with 

temperatures, precipitation, and aridity (as measured by the heat moisture index, which is a 

function of the ratio of temperature to precipitation). Regression equations of traits on source 

climates used various combinations of temperature, precipitation, or aridity variables. Regression

models were used to map patterns of variation for biomass, inflorescence number, leaf ratio, and 

bloom date. The R2 values ranged from 0.29 to 0.36. Although the magnitudes and nature ofthe 

relationships between the source climates and traits differ somewhat between the four traits, 

general patterns of variation emerged. Populations from the Columbia Basin had less biomass, 

narrower leaves, fewer inflorescences, and later phenology when grown together in a common 

environment. Populations from the Blue Mountains had greater biomass, wider leaves, more

inflorescences, and earlier phenology. Populations from the northern Great Basin were similar to 

the Columbia Basin in having narrow leaves and lower biomass but were more similar to the
3

Trait

X env. correlation

Blue Mountains in having earlier phenology. The few populations from western Oregon had 

wider leaves, greater biomass, and the latest phenology. From an adaptation perspective, we 

expect narrow leaves from areas of greater aridity, less precipitation, and warmer summer 

temperatures, and we expect earlier phenology from areas with colder winter temperatures(i.e., 

selection for quicker responses to springtime warming).

To determine whether climatic seasonality at source locations was related to traits expressed in 

the common gardens, Pearson correlations were run between monthly climate variables (monthly 

average temperature, and monthly average precipitation) and three important phenotypic traits 

(heading date, leaf length:width ratio, and biomass; Figure 1). High leaf length:width ratio 

values, thinner leaves with respect to size, were strongly correlated with high temperaturesfrom 

March through November, whereas high biomass and later heading date values were correlated 

with high temperatures for October or November through March. This makes sense, given that 

the environments conducive to high biomass and late heading tended to be the milder climates, 

with warmer winter temperatures, of western Washington and Oregon on the western side of the 

Cascades, while the environments conducive to thin leaves tended to be the more continental

climates of the Intermountain West, with their hot summers. High biomass and late heading were 

also correlated with high precipitation from October through March or April, again indicating an 

association with the environments west of the Cascades. Whereas, high values of leaf 

length:width ratio were negatively correlated with precipitation throughout the entire year, 

indicating that thin leaves are generally associated with dry areas.

The general patterns that emerged from this preliminary analysis were largely congruent with 

large-scale ecoregions, suggesting that Omernick’s Level III ecoregions may be useful forseed 

zones. These seed zones are also similar to other published bunchgrass seed zones. Preliminary 

seed zones are presented in Figure 2.

0.5 0.5

0 0

-0.5 -0.5

Month of average temperature value Month of average precipitation value

Figure 1. Trait correlations with monthly average temperature (left graph) and monthly 

average precipitation (right graph) for heading date (blue line), leaf width:length ratio 

(red), and biomass (orange). Areas outside the dashed lines represent significance of 

greater than 0.05.

Trait

X env. correlation

Jan

Feb 

Mar 

Apr 

May 

Jun 

Jul 

Aug 

Sep 

Oct 

Nov 

Dec

Jan

Feb 

Mar 

Apr 

May 

Jun 

Jul 

Aug 

Sep 

Oct 

Nov 

Dec
4

Figure 2. Preliminary seed zones for prairie junegrass where each color represents a 

different seed zone. Seed zones are based on dividing the area in three principle

components that represent small, intermediate, or large plant size, early or late phenology, 

and narrow or wide leaves.

Squirreltail

Squirreltail (Elymus elymoides) is a cool-season, short-lived, self-pollinating, perennial 

bunchgrass of semi-arid regions of western North America. It is found in a wide variety of 

habitats from the Pacific Coast to the Great Plains and from Canada to Mexico at elevationsfrom 

2,000 to 11,500 feet (600-3,500 m). Squirreltail is considered a “workhorse” species in native 

plant restoration. It can rapidly colonize disturbed sites, is relatively fire-tolerant, and is a 

potential competitor with medusahead (Taeniatherum caput-medusae) and cheatgrass (Bromus 

tectorum). There are seven cultivars and pre-variety germplasm (PVG) collections ofsquirreltail 

that have been or are being developed for restoration purposes; however, only one variety, Toe 

Jam Creek, is currently used extensively. Previous studies indicate that there is strong ecotypic 

divergence in squirreltail. However, few studies have evaluated genetic variation in relation to 

climatic factors across the Great Basin or the greater range of the species in a large set of diverse 

populations or to compare the mean and variation of cultivars with that of the species as a whole. 

Determining the extent to which adaptive genetic variation is related to climatic variation is 

needed to ensure that the proper germplasm is chosen for revegetation and restoration.
5

Furthermore, comparisons of cultivars with the natural range of variation will address questions 

of the suitability of cultivars over larger areas.

Objectives

1) Explore genetic variation in putative adaptive traits in bottlebrush squirreltail from a wide 

range of source environments in the inland West.

2) Relate genetic variation to environmental variation at collection locations.

3) Develop seed transfer guidelines.

Methods

With the help of collaborators from a variety of organizations interested in restoration using 

squirreltail (including the Forest Service, Bureau of Land Management, National Park Service, 

Fish and Wildlife Service, Department of Defense, the Nature Conservancy, Washington 

Department of Natural Resources, and the Yakama Nation), seed was collected from two 

maternal families from 110 populations from five western states, primarily from the northern and 

central Great Basin. In addition to the squirreltail populations, we obtained seed from cultivars 

and PVGs of squirreltail for inclusion in the common garden studies. The cultivars and PVGs 

included Toe Jam Creek, Fish Creek, Rattlesnake, Tusas, Pleasant Valley Antelope Creek, and 

Sand Hollow (E. multisetus). Common garden studies have been established at three contrasting 

sites: 1) Reno, Nevada, 2) Powell Butte, Oregon, and 3) Central Ferry, Washington. A variety of 

traits were measured to assess plant growth, morphology, phenology, and fecundity in 2009 and 

2010. Analyses have been done to evaluate differences among test sites, years, populations, 

families within populations, and interactions between these factors. Analyses have also 

investigated the relationship between population variation and climatic variation at source 

locations. All wild collected populations were identified to subspecies. Maps of genetic variation 

across the landscape are in progress, and preliminary seed zones will be delineated based on the 

results.

Results

Site, year, subspecies, population, and family were all significant contributors to variation in 

most measured traits. Inflorescence number was not significantly affected by either subspecies or 

population, and family did not significantly affect flower maturity or leaf length. When broken 

down into components, most of the explained variation in all traits was due either to year or site 

effects. Year explained 72.7 to 78.8% of the variation in phenological traits, 41.4% of the 

variation in reproductive traits, and between 3.0 to 32.1% of the variation in size traits. Site 

explained between 6.0 and 9.0% of the variation in phenological traits, 33.3% of the variation in 

reproductive traits, and between 21.6 and 69.2% of the variation in size traits. Subspecies never 

explained more than 4.8% (biomass) and population never explained more than 7.5% (leaf 

width) of the variation in any traits. The unexplained variation ranged from 9.7 to 39.0%.

Altogether, this suggests that squirreltail traits are highly plastic, with phenological traits varying 

primarily by year and size traits varying primarily by planting site.

To determine the relationships between squirreltail traits and climate variables, correlations 

between phenotypic traits and annual climate variables were run. In this report, the strong 

correlations between three traits (heading date, inflorescence number, and biomass) and four 

climate variables (mean warmest month temperature [MWMT], temperature differential [TD],
6

mean annual precipitation [MAP], and annual heat:moisture index [AHM]) are shown (Figure 3). 

For each of the shown traits, the correlations are in the same direction, though the direction of 

the correlation changes depending on the climate variable. For MWMT, correlations for heading 

date, inflorescence number, and biomass are all negative, suggesting that small plants with low 

reproduction and late flowering are associated with hotter environments. Negative correlations 

are also present for TD and AHM, again suggesting that small, late, and low reproduction 

populations are associated with more continental (TD) and more arid (AHM) environments. All 

three traits were positively correlated with MAP, indicating that large, early, high-reproduction 

plants are associated with wetter environments. These correlations, particularly associations with 

above ground biomass, are generally in line with other bunchgrassstudies.

Leaf length:width ratio, which has tended to have a strong correlation with aridity variables,such 

as annual heat:moisture index, in other bunchgrasses in the western United States, did not show 

as strong correlations in squirreltail (R = 0.23 for leaf length:width ratio × AHM). There are two 

reasonable possibilities for this discrepancy. First is the possibility of phenotypic plasticity

driven by garden site by trait interactions reduced the strength of the correlations (R for leaf 

length:width ratio × AHM for Central Ferry = 0.14, Powell Butte = 0.29, Reno = 0.04). This 

makes sense, given that Reno was the driest site tested and also had the lowest correlation, 

indicating that all plants in Reno demonstrated similar leaf thinness. Second, many more 

squirreltail populations tested in this study tended to be primarily from more arid locationsthan 

the tested populations of other bunchgrasses. For example, the average AHM for tested 

squirreltail populations was 59 (± 22 S.D.) whereas the average AHM for tested prairie junegrass 

populations was 34 (±15 S.D.). This indicates that a majority of squirreltail populations may 

have been under selection pressure to reduce moisture loss, which may have reduced the 

variation in leaf thinness among populations.

Management Applications and Seed Production Guidelines:

These studies indicate that populations of prairie junegrass and squirreltail differ greatly across 

the landscapes of the Great Basin, Columbia Basin, Blue Mountains, and adjacent ecoregions for 

traits of plant size, flowering phenology, and leaf width. Much of the variation in prairie 

junegrass is associated with climates of the source locations, indicating adaptive significance.

However, even though genetic differences exist, squirreltail appears to have highly plastic traits, 

suggesting that most ecotypes are broadly adapted. Seed zones for prairie junegrass are presented 

for recommendations of bulking seed collections and producing plant material for adapted, 

diverse, and sustainable populations for revegetation and restoration. Different levels of 

acceptable risk may be accommodated by choosing whether or not to bulk materials from the 

same zones but in different Level III ecoregions. Results indicate that sources may be 

consolidated over fairly broad areas of similar climate, and thus may be shared between districts, 

forests, and different ownerships.
7

R = -051 R = -0.38

R = -0.51 R = -0.32

R = 0.52 R = 0.33

R = -0.58 R = -0.35 Inflorescence nu

mber Inflorescence nu

mber Inflorescence nu

mber Inflorescence nu

mber

Bio

mass Bio

mass Bio

mass Bio

mass

180

170

200 80

150 60

160 100 40

150 50 20

140

12 16 20 24 28

MWMT

0

12 16 20 24 28

MWMT

0

12 16 20 24 28

MWMT

180

170

200 80

150 60

160 100 40

150 50 20

140

12 16 20 24 28

TD

0

12 16 20 24 28

TD

0

12 16 20 24 28

TD

180 200 80

170 150 60

160

150

140

0 500 1000 1500

MAP

100

50

0

0 500 1000 1500

MAP

40

20

0

0 500 1000 1500

MAP

180 200 80

170 150 60

160 100 40

150 50 20

140

0 50 100 150

AHM

0

0 50 100 150

AHM

0

0 50 100 150

AHM

Figure 3. Correlations between squirreltail traits heading date, inflorescence 

number, and biomass and source climate variables mean warmest month temperature 

(MWMT), temperature differential (TD), mean annual precipitation (MAP), and 

annual heat:moisture index (AHM). Pearson correlation values are displayed.

R = -0.43

R = -0.42

R = 0.53

R = -0.54

Heading date Heading date Heading date Heading date
8

Publications

Kilkenny, F. F.; St. Clair, J. B.; Darris, D.; Horning, M.; Erickson. V. [In preparation]. 

Genecology of prairie junegrass (Koeleria macrantha). Evolutionary Applications.

Kilkenny, F. F.; St. Clair, J. B.; Horning, M. 2013. Climate change and the future of seed zones. 

National Proceedings: Forest and Conservation Nursery Associations - 2012. Proceedings 

RMRS-P-69. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station. p. 87- 89.

St. Clair, J. B.; Kilkenny, F. F. Genecology of prairie junegrass (Koeleria macrantha): Final 

Report. Technical report for NRCS Corvallis Plant Materials Center.

Presentations

Kilkenny, F. F.; 2015. Seed Zones and Climate Change. Great Basin Fires Science Exchange, 

2015 Webinar Series; 2015 May 20th.

Kilkenny, F. F. 2015. Predicting the effects of climate change on bunchgrass populations using 

common garden studies. 3rd National Native Seed Conference; 2015 April 13-16; Santa Fe, NM.

Kilkenny, F. F. 2014. Characterization of current and future climates within and among seed 

zones to evaluate options for adapting to climate change. Society for Ecological Restoration, 

Northwest and Great Basin Chapters, Joint Regional Conference; 2014 October 6-10; Redmond, 

OR.

Kilkenny, F. F. 2013. Seed zones. Restoration and reforestation technical exchange study tour; 

USFS Lucky Peak Nursery, Boise, ID; 2013 September 9. USAID sponsored tour for Ecuadoran 

government officials.

Kilkenny, F. F.; St. Clair, J. B.; Horning, M.; Johnson, R. C.; Leger, E.; Shaw, N. 2013. 

Genecology of three native bunchgrasses: implications for management during climate change. 

Intermountain Native Plant Summit; 2013 March 26-27; Boise, ID.

Kilkenny, F. F.; St. Clair, J. B. 2013. Characterization of current and future climates within and 

among seed zones to evaluate options for adapting to climate change. 2nd National Native Seed 

Conference; 2013 April 8-11; Santa Fe, NM.

Kilkenny, F. F.; St. Clair, J. B.; Horning, M.; Johnson, R. C.; Leger, E.; Shaw, N. 2013. 

Genecology and seed zones for prairie junegrass and bottlebrush squireltail. 2nd NationalNative 

Seed Conference; 2013 April 8-11; Santa Fe, NM.

Kilkenny, F. F.; St. Clair, J. B.; Shaw, N.; Richardson, B. 2013. Efficiency of using speciesspecific seed zones for re-establishment of native bunchgrass populations after fire. Poster 

presented at the 2nd National Native Seed Conference; 2013 April 8-11; Santa Fe, NM.
9

Additional Products

This study provides seed zones and seed transfer guidelines for developing adapted plant 

materials of prairie junegrass for revegetation and restoration in the Great Basin and adjacent 

areas and guidelines for conservation of germplasm within the National Plant Germplasm 

System. Results for prairie junegrass are being submitted to a peer-reviewed journal and will be 

disseminated through symposia, field tours, training sessions, and/or workshops.
10

Project Title Testing the Efficacy of Seed Zones for Re-Establishment

and Adaptation of BluebunchWheatgrass 

(Pseudoroegneria spicata)

Project Agreement No. 13-IA-11221632-161

Principal Investigators and Contact Information

Holly Prendeville, Research Geneticist 

USDA Forest Service

Pacific Northwest Research Station 

3200 SW Jefferson Way

Corvallis, OR 97331-4401

541.750.7300, FAX 541.750.7329

hollyrprendeville@fs.fed.us

Francis Kilkenny, Research Biologist 

USDA Forest Service

Rocky Mountain Research Station 

322 E Front Street, Suite 401

Boise, ID 83702

208.373.4376, FAX 208.373.4391

ffkilkenny@fs.fed.us

Brad St. Clair, Research Geneticist 

USDA Forest Service

Pacific Northwest Research Station 

3200 SW Jefferson Way

Corvallis, OR 97331-4401

541.750.7294, FAX 541.750.7329

bstclair@fs.fed us

Project Description

Objectives

 Evaluate adaptation (i.e. establishment, survival, growth and reproduction) of bluebunch 

wheatgrass (Pseudoroegneria spicata) populations from local seed zones relative to nonlocal seed zones.

 Model adaptation as a function of source location and common garden climates.

 Characterize traits and climatic factors important for adaptation.

 Model effects of climate change on native populations and consequences of assisted 

migration for responding to climate change.

Methods

Previous research funded by the Great Basin Native Plant Project (GBNPP) found that 

bluebunch wheatgrass (Pseudoroegneria spicata) populations differed in traits important for 

adaptation to precipitation and temperature (St. Clair et al. 2013). From this work, seed zones
11

were delineated for the Interior Northwest including the Great Basin, Snake River Plain, 

Columbia Plateau, and Blue Mountains (St. Clair et al. 2013). In this study we investigate the 

efficacy of these seed zones by comparing differences in establishment, survival and 

reproduction of bluebunch wheatgrass populations from local seed zones compared to non-local 

seed zones. We hypothesize that in the long-term, populations from local seed zones will better 

establish, survive and reproduce. To test this hypothesis, we established a reciprocal transplant 

study in two broad regions (i.e., transects) each with eight seed zones and each seed zone 

represented by four to five wild populations. The two broad regions include (1) a transect from 

the hot, dry climates of the lower Snake River Plain to the cool, somewhat wet climates of the 

transverse ranges of the Great Basin to the cold, dry climates of the upper Snake River Plain; and

(2) a transect from the hot, dry climates of the Columbia Plateau to the cool, wet climates of the 

Blue Mountains to the cold, dry climates east of the Blue Mountains.

Eight common gardens were established in each transect with each garden representing the 

climate of the local seed zone. Each common garden was planted with bluebunch wheatgrass 

from each seed zone within a region (transect). In addition to bluebunch wheatgrass populations 

from each seed zone, one widely used variety and one selected germplasm were included at each 

common garden. The experimental design at each common garden is a split-plot design with a 

plot represented by 25 plants (five plants per population) from each seed zone and the subplots as 

five populations per seed zone. Each site includes four replicates. Planting spacing is 0.6 m x 0.8 

m between plants.

Results and Discussion

This study was initiated in 2013 beginning with seed collections from 138 populations from 

throughout the two broad regions. From these, 78 populations were chosen for inclusion in the 

study based on their distribution within each of the seed zones and the amount of available seed 

(only four populations, not five, were available from the hottest, driest seed zone within each 

transect). The University of Idaho was contracted to produce the containerized planting stock for 

the study in 2014. Potential collaborators were contacted to help with providing common garden 

sites on their lands. In August-December 2014, 16 common gardens were prepared and 

established. Most common gardens are protected from large mammal herbivory with fencing.

Most common gardens were planted with four replicates of 39 populations from eight seed 

zones, one cultivar, and one selected germplasm for a total of 1,000 plants surrounded by two 

rows of buffer plants. A few sites included an extra treatment of a seed zone from the other 

transect or an extra selected germplasm.

Future Plans

The remaining common gardens at risk of large mammal herbivory will be fenced. In the spring 

and summer of 2015 and 2016, monitoring and data collection for each of the 16 common 

gardens will occur. In the spring, plant survival, reproductive phenology, occurrence of 

herbivory, and presence of fungus will be recorded at each common garden. Weather data 

loggers will be placed at each common garden to record local temperature during the course of 

the study. Common gardens will be managed to control weeds to allow for establishment to be 

largely a function of local climate rather than differences among test sites in competing 

vegetation. At each common garden, plant survival and reproduction will be recorded along with 

occurrence of herbivory and presence of fungus. Crown width, a measure of plant size including
12

biomass, will be measured on each plant. A single representative leaf and reproductive stalk will 

be collected from each plant to measure leaf width and seeds per reproductive stalk, respectively. 

These measurements will be repeated in 2016. Measurements will continue at each site, every 

year for an indefinite length of time. Reporting of early results will be presented after the first 

two growing seasons (2015 and 2016).

Management Applications and/or Seed Production Guidelines

Bluebunch wheatgrass is an ecologically important restoration species (Daubenmire 1942; Harris 

1967). This study will examine the efficacy of seed zones for bluebunch wheatgrass to ensure 

successful establishment and allow for long-term adaptation by maintaining genetic diversity.

The results of this study will help land managers determine sources of bluebunch wheatgrass 

populations for post-fire restoration of an area. Furthermore, this study will explore the 

consequences of changing climates for adaptation by substituting space for time to evaluate 

different populations in different climates. Results from this portion of the study will aid land 

managers in maintaining and restoring areas considering future climate changes. Long-term 

productivity and adaptation will be modeled to allow evaluation of trade-offs between different 

management options for current and future climates. Population movement guidelines and 

associated seed zones can be adjusted based on results from this study and management 

objectives.

Presentations

St. Clair, B.; Kilkenny, F.; Shaw, N. 2014. Progress on establishment of reciprocal transplant

tests to study adaptation of bluebunch wheatgrass. Great Basin Native Plant Annual Meeting; 

2014 March 17-18; Boise, ID.

St. Clair, J. Bradley; Kilkenny, Francis F.; Shaw, Nancy L. 2014. Testing the efficacy ofseed 

zones for bluebunch wheatgrass. Great Basin Native Plant Project Annual Meeting; 2014 March 

17-18; Boise, ID. [Available at

http://www.fs.fed.us/rm/boise/research/shrub/projects/PowerPoint_Presentations/2013/StClair.pd

f].

St. Clair, J. Bradley. 2014. Development and use of seed zones to ensure adapted plant materials 

for restoration: Introduction. Society of Ecological Restoration Regional Conference, Northwest 

and Great Basin Chapters; 2014 October 6-10; Redmond, OR.

Additional Products

Seed zone maps and map layers for bluebunch wheatgrass are available for download and use at 

the USFS Western Wildland Environmental Threat and Assessment Center. [Available at

http://www. fs. fed. us/wwetac/threat_map/SeedZones_Intro.html].

References

Daubenmire, R. F. 1942. An ecological study of the vegetation of southeastern Washington and 

adjacent Idaho. Ecological Monographs. 12: 53-79.

Harris, G. A. 1967. Competitive relationships between Agropyron spicatum and Bromus 

tectorum. Ecological Monographs 37: 89-111
13

St. Clair, J. B.; Kilkenny, F. F.; Johnson, R. C.; Shaw, N. L.; Weaver, G. 2013. Genetic variation 

in adaptive traits and seed transfer zones for Pseudoroegneria spicata (bluebunch wheatgrass) in 

the northwestern United States. Evolutionary Applications 6: 933-948
14

Project Title Conservation, Adaptation and Seed Zones for KeyGreat

Basin Species

Project Agreement No. 10-IA-11221632-023 and 14-IA-11221632-013

Principal Investigators and Contact Information

R. C. Johnson, Research Agronomist 

USDA-Agricultural Research Service 

Western Regional Plant Introduction Station 

Box 646402, Washington State University 

Pullman, WA 99164

509.335.3771

rcjohnson@wsu.edu

Erin K. Espeland, Research Ecologist 

USDA- Agricultural Research Service 

Northern Plains Agricultural Research Lab 

Pest Management Research Unit

1500 N Central AVE, Sidney MT 59270 

406.433.9416

erin.espeland@ars.usda.gov

Matt Horning, Geneticist

U. S. Forest Service, Pacific Northwest Region 

63095 Deschutes Market Road

Bend, OR 97701

541.383.5519

mhorning@fs.fed.us

Elizabeth Leger, Associate Professor of Plant Ecology 

Department of Natural Resources and Environmental Science 

Fleishmann Agriculture, Room 130

University of Nevada, Reno 

775.784.7582

ealeger@gmail.com

Mike Cashman, Biologist

USDA-ARS, Western Regional Plant Introduction Station 

Box 646402, Washington State University

Pullman, WA 99164

509.335.6219 FAX 509.335.6654

mjcashman@wsu.edu
15

Ken Vance-Borland, GIS Coordinator 

Conservation Planning Institute

NW Wynoochee Drive 

Corvallis, OR 97330

kenvb@consplan.net

Project Description

Native plant species are critical for wildlife habitat, livestock grazing, soil stabilization, and 

ecosystem function in the Western United States and the Great Basin. Yet current germplasm 

releases of these species do not specifically match natural genetic diversity with local climates 

which often drive adaptation. We are developing seed zones that match climate with genetic 

variation that facilitate management decisions, ensure restoration with adapted plant material, 

and promote biodiversity needed for future natural selection with climate change. Over the 

course of the project, seed zones have been completed and published for tapertip onion (Allium 

acuminatum Hook.), Indian ricegrass (Achnatherum hymenoides [Roem. & Schult. ] Barkworth), 

bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] Á. Löve) and Sandberg bluegrass (Poa 

secunda J. Presl; Johnson et al. 2012; Johnson et al. 2013; St. Clair et al. 2013; Johnson et al.

2015). Ongoing work with additional Great Basin species includes Basin wildrye (Leymus 

cinereus [Scribn. & Merr.] Á. Löve), Thurber's needlegrass (Achnatherum thurberianum [Piper] 

Barkworth), bottlebrush squirreltail (Elymus elymoides [Raf.] Swezey), and Sulphur-flower 

buckwheat (Eriogonum umbellatum Torr.).

Figure 1. Schematic showing the steps for genecology studies to establish seed zones.

Our objectives are to:

1. Collect key Great Basin species and establish common garden studies quantifying genetic 

variation in plant traits associated with phenology, production, and morphology.

2. Uncover adaptive plant traits in common gardens linked to seed source location climates 

and develop composite plant traits using multivariate statistics.

3. Develop regression models linking composite plant traits with source climates and map 

seed zones for use in restoration.

Germplasm

collection

Common garden

evaluations for 

genetic traits

Link plant

traits to 

source

Multivariate

traits for data 

reduction

Regression modeling of

plant traits with source 

location climate

GIS mapping plant

of plant traits with
16

Sandberg bluegrass

Genetic variation for potentially adaptive traits of the key restoration species Sandberg bluegrass 

(Poa secunda J. Presl; Johnson et al. 2012, 2013, 2015; St. Clair et al. 2013) was assessed over 

the Intermountain Western United States in relation to source population climate (Johnson et al. 

2015). Common gardens were established at two Intermountain West sites with progeny from 

two maternal parents from each of 130 wild populations. Data was collected over 2 years at each 

site on 15 plant traits associated with production, phenology, and morphology. Analyses of 

variance revealed strong population differences for all plant traits (P<0.0001), indicating genetic 

variation. Both the canonical correlation and linear correlation

established associations between 

source populations and climate 

variability. Populations from 

warmer, more arid climates had 

generally lower dry weight, earlier 

phenology, and smaller, narrower 

leaves than those from cooler, 

moister climates. The first three 

canonical variates were regressed 

with climate variables resulting in 

significant models (P < 0.0001) used

to map 12 seed zones. Of the 700 

981 km2 mapped, four seed zones 

represented 92% of

the area in typically semi-arid and 

arid regions (Figure 2). The 

association of genetic variation with 

source climates in the Intermountain 

West suggested climate driven 

natural selection and evolution 

(Johnson et al. 2015). We

recommend seed transfer zones and

population movement guidelines to 

enhance adaptation and diversity for 

large-scale restoration projects

Figure 2. Proposed seed zones for Sandberg bluegrass in

the Intermountain West and Great Basin regions.

involving Sandberg bluegrass. Populations of plants from seed zones are being grown in the 

greenhouse and will be transplanted to the field for seed increase in spring 2015, and then further 

increased for use in large scale restoration.

Thurber's needlegrass

Thurber's needle grass common gardens were established at Central Ferry, WA, and Reno, NV in 

2010. The study consisted of 67 source locations predominantly from the Northern and Central 

Basin and Range ecoregions. Data were collected in 2012 and 2013 on numerous phenology, 

production, and morphological traits. Data analyses revealed extensive genetic variation in 

Thurber’s needlegrass across the Great Basin sampling region (Table 1). Genetic variation in 

wild populations of grasses and forbs has been reported for other species from the Great Basin
17

and Intermountain West (Johnson et al. 2010, 2012, 2013, 2015; St. Clair et al. 2013) and is 

generally observed in common garden research in other regions (St. Clair et al. 2005; Leimu and 

Fischer 2008; Horning et al. 2010). Interactions between source populations in different garden 

environments showed the expected presence of plasticity, that is, variation associated with 

different environments (Johnson et al. 2013, 2015; St. Clair et al. 2013).

Table 1. Summary of F-ratios and P-values from analyses of variance for Thurber’s needlegrass traits measuredin 

common gardens at Central Ferry, WA and Reno, NV on source populations collected from Great Basin overtwo 

years (2012 and 2013).

Trait mean Site(S) Pop. (P) SxP Year(Y) YxS

F P F P F P F P F P

Phenology†

Heading, day of year 135 27.9 <.01 8.9 <.01 2.01 <.01 187.6 <.01 27.8 <.01

Blooming, day of year 133 17.2 <.01 6.9 <.01 1.8 <.01 78.8 <.01 2.0 0.16

Maturity, day of year 161 36.1 <.01 3.4 <.01 1.47 0.01 2109.8 <.01 . .

Morphology‡

Leaf length x width, 

cm

2 1.64 263.2 <.01 4.2 <.01 1.47 0.01 125.4 <.01 147.4 <.01

Leaf length to width 74.5 41.4 <.01 1.7 <.02 1.42 0.02 27.9 <.01 85. 6 <.01

Culm length, cm 37.0 715.2 <.01 3.2 <.01 2.67 <.01 98.6 <.01 0.3 0.56

Panicle length, cm 12.2 3.9 0.07 4.7 <.01 2.32 <.01 30.7 <.01 7.2 <.01

Awn length, cm 4.1 0.0 0.91 6.6 <.01 1.27 0.09 2.3 0.13 1.6 0.21

Production§

Survival 0.71 11.2 <.01 6.5 <.01 3.01 <.01 54.9 <.01 11.0 <.01

Panicle, number 32.4 145.8 <.01 2.5 <.01 1.61 <.01 186.7 <.01 96.4 <.01

Biomass, g 12.8 230.2 <.01 3.0 <.01 2.66 <.01 17.4 <.01 14.3 <.01

Crown area, cm

2 62.8 96.5 <.01 2.0 <.01 1.67 <.01 213.0 <.01 13.7 <.01

†Heading was at complete emergence of a lead panicle from its sheath, blooming was the initial appearance of anthers, and

maturity when seed appeared mature in more than half the panicles.

‡Leaf length and width were measured on the last fully emerged leaf of a lead culm after heading; culm length, panicle length,

and awn length (twice geniculate measured in three sections and summed) were measured on a lead culm after fullextension.

§Panicles on each plant were counted after heading, harvest for dry weight was after maturity to within 4 cm of the soil, and

after harvest crown area was estimated as the product of two perpendicular crown diametermeasurements.

To simplify canonical correlation, traits were averaged over sites and years when correlations 

between specific traits were positive and significant (P < 0.01), indicating considerable 

correspondence for plants growing in different environments. This process reduced traits from a 

potential of 48 for all site-year combinations (12 traits times 4 environments) to 24 (Table 2).

These were used for canonical correlation with 21 ‘annual’ climate variables (Table 3) as defined 

by Wang et al. (2012). Since only the first canonical variate was significant (P < 0.001), 

explaining 44% of the variation, it was the only variate used in regression modeling with the 21 

climate variables.
18

Table 2. Plant traits used in canonical correlation of Thurber’s 

needlegrass growing at common gardens at Central Ferry (CF), WA and 

Reno (RE), NV in 2012 and 2013.

Heading† Panicle length RE12

Blooming Panicle length RE13

Maturity CF Awn length

Maturity RE Survival CF

Leaf length x width Survival RE

Leaf length to width CF 2012 Panicle number CF

Leaf length to width CF 2013 Panicle number RE

Leaf length to width RE 2012 Biomass CF

Leaf length to width RE 2013 Biomass RE

Culm length CF Crown area CF

Culm length RE Crown area RE 2012

Panicle length CF Crown area RE 2013

†See Table 1 for detailed descriptions of plants traits. Plant traits were averaged

over years and/or sites if correlations among of populations between year and

site combinations were positive and significant at P<0.01

Pearson’s linear correlations between climate variables and canonical variate 1 were often highly 

significant (P<0.01; Table 3) establishing a strong link between plant trait variation and climate 

across seed sources. The best regression model for canonical variate 1 and climates had 14 

climate variables and explained 96% of the variation (Table 4), an exceptionally high amount.

Regression R2 for other studies using multivariate plant traits with climateperennial 

bunchgrasses in the Intermountain West were substantially less. For Sandberg bluegrass, 

Johnson et al. (2015) reported regression model R2 values of 0.71 to 0.48 for canonical variates1 

and 2, respectively. For bluebunch wheatgrass, St. Clair et al. (2013), found regression R2 values 

of 0.51 and 0.28 for principal component (PC) 1 and PC 2, and Johnson et al. (2010) reported R2 

values between 0.40 and 0.46 for mountain brome PC 1 and PC 2. For Indian ricegrass, 

regression models with canonical variates and climate ranged from 0.16 to 0.43 (Johnson et al.

2012). Among conifers, Douglas fir is considered closely adapted to differing environments and 

regression models between canonical variates and climate resulted in R2 values of 0.50 and0.68 

(St. Clair et al. 2005). Horning et al. (2010) developed a regression model for oceanspray 

(Holodiscus discolor [Pursh] Maxim) from the first PC and climate with an R2 of 0.86,which 

approached our model for Thurber’s needlegrass. Thus the high R2 for the regression modelfor 

Thurber’s needlegrass (Table 5) indicated that canonical variate 1 was closely associated with 

climate variability across the Great Basin and suitable for seed zone development and mapping.

As in other genecology studies, Thurber’s needlegrass showed plant trait variation likely 

resulting from climate driven natural selection and evolution. The regression model (Table 4) 

will be used to develop seed zones and maps for Thurber’s needlegrass to guide the choice of 

germplasm for Great Basin restoration projects.
19

Table 3. Pearson correlation coefficient (r) between canonical variate 1 for plant 

traits and 21 climatic variables derived from each source population ofThurber’s 

needlegrass (n=60).

Climate variable† r P-value

Mean average temperature (MAT) 0.53 <.01

Mean warmest monthly temperature (MWMT) 0.58 <.01

Mean coldest monthly temperature (MCMT) 0.20 0.13

Continentality, MWMT-MCMT 0.43 <.01

Mean Annual Precipitation (MAP) -0.15 0.26

Mean summer precipitation (MSP) 0.04 0.78

Annual heat moisture index, ( MAT+10)/(MAP/1000) 0.37 <.01

Summer heat moisture index ((MWMT)/(MSP/1000)) 0.30 0.02

Chilling degree days (degree days <0°C) -0.27 0.04

Growing degree days (degree days >5°C 0.58 <.01

Heating degree days (degree days<18°C) -0.50 <.01

Cooling degree days (degree days>18°C) 0.53 <.01

Number of frost free days (NFFD) 0.35 <.01

Frost free period (FFP) -0.41 <.01

Day of year when FFP begins (bFFP) 0.38 <.01

Day of year when FFP ends(eFFP) 0.42 <.01

Precipitation as snow (PAS) -0.32 <.01

Extreme minimum temperature over 30 years (EMT) 0.14 0.29

Extreme maximum temperature over 30 years (EXT) 0.48 <.01

Hargreaves reference temperature (Eref) 0.58 <.01

Hargreaves climatic moisture index (CMD) 0.45 <.01

† See Wang et al. 2012 for additional climate variable definitions and details

Table 4. Coefficient of determination (R2) and the regression equation developed between canonical variate 

1 and climatic variables for Thurber’s needlegrass.

Trait R2 Regression equation†

Canonical variate 

1

0.96 27.79+5.957(MAT)- 0.0132(MAP) +0.0387(MSP) +0.0118(SHM)

+0.0432(DD<0)-0.0204(DD>18)

-0.2356(NFFD)-0.0568(bFFP) +0.0623(FFP) +0.0212(PAS)

+0.7153(EMT)-0.7993(EXT) +0.0235(Eref)-0.0193(CMD)

†Annual climatic variables as given in Table 3 and by Wang et al. (2012)
20

Basin wildrye

Collections of Basin wildrye were made in the Columbia 

Plateau in Washington State where the octoploid form is 

common, and in the Great Basin where the tetraploid and 

octoploid can be found. Ploidy for all collection locations 

was determined as described by Jakubowski et al. (2011). 

The analysis of field collected sources revealed 58 

octoploids and 56 tetraploids.

Common gardens of Basin wildrye were established at 

Pullman and Central Ferry, WA, sites in 2010 with plants 

from 114 collection locations that were randomized in

complete blocks with six replications. Data collection for

phenology, production, and morphology traits was 

completed 2011, 2012, and 2013. Initial data analyses 

indicate considerable plant trait variation, as expected. 

There were also differences associated with ploidy, so 

ploidy will be considered in climate adaption models. Seed 

zones for Basin wildrye will be developed in 2015.

Bottlebrush squirreltail

Common gardens of Bottlebrush squirrel-tail were 

established in at Central Ferry WA, Powell Butte, OR, and 

Reno, NV, and data collection at Central Ferry was 

completed for numerous phenology, production, and 

morphological traits. Data has been sent to Francis 

Kilkenny at the Rocky Mountain Research Station for 

consolidation with other sites, data analysis, and 

publication.

Sulphur-flower buckwheat

Collection of Sulphur-flower buckwheat was completed by

Figure 2. Distribution data for Seeds of

Success accessions within the National Plant

Germplasm System from 2006 through 2014.

the Western Regional Plant Introduction Station (WRPIS) in 2012 resulting in 21 accessions. 

There has also been ongoing collection through the BLM Seeds of Success (SOS) program 

resulting in 48 accessions. Seeds from the WRPIS and SOS collections were used to establish 

plants in the greenhouse in 2014. In the fall of 2014, these were transplanted to a common 

garden at the WRPIS farm at Pullman, WA. Taxonomy will be verified and data collected on 

phenology, production, and morphology plant traits in 2015 and 2016.

Seeds of Success activities

Seeds of Success (SOS) and the National Plant Germplasm System (NPGS) have partnered to 

collect, distribute, and evaluate key native plant materials needed for conservation and 

restoration. Accession and distribution numbers continue to grow. In 2014, 1,403 new native 

plant accessions were added to the SOS-NPGS collection, which now totals 10,515 accessions. 

Since 2006, nearly 4,500 seed distributions have been made for research projects including 

federal, state, U. S. commerce and private cooperators (Figure 2).

Cumulative SOS

4500 distrutions

4000

3500 Packets

3000 Taxa

2500 Orders

2000

1500

1000

500

0

06 07 08 09 10 11 12 13 14

SOS distributions by year

1200

1000

800

600

400

200

0

Packets

Taxa 

Orders

06 07 08 09 10 11 12 13 14
21

Publications

Johnson, R. C.; Horning, M. E.; Espeland, E.; Vance-Borland, K. 2015. Genetic variation and 

climate interactions of Poa secunda (Sandberg bluegrass) from the Intermountain Western 

United States. Evolutionary Applications 8: 172-184.

Dugan F. M.; Cashman, M. J.; Johnson, R. C.; Wang, M.; Xianming C. 2014. Differential 

resistance to stripe rust (Puccinia striiformis) in collections of basin wild rye (Leymus cinereus). 

Plant Health Progress 15: 34-38.

Johnson, R. C. 2014. Genecology and seed zone research for native forbs and grasses of the 

interior West. In: Proceedings Society of Ecological Restoration Regional Conference, 

Redmond, Northwest and Great Basin Chapter; 2014 October 6-10; Redmond, OR.

Presentations

Conservation, Adaptation, and Seed Zones for Key Great Basin Species for the Great Basin 

Native Plant Project Annual Meeting; 2014 March 17; Boise, ID.

Management Applications and Seed Production Guidelines

 Maps visualizing the interaction of genetic variability and climate for restoration species 

allow informed decisions regarding the suitability of genetic resources for restoration in 

varying Great Basin and Southwestern environments.

 The recommended seed zone boundaries may be modified based on management 

resources and land manger experience without changing their basic form or the 

relationship between genetic variation and climate.

 We recommend utilization of multiple populations of a given species within each seed 

zone to promote the biodiversity needed for sustainable restoration and genetic 

conservation.

 Collections representing each seed zone should be released, grown, and used for ongoing 

restoration projects.

Products

 Seed zones are for key Great Basin restoration species to ensure plant materials used for 

restoration are adapted, suitable, and diverse.

 Collection and conservation of native plant germplasm though cooperation between the 

National Plant Germplasm System (NPGS), the Western Regional Plant Introduction 

Station, Pullman, WA, the U. S. Forest Service, and the Seeds of Success (SOS) program 

under BLM. This ensures availability of native plant genetic resources that otherwise 

may be lost as a result of disturbances such as fire, invasive weeds, and climate change.
22

References

Horning, M. E.; McGovern, T. R.; Darris, D. C.; Mandel, N. L.; Johnson, R. 2010. Genecology 

of Holodiscus discolor (Rosaceae) in the Pacific Northwest. U. S. A. Restoration Ecology 18: 

235-243.

Jakubowski, A. R.; Jackson, R. D.; Johnson, R. C.; Hu, J.; Casler, M. D. 2011. Genetic diversity 

and population structure of Eurasian 1 populations of reed canarygrass: cytotypes, cultivars, and 

interspecific hybrids. Crop and Pasture Sciences 62: 982-991.

Johnson, R. C.; Erickson, V. J.; Mandel, N. L.; Bradley St. Clair, J. B.; Vance-Borland, K. W. 

2010. Mapping genetic variation and seed zones for Bromus carinatus in the Blue Mountains of 

Eastern Oregon, U. S. A. Botany-Botanique 88: 725-736.

Johnson, R. C.; Cashman, M. J.; Vance-Borland, K. 2012. Genecology and seed zones for Indian 

Ricegrass across the Southwest USA. Rangeland Ecology and Management 65: 523-532.

Johnson, R. C.; Hellier, B. C.; Vance-Borland, K. W. 2013. Genecology and seed zones for 

tapertip onion in the US Great Basin. Botany 91: 686-694.

Johnson, R. C.; Horning, M. E.; Espeland, E.; Vance-Borland, K. 2015. Genetic variation and 

climate interactions of Poa secunda (Sandberg bluegrass) from the Intermountain Western 

United States. Evolutionary Applications 8: 172-184.

Leimu, R.; Fischer, M. 2008. A meta-analysis of local adaptation in plants. PLoS One 3: e4010.

St. Clair, J. B.; Mandel, N. L.; Vance-Borland, K. W. 2005. Genecology of Douglas fir in 

western Oregon and Washington. Annals of Botany 96: 1199–1214.

St. Clair, J. B.; Kilkenny, F. F.; Johnson, R. C.; Shaw, N. L.; Weaver, G. 2013. Genetic variation 

in adaptive traits and seed transfer zones for Pseudoroegneria spicata (bluebunch wheatgrass) in 

the northwestern United States. Evolutionary Applications 6: 933-948.

Wang, T.; Hamann, A.; Spittlehouse, D. L.; Murdock, T. Q. 2012. ClimateWNA–Highresolution spatial climate data for western North America. Journal of Applied Meteorology and 

Climatology 51: 16–29.
23

Project Title Ecological Genetics of Big Sagebrush (Artemisia

tridentata): Genetic Structure and Climate-based Seed 

Zone Mapping

Project Agreement No. 13-IA-11221632-161

Principal Investigators and Contact Information

Bryce Richardson, Research Geneticist 

USDA Forest Service

Rocky Mountain Research Station 

Shrub Sciences Laboratory

735 N. 500 E.

Provo, UT 84606

801.356.5112

brichardson02@fs.fed.us

Nancy Shaw, Research Botanist

USDA Forest Service

Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4360

nancylshaw@fs.fed.us

Matthew Germino, Supervisory Research Ecologist

U. S. Geological Survey

Forest and Rangeland Ecosystem Science Center 

Boise, Idaho 83702

208.426.3353

mgermino@usgs.gov

Project Description

Plant species that occupy large geographic distributions contend with highly varied climates. 

Such climatic differences frequently require genetic adaptation. Failure to understand how these 

species are adapted to climate will impede restoration. In this study we assess the growth and 

seed yield of big sagebrush (Artemisia tridentata subspecies tridentata, vaseyana and 

wyomingensis) from two common gardens over four years. Using linear-mixed models, growth 

and seed yield variation is associated with source-location derived climate variables 

(temperature, precipitation and their interactions). Overall, continentality strongly affects climate 

adaptation in big sagebrush. We discuss the genetic patterns and development of seed transfer 

zones based on these data.
24

Objectives

1) Establish common gardens from collected seed sources across the range of big sagebrush.

2) Determine climatic factors important to adaptation within and among big sagebrush 

ecological races.

3) Develop climate based seed transfer zone maps across the range of big sagebrush.

Methods

Common Garden Establishment and Measurements

Collection of seed and plant tissue began in autumn of 2009. A total of 93 seed sources were 

collected, largely by collaborators, in 11 western states. In January 2010, seeds were planted in 

greenhouse containers. Up to ten families from each of 55 seed sources were outplanted at each 

of the common gardens (Table 1). Outplanting of seedlings occurred in May and June of 2010. 

First-year measurements were conducted in October and November of 2010. Plants were 

supplemented with water until August 2010 and mortality was minimal for the first year at <2%. 

No supplemental water was added in subsequent years.

Plant growth was determined by height and crown area. Plant heights were measured from the 

ground to the highest, living, non-flowering stem. Crown areas were calculated from overhead 

digital photos. Photos were analyzed by cropping green leaves from a white background placed 

under the plant using the color threshold feature in ImageJ ver 1.45. Pixel size of the cropped 

area was scaled to a known standard on each photo. Measurements were taken in October 2010 

and in May 2011 and 2012. Heights and crown areas were used to estimate volume based on an 

ellipsoid volume equation from measurements that occurred within 2 weeks of each other.

Seed yield was estimated for two plants per population in both gardens during 2012 and 2013 

growing seasons. Plants were randomly selected from each population. In August, two flowering 

stems of approximately the same size were fitted with 13x45cm bags constructed from mosquito 

netting. The bags were tied at the base of the stems with string. The remaining unbagged 

flowering stems were counted. The bags were then harvested in early December afterseed 

ripening. Contents of each bag were cleaned to remove any chaff and the seed was weighed. An 

estimate of total seed yield per plant was calculated by multiplying the seed yield from each bag 

and the total number of inflorescence stems.

Table 1. Location information of three big sagebrush common garden sites.

Site name Latitude Longitude Elevation (m)

Orchard, Idaho 43.328 -116.003 976

Ephraim, Utah 39.369 -111.580 1686

Majors Flat, Utah 39.337 -111.521 2088

Common Garden Analyses

In this report we focus on modeling climate associated with growth and seed yield of subspecies 

tridentata and wyomingensis from Majors Flat and Orchard gardens. Seed and growth were the 

response variables for linear-mixed models. The predictors in these models were divided into 

fixed and random effects. Fixed effects included original seed-source climate variables and 

random effects included a nested hierarchy of garden, year (seed yield only), population and 

family. The original seed-source climate variables were estimated from a spline model based
25

upon weather station data recorded between 1961-1990

(http://forest.moscowfsl.wsu.edu/climate/). Mapping of the model outputs was completed using 

the slope(s) and intercept from the fixed (climate) effects and individual cell values, 0 00833o (~ 

1 km2) from the relevant climate-surface variables. These grids were mapped with the predicted 

climatic niche of a subspecies wyomingensis bioclimate model (Still and Richardson 2015).

Results and Discussion

Among the climate (fixed) effects, summer dryness index (SDI) was an important predictor for 

both growth and seed yield (p < 0.0001 and p < 0.05, respectively). In addition, winter 

precipitation was important for seed yield (p < 0.001). The predicted values from the growth and 

seed yield models were highly and positively correlated (r = 0.72, p < 0.0001). Among random 

effects, subspecies-ploidy groups were significant for both growth and seed yield (p < 0.0001). 

Population variance was significant for growth (p < 0.0001) and family variance wassignificant 

for seed yield (p < 0.0001). It is important to note that subspecies-ploidy, population and family 

variances were nested within garden. However, garden alone was not a significant random effect 

in either model.

To illustrate the model prediction, fixed (climate) effects of seed yield and growth are projected 

within the predicted bioclimate niche of subspecies wyomingensis. For seed yield, the mapped 

patterns predict that tridentata and wyomingensis populations from areas with greater winter 

precipitation and drier and warmer summers have higher fecundity at the common gardensthan 

populations that receive more summer precipitation and cooler summers (Figure 1). Differences 

in seed yield among populations appear especially true for the Orchard garden, which 

experiences relatively warmer temperatures and precipitation that principally falls in winter 

compared to regions further east. For example at the Orchard garden, seed yield ranged from an 

average of 0.1g from populations in Montana and southern Utah to 41g from the local Orchard, 

Idaho source.

As mentioned above, growth followed a similar pattern as seed yield. Plants from warmer, drier 

summer climates had greater growth than plants originating from areas further east that receive 

more frequent summer precipitation and cooler temperatures (Figure 2). These differences in 

growth were most pronounced at the Orchard garden compared to Majors Flat. It is expected that 

if a garden were established in a region that received greater summer precipitation (e.g., 

Montana) the reciprocal would be true. Plants from areas acclimated to receive greatersummer 

precipitation would have greater growth.

Future plans

Data collection for the project is complete and final data analyses and manuscripts will be 

submitted to peer-reviewed journals. Confidence intervals will need to be calculated to break the 

continuous variation (Figure 1 and Figure 2) into climatypes and seed, growth and survivorship 

(not shown) models will be combined to construct seed transfer zones. The association between 

trait responses and physiology remains.
26

Management Applications

The highlighted data and models are the foundation for development of seed transfer zonesfor 

big sagebrush. Synthesis of these models into seed zones is underway. Published seed transfer 

guidelines will be developed into a web-based tool for land managers.

-125° -120° -115° -110° -105° -100°

50°

45°

40°

35°

-120° -115° -110° -105° -100°

Figure 1. Mapped linear mixed model of fixed effects (climate) of seed yield (g) for 

basin big sagebrush (Artemisia tridentata ssp. tridentata) and Wyoming big sagebrush 

(Artemisia tridentata ssp. wyomingensis). Colors corresponded to the predicted values 

of the model displayed as log (1+seed yield). For example, values of 4.09 and 0.68 

equate to 58.7 and 0.9g of seed per plant. The values are mapped within the predicted 

climate niche of Wyoming big sagebrush (Still and Richardson 2015).

log(1+ seed yield)

Value

High : 4.09204

Low : 0.684407

45°

40°

35°

30° 0 80 160 320 Km
27

-125° -120° -115° -110° -105° -100°

50°

45°

40°

35°

-120° -115° -110° -105° -100°

Figure 2. Mapped linear mixed model of fixed effects (climate) of growth (m3) for basin 

big sagebrush (Artemisia tridentata ssp. tridentata) and Wyoming big sagebrush 

(Artemisia tridentata ssp. wyomingensis). Colors correspond to the predicted values of 

the model displayed as log (1+seed yield). For example, values of 0.89 and 0.44 equate to

1.43 and 0.55m2 of growth. The values are mapped within the predicted climate nicheof 

Wyoming big sagebrush (Still and Richardson 2015).

45°

40°

35°

log(1 + growth)

Value

High : 0.899633

Low : 0.444504

30° 0 80 160 320 Km
28

Presentations

Richardson, B. Development of tools and technology to improve the success and planning of 

restoration of big sagebrush ecosystems. Great Basin Landscape Conservation Cooperative 

Webinar; 2014, September 16.

Richardson, Bryce R.; Shaw, Nancy L.; Germino, Matt; Ortiz, Hector; Carlson, Stephanie; 

Sanderson, Stewart; Still, Shannon. Development of tools and technology to improve the success 

and planning of restoration of big sagebrush ecosystems. Great Basin Consortium Conference; 

2013 December 9-10; Reno, NV. Abstract and Poster.

Richardson, B. A.; Shaw, N. L.; Germino, M. J.; Still, S. M. Comparisons of contemporary and 

future predictions of the climate niche and ecological genetics of big sagebrush and blackbrush. 

Society for Ecological Restoration Regional Conference, Northwest and Great Basin Chapters; 

2014 October 6-10; Redmond, OR.

Richardson, B.; Shaw, N. Germino, M.; Still, S. Relationships between common garden

responses and climate in big sagebrush (Artemisia tridentata). Great Basin Native Plant Annual 

Meeting; 2014 March 17-18; Boise, ID.

Additional Products

 Leveraged funding: National Fire Plan 2013-2016 ($72,000 per year).

 Leveraged funding: Great Basin LCC 2013-2015 ($35,000).

References

Still, S. M.; Richardson, B. A. 2015. Projections of Contemporary and Future Climate Niche for 

Wyoming Big Sagebrush (Artemisia tridentata subsp. wyomingensis): A Guide for Restoration. 

Natural Areas Journal 35: 30–43.
29

Project Title Factors Affecting Initial Establishment of Big Sagebrush

Outplants: Land Treatments, Seed Sources, and Climate 

Effects

Project agreement No. 14-IA-11221632-012 and 11-IA-11221632-098

Principal Investigators and Contact Information

Matthew J. Germino, Supervisory Research Ecologist

U. S. Geological Survey

Forest and Rangeland Ecosystem Science Center 

970 Lusk St.

Boise, ID 83706

208.426.3353, FAX 208.426.5210

mgermino@usgs.gov

Martha Brabec, Biological Sciences Technician

U. S. Geological Survey

Forest and Rangeland Ecosystem Science Center 

970 Lusk St.

Boise, ID 83706

208.426.5207, FAX 208.426.5210

mbrabec@usgs.gov

Bryce Richardson, Research Geneticist

USDA Forest Service, Rocky Mountain Experiment Station 

735 N. 500 E.

Provo, UT 84606

801.356.5112

brichardson02@fs.fed.us

This information is preliminary and is subject to revision. It is being provided to meet the need 

for timely best science. The information is provided on the condition that neither the U. S. 

Geological Survey nor the U. S. Government may be held liable for any damages resulting from 

the authorized or unauthorized use of this information.

Project Description

Increased wildfire activity poses a major challenge for big sagebrush (Artemisia tridentata) and 

its dependent species due to a lack of fire adaptation in big sagebrush. Big sagebrush seeding and 

planting are becoming increasingly common, particularly in light of conservation concerns of 

sage grouse (Centrocercus urophasianus), but success has been mixed. Climate and weather 

variability, appropriateness of seeds, and site conditions relating to management are all factors
30

that may help explain success of sagebrush restoration efforts. Consideration of these factors 

may improve seeding or planting success.

Big sagebrush has high intraspecific diversity, and selection of appropriate seed source is 

increasingly a concern in designing and implementing big sagebrush restoration treatments. 

Diversity in big sagebrush results from subspecies, cytotypes, and population-level variability. 

Seedling survival is recognized as a primary bottleneck for sagebrush establishment, and 

information on climate adaptation and ecological responses at this stage are a priority need.

Big sagebrush seeding or planting treatments are often accompanied by treatments that reduce 

wildfire fuel loads and invasive annuals or increase native herbs through mowing, herbicide 

applications, or rangeland drill seeding. Neighboring herbs can have strong effects on big 

sagebrush seedlings, which can relate to competition for water and other soil resources.

Objectives

1. Assess the influence of management treatments of herbs on seedlings of differentseed 

sources of big sagebrush.

2. Compare growth, survival, and underlying ecophysiological mechanisms and thresholds 

among seedlings of the different subspecies, cytotypes, and populations of big sagebrush.

Methods

Objective 1: Effect of Management Treatments

We planted seedlings in 48 1-ha experimental plots in the Birds of Prey National Conservation 

Area (BOP NCA) in Idaho, on an A. tridentata wyomingensis site that burned in 1998 and had 

become dominated by cheatgrass and Sandberg’s bluegrass (Poa secunda). Experimental plots 

were placed in three replicate blocks using a full-factorial, completely randomized combination 

of four management practices (no treatment, 4 oz/acre of Impazic herbicide, mowing, and 

mowing plus herbicide) and two seeding treatments (seeded and unseeded). All treatments were 

implemented in fall 2012. A native seed mix containing grasses, forbs, and shrubs was applied 

using a Truax minimum-till seed drill.

Seven-month old sagebrush seedlings of cone-tainer stock (shoots were <~4 cm tall) were

outplanted in spring 2013. Seedlings of four subspecies of A. tridentata ssp. tridentata were 

bare-root transplanted into small holes and outplanted into all 48 plots (nine seedlings per 

subspecies per plot). Seed sources included dipoids local to the study site (IDT2) or from a 

relatively dry site in California (CAT-2) or a warm and dry site in Oregon (ORT2, known to be 

fast-growing), and a tetraploid population from a relatively dry site in New Mexico (NMT2).

Survival of outplants was measured quarterly from April 2013-May 2014. For Objective one 

(and two), differences in survival and significance of factors related to survivorship were 

determined using Cox’s hazard of death analyses and general estimating equations along with 

conditional logistic regression (Brabec 2014).
31

Objective 2: Variation Among Seedlings of Different Subspecies, 

Cytotypes, and Populations

Eleven different seed sources were utilized to assess how initial establishment and factors 

affecting it vary among subspecies, cytotypes, and population of big sagebrush (Table 1) and 

seedlings were reared as in Objective 1. Plots were established in five separate sites (statistical 

blocks) that burned in the summer of 2012 at BOP NCA, south of Boise, Idaho, in a randomized 

complete block design. Each block was fenced to prevent cattle disturbance and grazing on the 

plots. Seedlings were outplanted in November 2012 and average seedling heights ranged from 

2.25-3.83cm, with ten seedlings per plot and 10cm spacing between seedlings. Height and 

survival of sagebrush seedlings were recorded in monthly increments to May 2014. Soil moisture 

and temperatures in the seedling’s depth or height of roots or shoots (respectively) were

measured in each plot and are reported in Brabec (2014).

Table 1. Description of big sagebrush seed sources utilized in this study. 

Provenance State Location Subspecies Climate-of-origin Ploidy

IDT-2 ID Orchard (Local) Tridentata Local 2N

BOP-W ID Birds of Prey Wyomingensis Local 4N

IDV-2 ID (Nearest Mtn) Vaseyana Cool/Wet 2N

NMT-2 NM San Luis Mesa Tridentata Dry 4N

UTT-1 UT Canyon B. Tridentata Cool/Wet 2N

IDW-2 ID Orchard (Local) Wyomingensis Local 4N

ORT-2 OR Echo Tridentata Warm/Dry 2N

MTW-3 MT Montana Wyomingensis Cool/Wet 4N

ORV-1 OR Lookout Mountain Vaseyana Cool/Wet 4N

IDW-3 ID Sommer Camp Wyomingensis Dry 4N

IDV-3 ID Stanley Vaseyana Cool/Wet 2N

Freezing points were determined from exothermic heat pulses released when leaves reached their 

freezing points during controlled cooling experiments in the lab. To assess freezing tolerance, 

we measured Fv/Fm50, the temperature causing 50% loss of chlorophyll a fluorescence. ANOVA 

was used to assess physiological differences among subspecies, cytotype, and climate-of-origin 

to freezing. Freezing resistance mechanisms (avoidance or tolerance) were determined by 

assessment of the difference between Fv/Fm50 and freezing point.

Results and Discussion

We observed high mortality during the first year of the study, which corresponded to unusually 

cold and dry conditions.
32

Objective 1: Effect of ManagementTtreatments 

Management treatments, drill seeding, and big sagebrush 

population seed source all influenced seedling survival 

times (χ2

=69.8, p<0.001). Mortality risks increased 2-to 

15-fold on drill-seeded compared to unseeded plots that 

also had treatment combinations resulting in reduced 

herbaceous cover (i.e., mowing, mowing+herbicide; 

Figure 1), although substantive emergence of seeded 

species was not readily detected in the treatments. Mean 

cheatgrass cover was 25.21±7.9% on seeded plots and 

18.02±6.2% on non-seeded plots, and seeded plots also 

had less mean bare soil cover (19.84±2.7%) compared to 

non-seeded plots (25.03±2.6%).

A hypothetical explanation for our results is that 

establishment of sagebrush outplants was negatively 

affected by increased competition from invasive 

herbaceous species, particularly in drill-seeded plots. 

Reduced spacing between neighboring herbs such as

cheatgrass and sagebrush seedlings on the drill-seeded 

plots could have exacerbated competition. Conversely, the 

reduced intensity of plant interactions in communities on 

non-seeded plots may have increased water and nutrient 

availability for seedlings and thus survival.

Objective 2: Variation Among Seedlings of Different 

Subspecies, Cytotypes, and Populations

As predicted, seedling survival over the entire duration of 

the experiment was less for A. t. vaseyana relative to both

A. t. tridentata and ssp. wyomingensis. Survival of 

tetraploid groups was consistently greater than diploids, for 

comparisons within subspecies; χ2=6.8, p=0.009 for A. t.

vaseyana and χ2

=4.4, p=0.036 for A. t. tridentata. Diploid A. t. 

tridentata had the greatest average heights of all populations 

and their survivorship was at 12% and 16% (6.13 and 5.76 cm

Figure 1. Differences in 

survivorship among 8 treatments in 

April, May, and June 2013 (from 

Brabec et al., in review).

respectively). Survival of other populations was 20-30%, and heights ranged 3.0-4.5 cm. 

Survival of A. t. wyomingensis was 24%, and height averaged 4.65 cm.

Mortality patterns over all seedlings were related more to minimum temperatures of the soil 

surface than to growing degree days >10°C or volumetric water content (from conditional 

logistic regression; Information criterion coefficient =1378 compared to 1462 or 1466, 

respectively). The sensitivity of survival to minimum temperatures did not relate to climate-oforigin of populations, but statistical inference was complicated by high mortality among blocks.
33

Freezing points of A. t. vaseyana populations were 2-3°C significantly warmer than for A. t. 

wyomingensis populations (Figure 2). A. t. tridentata froze at -10.51 ± 0.71°C. The range of 

Fv/Fm50 among subspecies x cytotypes was much smaller than that observed for freezing point 

(Figure 2). Among subspecies, A. t.

wyomingensis had greater freezing resistance 

than both A. t. vaseyana and ssp. tridentata by 

approximately 1.1°C (p = 0.0017). There were 

no differences in Fv/Fm50 between cytotypes 

within each subspecies, but A. t. wyomingensis 

had 1.4°C greater freezing resistance than 

diploid A. t. vaseyana. The difference (in °C) 

between Fv/Fm50 and freezing point (Fv/Fm50

─freezing point) revealed variable freezing 

response strategies among subspecies and 

cytotypes (Figure 2). Tetraploid A. t. tridentata 

had a significantly higher Fv/Fm50 ─freezing 

point than tetraploid A. t. vaseyana indicating. 

Fv/Fm50 ─freezing point was not associated with 

climate-of-origin for any subspecies.

Cytotype appears to be an important factor in 

explaining functional diversity, possibly even 

as important as subspecies itself for seedling 

establishment and survival. Using the criteria 

for freezing avoidance versus tolerance, our 

data suggest big sagebrush’s freezing 

resistance strategy is classified as freezing 

tolerance, except for tetraploid A. t. tridentata,

an avoider (Figure 2). “Avoiders” had the most 

mortality in March, whereas freezing “tolerators” 

experienced consistent morality due to minimum 

temperatures from April to June. A putative 

investment in freezing tolerance at the expense of 

other growth (competitive) traits may limit the 

ability of seedlings to compete with faster growing 

species when freezing events are infrequent, 

resulting in mortality.

Figure 2. The relationship of

temperature causing 50% loss of 

Fv/Fm (Fv/Fm50) tofreezing point (A, 

R2 all three regressionlines

<0.05) in contrast to the relationship 

of the difference between Fv/Fm50 and 

freezing point (“Fv/Fm50 ─ Freezing 

Point °C”) to freezing point by 

subspecies and cytotype (R2

=0.776)

(from Brabec et al., in review).

In spite of its freezing tolerance strategy, A. t. vaseyana had the greatest vulnerability to 

minimum temperatures and thus least capacity to cope with freezing. The relatively higher 

elevations of A. t. vaseyana habitat may on one hand be cooler, but on the other hand is

more likely to be insulated from temperature extremes by snow cover.
34

Management Applications and/or Seed Production Guidelines

Interactions among species in seed mixes are rarely considered, except where aggressive nonnative species (e.g. crested wheatgrass or forage kochia) are part of the seed mix. Our 

preliminary data suggest that treatments resulting in temporary reduction of herb cover may 

increase success in situations where establishment of big sagebrush is a priority. Our 

preliminary data indicate that seedling mortality, growth, and response to freezing differ at the 

subspecies, cytotype, and population level. Cytotype may be as important as subspecies in 

explaining adaptation and functional diversity in big sagebrush. Consideration of this functional 

diversity may improve sagebrush planting success.

Presentations

Germino, M. J. Evolution and conservation of sagebrush. Ecology and Evolution Class, College 

of Idaho; 2014 October 29; Caldwell, ID.

Germino, M. J. 2014. Sagebrush responses to shifting climate and fire disturbances: basic 

insights for restoration. Sage Steppe Partner Forum Webinar Series; 2013 January 21; Boise, ID.

Brabec, M. M.; Germino, M. J.; Richardson, B. A.; Vander Veen, J.; Lazarus, B. 2014. Big 

sagebrush changing climate context; the effects of genotype and climate variability on success of 

sagebrush seedlings. Ecological Society of America Annual Meeting, Sacramento CA; Aug 10-

15. Poster.

The loss of big sagebrush (Artemisia tridentata) due to wildfire has motivated effortsto 

restore it through seeding and planting. Restoration success has been mixed, despite large 

investment. Big sagebrush displays high intraspecific diversity, and appropriate seed source is an 

increasing management concern in post-fire restoration. Currently, there are unmet data needsfor 

establishing seed transfer zones for big sagebrush under future climate regimes and a major 

pending question is whether seeds sources from warmer and drier sites will be better suited than 

local seed sources. Our primary objectives in this study were to assess climate responsiveness of 

seedlings of different provenances on burned areas and relate ecophysiological-climate 

adaptation in seedlings to that of established plants from a common garden study. Experimental 

passive warming chambers paired with unwarmed control plots were installed at Birds of Prey 

National Conservation Area on five sites burned in 2012. We evaluated eleven different seed 

sources of big sagebrush from all three subspecies, dissimilar climates of origin, and different 

ploidy levels to assess how genotype affects initial establishment of sagebrush. A suite of 

dependent variables were measured; including survival, growth, chlorophyll fluorescence, 

isotopic assessment of water-use efficiency, and differential thermal analysis. Preliminary results 

suggest that experimental warming enhanced overall seedling survival revealing the sensitivity of

sagebrush to temperature (particularly daily minima). Among the three subspecies ofsagebrush, 

basin big sagebrush had the lowest mortality under warming, which may relate to 

access to deeper soil water resources (p=0.001). Overall, we did not observe consistent strong 

support for local adaptation. Local genotypes exhibited greater stress and photoinhibition after 

freezing and light treatments than distant genotypes (p=0.01). Isotopic assessment of water-use 

efficiency of distant vs. local seedlings revealed that seedlings from warmer and drier climates 

have greater water use efficiency under current climate conditions (p=0.04). During initial 

establishment, there is an emerging pattern that implies a tradeoff between growth and survival. 

Genotypes with most overall growth also had the highest overall mortality, and this trend 

contrasts with patterns observed in more mature sagebrush. These findings indicate the
35

importance of drawing inference on ecophysiological adaptation from multiple demographic 

stages while establishing seed transfer zones for big sagebrush restoration.

Germino, M. J. 2014. Sagebrush responses to shifting climate and fire disturbances: basic

insights for restoration. Society for Ecological Restoration Regional Conference of Northwest 

and Great Basin Chapters, 2014 October 6-10; Redmond, OR.

The Birds of Prey National Conservation Area (BOPNCA) has experienced considerable 

loss of big sagebrush as fire and invasive grasses have increased in recent decades, similar to 

many other regions in the western United States. Management efforts aimed at promoting 

recovery of sagebrush and wildlife habitat are faced with climate, weather, and other landscape 

challenges. Our research program is evaluating climate responses of adult and seedling 

sagebrush, and the effect of management options for post-fire seeding on recovery. To test the 

hypothesis that warming is hindering sagebrush persistence or post-fire establishment, and to 

evaluate genetic variability in responses, we established experimental warming and control plots 

on five new wildfire sites and on nearby unburned sites that still had sagebrush. Seedlings of a 

variety of populations of the three primary subspecies of big sagebrush were planted onto the 

burned plots. Preliminary data suggest little effect of warming on adult big sagebrush, which 

contrasts with positive effects observed elsewhere at higher elevations. Also in contrast, warming 

affected seedling survival and growth. Considerable variation was evident among subspecies and 

populations, indicating importance of selecting appropriate seed sources in restoration or 

rehabilitation seedings. In an associated pilot study, we attempted to evaluate how choice ofseed 

source has related to success of sagebrush recovery in >20 post-fire seedings in and around the 

BOPNCA, in light of weather, climate, and many other factors affecting establishment. In

contrast to the climate experiment, the pilot study did not reveal a strong effect of sagebrush seed 

source on seeding outcomes, in spite of a tendency for non-target subspecies from higher 

elevation climates to have been seeded. To provide a stronger test of how choice of sagebrush 

seed has affected seeding outcomes, we are expanding the analysis to include many more

seeding projects distributed throughout the Great Basin and elsewhere.

Brabec, M.; Germino, M. J.; Richardson, B. A. 2014. Big sagebrush in a changing climate 

context: effects of cytotype and climate variability on success of sagebrush seedlings. Society of 

Ecological Restoration Regional Conference of Northwest and Great Basin Chapters; 2014 

October 6-10; Redmond, OR.

Big sagebrush (Artemisia tridentata) is a genetically diverse species that shows high diversity 

in its response to climate. Three subspecies are widely recognized; mountain, basin, and 

Wyoming. Wyoming sagebrush is universally tetraploid and mountain and basin subspecies are 

often diploid, but tetraploid populations can occur. Data suggests that cytotype is as important a 

factor in explaining functional diversity as subspecies, and polyploidy and inter- and intraspecific 

hybridization are likely to be important mechanisms in big sagebrush adaptation and landscape

dominance. The loss of big sagebrush due to wildfire has motivated efforts to restore it, 

and appropriate seed source is an increasing management concern in post-fire restoration. A 

number of sagebrush common-garden studies have been conducted in the Great Basin and have 

revealed key ways that adult sagebrush differ by subspecies and cytotype, particularly in climate 

response. However, little is known about climate responses of seedlings during the initial 

establishment phase. To assess differences in climate responsiveness among sagebrush 

seedlings, we evaluated eleven different seed sources of big sagebrush from all three subspecies,
36

dissimilar climates of origin, and different ploidy levels at Birds of Prey National Conservation 

Area. A suite of dependent variables were measured including survival, growth, chlorophyll 

fluorescence, isotopic assessment of water-use efficiency, and freezing resistance and tolerance. 

Preliminary results suggest that freezing tolerance and resistance differ among cytotypes and 

may be an important factor in seedling survival. Exposure to cold temperatures in May resulted 

in higher probabilities of mortality in diploid basin big sagebrush and Wyoming big sagebrush. 

Tetraploid basin and mountain populations demonstrated higher survival rates relative to their 

diploid counterparts, providing further evidence ploidy level influences seedling stress responses. 

These findings indicate the importance of understanding climate responses of different 

subspecies and cytotypes when establishing seed zones for big sagebrush restoration.

Publications

Brabec, M.; Germino, M. J.; Richardson, B. [Submitted February 2015]. Intraspecific variation 

in big sagebrush seedling responses to post-fire weather indicate importance of minimum 

temperatures to establishment. Journal of Applied Ecology. (Paper submitted to the journal, but 

not yet accepted).

Brabec, M.; Germino, M. J. [Submitted February 2015]. Experimental warming supports 

importance of minimum temperature to post-fire establishment. Ecosphere. (Paper submitted to 

the journal, but not yet accepted).

Brabec, M.; Germino, M.; Shinneman, D.; Pilliod, D. S.; McIlroy, S.; Arkle, R. S. [Submitted 

February 2015]. Response of young sagebrush seedlings to management treatments of herbs.

Rangeland Ecology and Management. (Paper submitted to the journal, but not yet accepted).

Brabec, Martha. 2014. Big sagebrush in a shifting climate context: assessment of seedling 

responses to climate. Boise, ID: Boise State University. 95 p. Thesis.

Additional Products

We procured additional, leveraged funding for the management-treatment aspects of this project 

from the U. S. Fish and Wildlife Service contributions to the Great Basin Landscape 

Conservation Cooperative.

This research was made possible by cooperative efforts from D. Shinneman, R. Arkle, D. Pilliod, 

and S. McIlroy of the U. S. Geological Survey, Boise, ID; volunteer planting crews via Mary 

Dudley and Mike Young, Idaho Department of Fish and Game, University of Idaho class 

involvement for help in data collection through Beth Newingham, and supplementary funding 

from the Great Basin Landscape Conservation Cooperative.
37

Project Title Species and Population-level Variation inGermination

Strategies of Cold Desert Forbs

Project Agreement No. 13-JV-11221632-080

Principal Investigators and Contact Information

Elizabeth Leger, Associate Professor of Plant Ecology 

Department of Natural Resources and Environmental Science 

University of Nevada

Reno, NV 89557

775.784.7582, FAX 775.784.4789

eleger@cabnr.unr.edu

Sarah Barga, Ph. D. Student

Department of Natural Resources and Environmental Science 

University of Nevada

Reno, NV 89557

306.528.7368, FAX 775.784.4789

sarahcatherinebarga@gmail.com

Project Description

Background

There is an increasing interest in the restoration of degraded sagebrush communities that lack a 

native, herbaceous understory. Native forbs are not only desired for their value to wildlife, but 

also for their potential to suppress annual invaders, such as cheatgrass. Currently, there is a lack 

of knowledge regarding the germination strategies of many cold desert forbs, making it difficult 

for land managers to successfully incorporate herbaceous species into their restoration protocols.

Increasing our knowledge of species specific germination strategies of native forbs can inform 

seed increase protocols, the choice of restoration species in different areas, and the relative 

importance of local seed sourcing for different species. The seed germination of forbs can be 

more challenging than germination of many grasses, and exposing forb seeds to their preferred 

dormancy breaking conditions can dramatically increase their germination success. In addition, 

different species express different amounts of within-species variation in germination cues, 

which may positively correlate with the importance of local seed sourcing for a species.

Objectives

The objective of this project was to assess the germination strategies of ten cold desert forbs, 

including identifying their dominant germination strategy and their level of within-species 

variation in germination cues. We determined seed germination requirements for some common 

species that have not yet been studied and identified species that possess high levels of seed 

dormancy.
38

Methods

This study focused on 10 species of annual and short-lived native forbs that are commonly 

found in sagebrush steppe ecosystems and are of interest due to their potential importance as a 

food source for sage-grouse during early life stages (Figure 1 and Figure 2).

Seeds were collected from multiple populations in the vicinity of Reno, NV and Carson City, NV 

in the spring and summer of 2013. Seeds were placed in petri dishes on wet germination paper, 

and treatments varied after-ripening temperatures (either room temperature or 40°C) and 

exposure to wet, cold conditions (2°C for 0, 2, 4, or 6 weeks) before placement in a moderate 

(15°C) temperature in a factorial design, with each treatment replicated in five dishes. These 

treatments were designed to determine if seeds require warm or cold temperatures to break 

dormancy, which are both common dormancy mechanisms in desert plants. The Survival 

package within Program R (2.12.2) was used to perform survival analysis using the germination 

data.
39

Results

Species varied in preference for after-ripening temperature and cold stratification (Figure 3). Hot 

after-ripening stimulated earlier germination in A. grandiflora, M. gracilis, and P. hastata; 

however, the total number of germinated seeds did not differ between after-ripening treatments. 

Cool after-ripening resulted in both earlier and higher total number of germinated seeds for C. 

douglasii. Species with high levels of germination during cold stratification (potential fall 

germinators) included A. grandiflora, B. scaber, C. parviflora, C. pterocarya, and M. gracilis.

Other species germinated best in warmer temperatures after exposure to cold, including B. 

scaber, C. douglasii, C. intermedia, and P. hastata. Those that germinated best when placed 

directly into 15oC included M. albicaulis and G. inconspicua.

A) B)

Figure 3. Histograms of species preferences for A) after-ripening temperature and B) cold 

stratification.

Figure 4. Examples of survival analysis using the results of cold stratification for

A) a generalist species and B) a species with high dormancy. As the colored lines fallfrom 

left to right, they indicate an increase in total seed germination with time.
40

Survival analysis compares differences in both the timing of germination and the overall quantity 

of germinated seeds based on different treatments (Figure 4). This analysis method allowed for 

identification of generalist species (A. grandiflora, C. parviflora, C. pterocarya, and M. gracilis) 

that germinate under a broad range of conditions and others that possess high levels of 

dormancy, including G. inconspicua, M. albicaulis, and P. hastata (Figure 4). While seed 

dormancy is a strategy than can increase plant fitness over longer time scales, it can be 

undesirable when restoration outcomes depend on seed germination over shorter time scales. We 

also identified population level differences in germination strategies for all species; although 

some differences were more dramatic than others (Figure 5).

Figure 5. Example of survival analysis 

showing dramatic population-level difference 

for C. intermedia.

Table 1. Species specific details for the results of the survival analyses for key treatments.
41

Conclusions

Our ten focal species represented a variety of germination strategies (Table 1). We identified 

some species as facultative winter germinators, if they germinated in cold temperatures. We also 

identified summer germinators as those that germinated best in 15oC, although some required 

cold stratification to achieve optimum germination. We found that all species exhibited 

population level differences in their germination strategies, indicating that seeds may possess 

germination strategies that are closely associated with the environmental conditions at their 

collection location. We are also in the process of assessing what additional environmental cues 

might break dormancy in species expressing high levels of dormancy in this experiment, 

including the addition of soil and smoke treatments.

Our current research aims to identify which environmental characteristics drive variation within 

a species. We are quantifying the level of within-species variation that exists for each species 

using our survival analyses. We will then determine if higher levels of variation in different 

environmental variables, such as precipitation and climate water deficit over the range of a 

species, are correlated with increased population variation in germination requirements. In a 

related project, we are also exploring how variation in type and quantity of disturbance can affect 

seed bank dynamics of understory forbs in sagebrush steppe ecosystems.

Management Applications

Screening plants for their ability to tolerate a variety of field growing conditions will provide 

seed developers with promising species to add to production fields. Further, knowledge about 

species-specific germination preferences and seeding techniques will help managers identify 

methods for incorporating native forbs into restoration protocols. This work will provide 

information on how to germinate species of interest for improving sage-grouse nesting habitat, 

and has the potential to improve recruitment success in areas where food resources are limiting 

for chick development.

Presentations

Barga, S.; Leger, E. A. Species and Population-Level Variation in Germination Strategies of 

Cold Desert Forbs. Oral presentation at the Ecological Society of America 99th Annual Meeting, 

2014 August 10-15; Sacramento, CA.

Barga, S.; Leger, E. A. 2013. Germination Ecology of Great Basin Forbs. Poster presentation at 

the Great Basin Consortium Annual Meeting, 2013 December 9-10; Reno, NV.
42

Project Title Dynamics of cold hardiness accumulation and lossin

Sulphur-flower buckwheat (Eriogonum umbellatum)

Project Agreement No. 13-JV-11221632-066

Principal Investigators and Contact Information

Anthony S. Davis, Associate Professor of Native Plant 

Regeneration and Silviculture

Department of Forest, Rangeland, and Fire Sciences 

P. O. Box 441133

University of Idaho 

Moscow, ID 83844

208.885.7211

asdavis@uidaho.edu

Matthew R. Fisk, Graduate Student, University of Idaho 

Biological Science Technician (Plants)

USDA Forest Service, Rocky Mountain Research Station 

Grassland, Shrubland and Desert Ecosystems Research Program 

Boise, ID 83702

208.373.4358

mfisk@fs.fed.us

Kent G. Apostol, Research Scientist

Department of Forest, Rangeland, and Fire Sciences 

University of Idaho

Moscow, Idaho 83843

651.253.9970

kapostol@uidaho.edu

Project Description

Objectives

Cold hardiness, defined as the capacity of a plant tissue to survive exposure to freezing 

temperatures, is variable across species geographic distribution and annual seasonality. It is a 

well-known indicator of species tolerance and distribution ranges and limits. This is an important 

factor to consider in Great Basin restoration work as diurnal and seasonal temperature 

fluctuations within the region can be extreme. Knowing where the thresholds of temperature 

tolerances lie within individual species is a key component in determining population and 

regional potential for restoration success.

The current research goals are to provide a streamlined strategy for plant producers and 

restoration specialists in the Great Basin to test cold hardiness of forb species and specific 

genetic sources in an effort to provide better guidelines for developing and testing seed zones, as
43

well as producing a greater understanding of the influence of changing climatic conditions on 

plant populations. Using common and valuable candidate Great Basin species, we are working 

with public and private collaborators to build a short guide to cold hardiness. This will greatly 

enhance our ability to both understand the role cold temperatures play in native plant 

establishment following planting, as well as potential implications of changing climatic 

conditions for restoration in the Great Basin.

Materials and Methods

Sulphur-flower buckwheat (Eriogonum umbellatum) is a common, native, valuable subshrub 

whose natural range covers much of the Great Basin region. It has been selected for initial 

investigation to help provide the basis for understanding seasonal cold tolerance accumulation 

and loss within Great Basin native species.

In the fall of 2006 a sulphur-flower buckwheat common garden consisting of seventeen distinct 

populations was established in Boise, ID (Figure 1). In October 2013, five accessions of the 

surviving populations were selected as representatives for the species geographic distribution 

(Table 1). From these five specific accessions, we examined different aspects of the species, 

including geographical range, longitudinal seed production, seed zones, ecoregions, and 

elevational and longitudinal gradients. A sixth population component was also added to the 

study. This sixth component was designed to include one original wildland collection site from 

the five selected accessions growing in the common garden to evaluate localized effects on 

populations.

Figure 1. Sulphur-flower buckwheat common garden located in Boise, ID.

Google Earth 2015
44

Table 1. Sulphur-flower buckwheat population site details.

The freeze-induced electrolyte leakage (FIEL) technique was deployed to evaluate the cold 

tolerance of leaf tissue samples. Index of injury, that is the relative amount of electrolytesleaked, 

was calculated across a range of temperatures and used to determine LT50 values. LT50, the 

temperature at which 50% of cell damage occurs, was selected as the basis of population 

comparison. Tissue samples were collected from the field across an entire calendar year, starting 

in October 2013 on a 6 week cycle. Nine sets of samples were collected and processed to 

complete the year. All samples were transported to Moscow, ID for processing at the University 

of Idaho Seedling Quality Assessment laboratory. Sample collection and processing all took 

place within 48 hours.

Results and Discussion

All field collections and sample processing has been completed. Statistical models are currently 

being designed to calculate LT50 values based on fitting 3-parameter Weibull curves.Initial 

results are showing geographic and seasonal differences among accessions. October 2013 data 

shows a difference of 9o

C across population LT50 levels (Figure 2). Figure 3 shows the 3-

parameter Weibull fitted curves for five September 2013 plant replications of population ERUM 

01 (M).

Future Plans

The remaining tissue samples from field collections not used in the FIEL tests have been oven 

dried, organized, and stored with the future potential to further investigate cold tolerance 

acclimation and loss via carbohydrate analysis or other means. These types of analysis are not 

time sensitive and can be completed at any future date. Interest in further investigation will be 

based on the results from the FIEL work.

A Master’s thesis and journal publications are expected to come out of this work in the next year.

Management Applications and/or Seed Production Guidelines

The techniques from this work can easily be deployed to other species. Upon further 

investigation on plant materials across geographic distributions, we will be able to determine the 

degree to which cold hardiness results can be extrapolated and applied across species and 

populations. The project will provide a streamlined strategy for plant producers and restoration 

specialists in the Great Basin to test cold hardiness of native species, particular genetic sources,
45

and other plant materials in an effort to provide better guidelines for developing and testing seed 

zones, as well as a greater understanding of the influence of changing climatic conditions on 

plant distributions. In order to fill major knowledge gaps, there is a definite need for further 

developed guidelines on cold hardiness.

Figure 2. Percent electrolyte leakage across temperatures for five distinct Sulphur-flower 

buckwheat populations and one wildland collection site.
46

Figure 3. 3-parameter Weibull fitted curves for five 

September 2013 sulphur-flower buckwheat replications 

from accession ERUM 01 (M) used to calculate LT50 of40o Celsius.
47

Presentations

Davis, A. S.; J. R. Pinto; Dumroese, R. K. The Target Plant Concept: case studies in improving 

restoration success. IUFRO World Congress, USA Booth; 2014 October 9; Salt Lake City, UT. 

Invited.

Davis, A. S.; Pinto, J. R. Using the target plant concept to improve dryland restoration success. 

Society for Ecological Restoration: Northwest and Great Basin joint meeting; 2014 October 8; 

Redmond, OR. Invited.

Fisk, M. R.; Pete, K. W.; Apostol, K. G.; Davis, A. S. Dynamics of Germination and Cold 

Hardiness for Great Basin Native Plants (Eriogonum umbellatum). Great Basin Native Plant 

Project Annual Meeting; 2014 September 17-18; Boise, ID. Invited.

Fisk, M. R.; Pete, K. W.; Apostol, K. G.; Pinto, J. R.; Dumroese, R. K.; Davis, A. S. Dynamics 

of Germination and Cold Hardiness for Great Basin Native Plants (Eriogonum umbellatum). The 

Latest & Greatest In Nursery Technology Joint Annual Meeting of The INC, WFCNA, And 

ICSGA; 2014 September 9-11; Boise, ID. Invited.

Pete, K.; Fisk, M. R.; Pinto, J. R.; Davis, A. S. Germination Characteristics of Northern Great 

Basin (Eriogonum umbellatum). American Indian Science and Engineering Society National 

Conference; November 13-15; 2014; Orlando, FL.

Moderator

Fisk, M. R. Re-establishing Big Sagebrush: Integration of Planting Stock to Leverage 

Restoration Success Session, Society for Ecological Restoration, Northwest & Great Basin Joint 

Regional Conference, October 6-10; 2014; Redmond, OR. Invited.

Instructor

Fisk, M. R. Seed Collection for Restoration and Conservation Training; National Training Center 

Course 1730-06; 2014 May 13-15; Boise, ID. Invited.

Fisk, M. R. Freeze Induced Electrolyte Leakage Training. 2014 February 23-28; Hilo, HI. 

Invited.

Exhibitor

Native Wildflower Seed Production Field Day. Ontario, OR. Great Basin Native Plant Selection 

and Increase Project; 2014 May 15; Ontario, OR.
48

Project Title Community Structure of Arbuscular Mycorrhizal Fungi

Colonizing Wyoming Big Sagebrush Seedlings

Project Agreement No. 10-JV-11221632-062

Principal Investigators and Contact Information

Marcelo D. Serpe, Professor 

Department of Biological Sciences 

Boise State University

1910 University Drive

Boise, ID, 83725-1515

208.426.3687, FAX 208.426.4267

mserpe@boisestate.edu

Bill E. Davidson, Graduate Student 

Department of Biological Sciences 

Boise State University

1910 University Drive

Boise, ID, 83725-1515

208.426.3687, FAX 208.426.4267

billdavidson@boisestate.edu

Project Description

Introduction

Arbuscular mycorrhizal fungi (AMF) are obligate biotrophs that obtain organic carbon from the 

host plant and in return facilitate nutrient uptake (Smith et al. 2010). Other effects of 

mycorrhizae on plants have received less attention, but there is evidence that mycorrhizae can 

enhance tolerance to drought (Finley 2008; Auge 2001; Jayne and Quigley 2014), and pathogens 

(Azcon-Aguilar and Barea 1996; Lindermann 2000). All AMF belong to the fungal phylum 

Glomeromycota, which comprises four orders and includes 200 described morphospecies 

(Redecker et al. 2013). However, molecular studies indicate that the number of morphospecies 

might vastly underestimate the true Glomeromycota diversity (Redecker and Raab 2006). AMF 

have low host specificity; a particular AMF species can colonize many plant species. Yet, the

effects of AMF on nutrient uptake and plant growth vary depending on the fungal/plant genotype 

involved in the association and can range from parasitic to mutualistic (Klironomos 2003; Jones 

and Smith 2004). The differential growth responses that result from specific plant/AM symbiosis 

appear to play a major role in determining seedling establishment and plant community 

composition (van der Heijden et al. 1998; van der Heijden et al. 2008).

Host plant preferences and variation in the abiotic environment can affect the composition of 

AMF assemblages (Bell et al. 2009; Torrecillas et al. 2012; Busby et al. 2013). In addition, 

inoculation of seedlings can alter the AMF taxa present in the roots after transplanting (Mummey 

et al. 2009). Results described in more detail in a previous report (Serpe and Davidson 2013) 

showed that inoculation of sagebrush seedlings with native AMF increased colonization of 

sagebrush seedlings over the levels naturally occurring in the soil. Moreover, the increase in
49

colonization was associated with an increase in seedling establishment. This increase in 

establishment may be simply related to higher rates of colonization, but could also reflect 

difference in the AMF taxa colonizing the roots. The AMF taxa used for inoculation were native 

to a sagebrush habitat, but the inoculum was developed from soil collected at only one site. This 

inoculum was introduced into a new site in one of the experiments, which could have led to 

differences in AMF composition between non-inoculated and inoculated seedlings. Furthermore, 

multiplication of AMF for the production of inoculum can change the AMF community 

(Sykorova et al. 2007; Carter et al. 2014). To assess the effect of inoculation on the AMF taxa 

colonizing the roots, the same samples that were used to determine colonization were analyzed 

for AMF community composition using molecular methods. An additional motivation for these 

analyses was to characterize the diversity of AMF taxa present in sagebrush seedlings and to 

investigate possible seasonal changes in AMF.

Objectives

1) Determine whether pre-inoculation with native AMF alters the structure of AMF 

communities colonizing Wyoming big sagebrush seedlings.

2) Gain knowledge of AMF taxa colonizing Wyoming big sagebrush seedlings.

3) Investigate possible seasonal changes in AMF taxa.

Methods

To produce mycorrhizal inoculum, sandy-loamy soil was collected from Kuna Butte, Idaho 

(43°26.161’N, 116°25.848’W). The AMF present in the soil were multiplied in pot cultures that 

were prepared as previously described using Sudan grass as a host (Carter et al. 2014). Wyoming 

big sagebrush seedlings were first grown in a greenhouse in sterilized soil (non-inoculated 

seedlings) or soil containing the mixture of native mycorrhizal species from the pot cultures 

(inoculated seedlings). Three-month old seedlings were transplanted outdoors to 24 L potsfilled 

with soil from Kuna Butte or to a recently burned sagebrush habitat near Bigfoot Butte, ID 

(43°18'48.43"N, 116°21'48.57"W), within the Morely Nelson Snake River Birds of Prey NCA. 

Seedlings were transplanted to pots (mesocosm experiments) in the spring and fall of 2011 and 

to the burned site in the spring and fall of 2012. Samples for analyses of colonization and AMF 

community composition were collected approximately 4 and 7 months after spring and fall 

transplanting, respectively. These samples were stored at -20°C until used for DNA extraction, 

amplification, and cloning. The AMF taxa colonizing individual seedlings were identified based 

on phylogenetic analysis of sequences from the large subunit-D2 rDNA region (Mummey and 

Rillig 2007). Sequences were also grouped into operational taxonomic units (OTUs) with 

sequence similarities ≥ 94% using the Mothur program (Schloss et al. 2009; Wang et al. 2011). 

Possible differences in AMF composition between non-inoculated and inoculated seedlingswere 

analyzed using non-metric multidimensional scaling (NMDS). This analysis was also used to 

characterize variations in AMF communities among seasons or between shallow and deep roots. 

NMDS is an unconstrained ordination method that arranges communities according to their 

similarities in species composition (Minchin 1987).

Results and Discussion

Phylogenetic and Diversity Analyses

The phylogenetic analysis revealed that all the sequences obtained were in the order Glomerales 

(Figure 1). About 5% of these sequences belonged to the family Claroideoglomeracea and the
50

rest to the Glomeraceae. Within the Claroideoglomeracea, the analysis resolved two phylotypes, 

Claroideoglomus I and II. These sequences had less than 94% similarity with Claroideoglomus 

claroideum; but had up to 99% similarity with several published sequences of uncultured 

glomeromycetes (Mummey and Rillig 2007; Verbruggen et al. 2010; Carter et al. 2014). Within 

the Glomeraceae, the analysis resolved four phylotypes. Two phylotypes were within the Glomus 

genus and were designated Glomus I and Glomus II. Sequences within Glomus I clustered with 

Glomus macrocarpum. Glomus II was the phylotype with more sequences. A few of these 

sequences clustered with Glomus microaggregatum. However, the majority of the sequences 

within Glomus II did not cluster with known species, but showed high similarity with published 

sequences of uncultured and unnamed glomeromycetes (Mummey and Rillig 2007; Torrecillas 

et al. 2012; Carter et al. 2014). Sequences were also identified within the Funneliformis and 

Rhizophagus genera. The former clustered with sequences obtained from single spore of F. 

mosseae or isolates of F. mossea. Similarly, sequences within the Rhizophagus phylotype 

clustered with Rhizophagus intraradices or R. irregularis.

The phylotypes obtained by the Maximum Parsimony analysis were compared with resultsfrom 

pairwise distance analysis. There was a good agreement between the results of these analyses; a 

particular OTU was only present in one phylotype (Figure 1). On the other hand, all phylotypes 

included more than one OTU. The largest number of OTUs was detected in Rhizophagus 

followed by Glomus II; which had eight and seven OTUs, respectively. Glomus II included 

sequences showing pairwise differences of up to 22%. Considerable genetic variation was also 

observed in the other phylotypes. Funneliformis, Glomus I, and Claroideoglomus I and II

contained 5, 4, 3, and 2 OTUs, respectively. Furthermore, pairwise differences within these 

phylotypes ranged from 12% in ClaroideoglomusII to 25% in Funneliformis.

AMF Community Composition in Non-inoculated and Inoculated Seedlings

To evaluate whether inoculation altered the AMF community of the roots, we compared the 

richness and diversity of OTUs in non-inoculated and inoculated seedlings. Within a particular 

experiment, no differences were detected in the richness index (S) between non-inoculated and 

inoculated seedlings (Table 1). Among experiments, the average S values ranged from 3 to 5, 

indicating that most seedlings were colonized by AMF in more than one OTU. However, the 

Shannon index of diversity (H’) was much lower, suggesting the predominance of certain OTUs 

over others (Table 1). Like for S, the H’ values did not differ among treatments, except for the 

spring field experiment. In this experiment, H’ was somewhat higher in non-inoculated than 

inoculated seedlings.

Although richness and diversity were similar, the particular OTUs colonizing the roots could 

have been different among treatments. This possibility was analyzed by nonmetric

multidimensional scaling. For both the mesocosm and field experiments, NMDS plots showed no 

significant differences between non-inoculated and inoculated treatments (Figure 2). In addition 

for the spring field experiment, we compared the AMF community of roots collected within the 

upper 20 cm of the soil with those collected below 30 cm. These communities showed little 

separation in the ordination space and their centroids were not significantly different (p = 0.89, 

data not shown).
51

Analysis of Seasonal Changes in the AMF Community

For the mesocosm and field experiments, we also compared the AMF communities of 

seedlings harvested during the summer (4 to 5 months after spring transplanting) to those 

harvested in early spring (7 to 8 months after fall transplanting). Seedlings from the 

mesocosm experiments showed more seasonal variation in their centroids than those from the 

field experiments (Figure 3, A and B). However, in both analyses the centroids were not

significantly different with p values of 0.16 and 0.73 for the mesocosm and field 

experiments, respectively

Table 1. Richness (S) and Shannon-Wiener Diversity (H’) indices of AMF OTUs detected in roots 

of (n) individual sagebrush seedlings. Mean ± SE for each study. Within a particular experiment 

and transplanting time, values that differ significantly from the non-inoculated seedlings are 

indicated by an asterisk.

Experiment Transplanting 

time Treatment n Richness (S) Diversity (H’)

Mesocosm

Spring

Non-inoculated 5 4.70 ± 1.20 1.21 ± 0.32

Inoculated 5 5.00 ± 0.58 1.13 ± 0.09

Fall Non-inoculated 8 4.13 ± 0.72 0.95 ± 0.22

Inoculated 8 3.40 ± 0.42 0.74 ± 0.14

Field

Spring Non-inoculated 6 5.00 ± 0.52 0.96 ± 0.10

Inoculated 6 3.83 ± 0.31 0.64 ± 0.07 *

Fall Non-inoculated 6 3.67 ± 0.56 0.82 ± 0.15

Inoculated 6 2.83 ± 0.48 0.59 ± 0.17

The lack of significant differences in AMF communities between inoculation treatments, root 

depth, and seasons appeared to have been related to the predominance of certain OTUs. In all 

experiments and treatments, the most common OTU was GII-1 (Figure 4). Overall this OTU

was detected in 94% of the seedlings (Figure 4, E). Other OTUs that were present in relatively

high frequency were F1, GII-2, and R1, which were detected in 54, 51, and 40% of the

seedlings respectively. For these OTUs, similar patterns were in general observed in individual 

experiments (Figure 4, A-D). Funneliformis 1 and GII-2 were found in more than 50% of the 

seedlings in three of the four experiments, the exception was the fall 11 experiment, where only

18% and 31% of the seedlings had F1 and GII-2, respectively. The fourth most abundant OTU,

R1, showed more variation. In the mesocosm experiments (Figure 4, A and B), R1 was found in

at least half of the seedlings. In contrast, in the field experiments, R1 was not detected in one of

the treatments or it was found at low frequency on the other (Figure 4, C and D). Another major 

factor contributing to the dominance of certain OTUs over others was that several OTUs were 

only found in very few seedlings. Specifically, 15 of the 29 OTUs were detected in less than

5% of the seedlings and 10 of them in less than 2%. Thus, the overall contribution of these 

OTUs to the AMF community of sagebrush seedlings was probably nominal. Further work is 

needed to evaluate whether the differences in OTU abundance reflect preference by the host 

plant or differences in OTU frequencies within the soil.
52

Figure 1. Phylogenetic tree of 

representative Glomeromycota 

LSU-D2 rDNA sequences 

detected in this study with the 

FLR3/FLR4 primers and 

database sequences of known 

Glomeromycota. The values 

above the branches are bootstrap 

support values obtained by 

Maximum Parsimony analysis. 

Names on the right indicate the 

six phylotypes identified in the 

phylogenetic analysis. Smaller 

vertical lines within each 

phylotype indicate operational 

taxonomic units (OTUs) with a 

threshold level of 94% sequence 

similarity based on pairwise 

distance analysis.
53

Figure 2. Comparison of arbuscular mycorrhizal communities of non-inoculated and inoculated 

seedlings by nonmetric multidimensional scaling. Samples of Artemisia tridentata seedlings 

were collected several months after transplanting in the spring (A) and fall (B) mesocosm 

experiments, and the spring (C) and fall (D) field experiments. OTUs are represented by open 

triangles and seedlings with filled circles. The centroids for the each treatment are labeled(NI, 

non-inoculated seedlings; I, inoculated seedlings) and linked to the samples that relate to them. 

Ellipses represent 95% confidence limits around the centroids of each variable. OTUs are 

labeled based on their phylotype (CI, Claroideoglomus I; CII, Claroideoglomus II; F, 

Funneliformis; GI, Glomus I; GII, Glomus II; R, Rhizophagus) and OTU number within its 

phylotype.

A B

C D
54

Figure 3. Analysis of seasonal changes in arbuscular mycorrhizal communities by nonmetric 

multidimensional scaling. (A) Mesoscosm experiments, comparison of AMF communitiesin 

seedlings collected 5 and 8 months after spring and fall transplanting, respectively. (B) Field 

experiments, comparison of AMF communities in seedlings collected 4 and 8 months after 

spring and fall, respectively. OTUs are represented by open triangles and seedlings withfilled 

circles. The centroids for each categorical variable are labeled and linked to the samplesthat 

relate to them. Ellipses represent 95% confident limits around the centroids of each variable.

B

A
55

Figure 4. Percent of Artemisia tridentata seedlings colonized by different OTUs. 

Results from the spring (A) and fall (B) mesocosm experiments, and from the spring

(C) and fall (D) field experiments. Overall frequencies observed after combining the 

results from all the seedlings analyzed (E).
56

Management Applications

Restoration projects often use seedlings that have been pre-inoculated with commercial and nonnative mycorrhizae. This can alter the resident AMF community with potentially detrimental 

ecological consequences. In principle, alterations of the AMF community can also occur 

following inoculation with native AMF if the process of AMF multiplication favors certain taxa

over others. The results obtained in this study indicate that a native AMF community wasrather 

resilient to changes in composition during multiplication and subsequent inoculation.

Consequently, this approach appears to be a feasible strategy for increasing AMF colonization of 

sagebrush seedlings without the risks associated with changing the AMF community. The 

dominance of a few taxa may also simplify the production of inoculum. Production of 

monospecific cultures of these taxa and subsequent mixing would allow the development of a 

more precise and uniform inoculum.

Acknowledgments

Funding for this research was provided by the USDI Bureau of Land Management, Great Basin 

Restoration Initiative and the USDA Forest Service Rocky Mountain Research Station,Great 

Basin Native Plant Selection and Increase Project.

Presentations

Serpe, M. D.; Davidson, B. E. Pre-inoculation of Wyoming big sagebrush seedlings with native 

arbuscular mycorrhizae: effects on mycorrhizal colonization and seedling survival after 

transplanting. Great Basin Native Plant Annual Meeting; 2014 March 17-18; Bois, ID.

Davidson, B. E.; Smith, J. F.; Serpe, M. D. Inoculation of Artemisia tridentata ssp.

wyomingensis with arbuscular mycorrhizae: impacts on colonization and mycorrhizal 

composition of roots after transplanting. Botanical Society of America Conference; 2014 July 

26-30; Boise, ID. Poster. .

Davidson, B. E.; Serpe, M. D. Inoculation of Wyoming big sagebrush seedlings with native 

arbuscular mycorrhizae: impacts on root colonization, mycorrhizal community composition, and 

seedling survival after transplanting. Society for Ecological Restoration Regional Conference, 

Northwest Great Basin Chapter. 2014; Redmond, OR.

References

Auge, R. M. 2001. Water relations, drought and vesicular-arbuscular mycorrhizal symbiosis. 

Mycorrhiza 11: 3-42.

Azcon-Aguilar, C.; Barea, J. M. 1996. Arbuscular mycorrhizas and biological control ofsoilborne plant pathogens: an overview of the mechanisms involved. Mycorrhiza 6: 457-464.

Bell, C. W.; Acosta-Martinez, V.; McIntyre, N. E.; Cox, S.; Tissue, D. T.; Zak, J. C.2009. 

Linking microbial community structure and function to seasonal differences in soil moisture and 

temperature in a Chihuahuan Desert Grassland. Microbial Ecology 58: 827-842.

Busby, R. R.; Stromberger, M. E.; Rodriguez, G.; Gebhart, D. L.; Paschke, M. W. 2013. 
57

Arbuscular mycorrhizal fungal community differs between a coexisting native shrub

and introduced annual grass. Mycorrhiza 23: 129-141.

Carter, K. A.; Smith, J. F.; White, M. M.; Serpe, M. D. 2014. Assessing the diversity of 

arbuscular mycorrhizal fungi in semiarid shrublands dominated by Artemisia tridentata ssp. 

wyomingensis. Mycorrhiza 24: 301-314.

Finley, R. D. 2008. Ecological aspects of mycorrhizal symbiosis with special emphasis on the 

functional diversity of interactions involving the extraradical mycelium. Journal of Experimental 

Botany 59: 1115-1126.

Jayne, B.; Quigley, M. 2014. Influence of arbuscular mycorrhiza on growth and reproductive 

response of plants under water deficit: a meta-analysis. Mycorrhiza 24: 109-119.

Jones, M. D.; Smith, S. E. 2004. Exploring functional definitions of mycorrhizas: Are 

mycorrhizas always mutualisms? Canadian Journal of Botany-Revue Canadienne De Botanique 

82: 1089-1109.

Klironomos, J. N. 2003. Variation in plant response to native and exotic arbuscular mycorrhizal 

fungi. Ecology 84: 2292-2301.

Lindermann, R. 2000. Effects of mycorrhizas on plant tolerances to diseases. In: Y. Kapulnik,D.

D. J. Douds (ed. s) Arbuscular mycorrhizas: physiology and function. Dordrecht, the 

Netherlands: Kluwer Academic Publishers: 345-365.

Minchin, P. R. 1987. An evaluation of the relative robustness of techniques for ecological 

ordination. Vegetatio 69: 89-107.

Mummey, D. L.; Antunes, P. M.; Rillig, M. C. 2009. Arbuscular mycorrhizal fungi pre-inoculant 

identity determines community composition in roots. Soil Biology and Biochemistry 41: 1173-

1179.

Mummey, D. L.; Rillig, M. C. 2007. Evaluation of LSU rRNA-gene PCR primers for analysis of 

arbuscular mycorrhizal fungal communities via terminal restriction fragment length 

polymorphism analysis. Journal of Microbiological Methods 70: 200-204.

Redecker, D.; Raab, P. 2006. Phylogeny of the Glomeromycota (arbuscular mycorrhizalfungi): 

recent developments and new gene markers. Mycologia 98: 885-895.

Redecker, D.; Schuessler, A.; Stockinger, H.; Stuermer, S. L.; Morton, J. B.; Walker, C. 2013. An 

evidence-based consensus for the classification of arbuscular mycorrhizal fungi 

(Glomeromycota). Mycorrhiza 23: 515-531.

Serpe, M. D.; Davidson, B. E. 2013. Pre-inoculation of Wyoming big sagebrush seedlings with 

native arbuscular mycorrhizae: effects on mycorrhizal colonization and seedling survival after 

transplanting. Great Basin Native Plant Project 2013 Progress Report: 61-67.

Schloss, P. D.; Westcott, S. L.; Ryabin, T.; Hall, J. R.; Hartmann, M.; Hollister, E.B.;
58

Lesniewski, R. A.; Oakley, B. B.; Parks, D. H.; Robinson, C. J.; Sahl, J. W.; Stres, B.;Thallinger,

G. G.; Van Horn, D. J.; Weber, C. F. 2009. Introducing mothur: open-source, platformindependent, community-supported software for describing and comparing microbial 

communities. Applied and Environmental Microbiology 75: 7537-7541.

Smith, S. E.; Facelli, E.; Pope, S.; Smith, F. A. 2010. Plant performance in stressful 

environments: interpreting new and established knowledge of the roles of arbuscular 

mycorrhizas. Plant and Soil 326: 3-20.

Sykorova, Z.; Ineichen, K.; Wiemken, A.; Redecker, D. 2007. The cultivation bias: different 

communities of arbuscular mycorrhizal fungi detected in roots from the field, from bait plants 

transplanted to the field, and from a greenhouse trap experiment. Mycorrhiza 18: 1-14.

Torrecillas, E.; del Mar Alguacil, M.; Roldan, A. 2012. Differences in the AMF diversity in soil 

and roots between two annual and perennial gramineous plants co-occurring in a Mediterranean, 

semiarid degraded area. Plant and Soil 354: 97-106.

van der Heijden, M. G. A.; Bardgett, R. D.; van Straalen, N. M. 2008. The unseen majority:soil 

microbes as drivers of plant diversity and productivity in terrestrial ecosystems. Ecology Letters 

11: 296-310.

van der Heijden, M. G. A.; Boller, T.; Wiemken, A.; Sanders, I. R. 1998. Different arbuscular 

mycorrhizal fungal species are potential determinants of plant community structure. Ecology 79: 

2082-2091.

Verbruggen, E.; Roling, W. F. M.; Gamper, H. A.; Kowalchuk, G. A.; Verhoef, H. A.; van der 

Heijden, M. G. A. 2010. Positive effects of organic farming on below-ground mutualists: largescale comparison of mycorrhizal fungal communities in agricultural soils. New Phytologist 186: 

968-979.

Wang, Y.; Huang, Y.; Qiu, Q.; Xin, G.; Yang, Z.; Shi, S. 2011. Flooding greatly affectsthe 

diversity of arbuscular mycorrhizal fungi communities in the roots of wetland plants. PLoS ONE 

6(9): e24512.
59

Project Title Smoke-induced Germination of Great Basin Native Forbs

Project Agreement No. 11-CR-11221632-005

Principal Investigators and Contact Information

Robert Cox, Assistant Professor 

Texas Tech University

Box 42125

Lubbock, TX 79409

806.742.2841, FAX 806.742.2280

robert.cox@tt.edu

Project Description

Objectives

Determine responsiveness to smoke of several species native to the Great Basin and how that 

responsiveness varies by population.

Methods

Step one is to conduct a wide literature review of as many species as possible. Step two is to test 

as many species as possible to determine smoke response. Step three is to further investigate 

responsive species to determine whether smoke response varies by population by testing seeds 

from as many populations of the species in question as possible. Step four will be to develop 

seeding protocols for the most responsive species that include smoke treatments.

Results and Discussion

The literature review has been completed, and is now being organized by species. We are 

preparing portions of this literature review as a publication that will analyze seed/species 

characteristics as a method for predicting possible smoke response. We have now tested 19 

species for smoke response, and are analyzing the results. Species tested are: Achillea 

millefolium, Achnatherum hymenoides, Achnatherum thuberianum, Agoseris grandiflora,

Artemisia tridentata, Atriplex canescens, Balsamorhiza hookeri, Balsamorhiza saggittata, 

Chrysothamnus nauseosus, Erigeron pumilus, Eriogonum heracleoides, Eriogonum umbellatum, 

Grayia spinosa, Lomatium dissectum, Lomatium grayi, Penstemon speciosus, Phacelia hastata, 

Poa secunda, and Pseudorognena spicata.
60

Project Title Herbicide Impacts on Forb Performance in Degraded

Sagebrush Steppe Ecosystems

Project Agreement No. 14-JV-11221632-067

Principal Investigators and Contact Information

Marie-Anne De Graaff, Assistant Professor 

Department of Biological Sciences

Boise State University 

Boise, ID 83725

208.426.1256, FAX 208.426.1040

Marie-annedegraaff@boisestate.edu

Aislinn Johns, Undergraduate Student 

Department of Biological Sciences 

Boise State University

Boise, ID 83725

208.713.5323, FAX 208.426.1040

aislinnjohns@u.boisestate.edu

Project Description

Introduction

Large areas of the sagebrush steppe ecosystem have been invaded by Bromus tectorum 

(cheatgrass), leading to a change in ecosystem structure (i.e. from shrub to grass dominated). 

This threatens sagebrush dependent species such as Centrocercus urophasianus (sage-grouse), a 

candidate for protection under the Endangered Species Act. An herbicide containing the active 

ingredient imazapic is commonly used to suppress cheatgrass invasion. Imazapic kills plants by 

inhibiting the production of branched chain amino acids, which are necessary for protein 

synthesis and cell growth.

Although imazapic is generally an effective herbicide for weed control, little is known about its 

impact on performance of non-target species, such as forbs native to the sagebrush steppe 

ecosystem that are essential for sage-grouse survival. The impact of imazapic on non-target 

species may hinge on its persistence in the soil from where it can effect plant growth. Imazapic is 

degraded primarily by soil microbial community, and has an average half-life in soil of 120 days. 

However, in dry lands, such as a semi-arid desert, microbial activity is frequently limited by 

water availability which may prolong the predicted half-life time of imazapic in soil. Further, 

these soils are frequently subjected to fire, which alters the soil microbial community and its 

activity, thus potentially changing the decay rate of imazapic.
61

Objectives

With this study we set out to improve our understanding of the impact of imazapic on forbs 

native to the sagebrush steppe ecosystem. Specifically we asked:

 How do different application rates (concentrations) affect forb performance?

 How does the timing of imazapic application relative to seeding of forbs impact forb 

performance?

 Does a fire-induced change in soil microbial activity mediate the impact of imazapic on 

soils?

Methods

To assess the impact of fire reduction treatments on 

the performance of native forb species, we 

conducted two greenhouse experiments in which 

we (1) applied a gradient of imazapic 

concentrations, and (2) varied the timing of 

application relative to seeding of Astragalus filipes, 

Achillia millefolium, and Sphaeralcea 

grossularifolia (Figure 1). Soils were collected in 

degraded-burned and unburned sagebrush 

ecosystems (0-10 cm depth) at the Morley Nelson 

Birds of Prey National Conservation Area. The soil 

was air-dried for 7 days, litter and roots were 

removed by using a 2mm sieve, and kept under dry 

conditions at 20-23°C until ready for planting.

A 15-day laboratory incubation with the soil was performed to confirm that 

the fire had altered microbial activity in the recently burned soil compared to 

the unburned soil. Soils from each treatment were placed in quart mason jars, 

then incubated in the dark for 15 days under controlled soil moisture (60% 

WHC) and temperature (20°C) conditions. Potential C decomposition rates 

were measured on days 1, 3, 5, 7 and 15 with a LI-7000 CO2/H2O analyzer

(http://www. licor. com/env/products/gas_analysis/).

Both burned and unburned soils were transferred to pots (6. 4 cm diameter and

12. 7cm depth) that were lined with a nylon sack (Figure 2). The nylon sack 

was filled with cleaned rocks and then sealed, after which the remainder of 

the pot was filled with 250g of soil. Soil was watered evenly to 60% water 

holding capacity.

Imazapic Concentration Study

The imazapic concentration experiment was set up as a complete 

randomized block design (n=10) (Figure 4). Imazapic concentrations of 0, 2, 

4 and 8 oz a. i. /acre were applied evenly to pots 2 weeks prior to planting

forbs. Three sage-grouse preferred forbs Achillea millefolium (A. m. ), Astragalus filipes (A. f), 

and Sphaeralcea grossulariifolia (S. g), were planted in pots for a controlled greenhouse 

experiment after the seeds were germinated according to their specific germination requirements

Figure 1. Greenhouse experiment

Figure 2. Pot

preparation
62

(Table 1). The seeds were planted with equal spacing (Figure 3) and the pots were randomized 

every 3 weeks. Plants were grown at 15°C-25°C for 41 days. We measured height, maturity,

emergence, and aboveground and belowground biomass on a biweekly basis. Data were

analyzed using a Two-Way Analysis of Variance (ANOVA) in SPSS.

Figure 4. Complete randomized block design of imazapic impacts on forb performance greenhouse

experiment. Soil A and B are unburned and burned soils, respectively. On each one of the soils, one of

three forb species (Astragalus filipes, Achillia millefolium, Sphaeralcea grossularifolia) is planted. All are

exposed to one of four imazapic concentrations (including a control treatment receiving wateronly).

Table 1. Germination protocol

Forb Scarify Stratify Surface 

Sterilize

Plant depth

Astragalus filipes 220 grit sandpaper

for 3 minutes (50

seeds X 8)

NO NO 3. 5 mm

Achillia 

millefolium

NO NO NO 0 mm

Sphaeralcea 

grossularifolia

18 M H2SO4 for 10

min (200 seeds X 4)

3/15/13- 4/29

Refrigerated (~5°C) 

in petri dishes on 

moist filter paper for 

7 weeks

NO 6. 5 mm

Bromus tectorum NO NO NO Light 

dusting

Imazapic Application

The imazapic application timing experiment was a controlled greenhouse experiment. Burned 

and unburned soils were composited and transferred to pots in the above method. Imazapic

 1

 2

 3

 4

 1

 2

 3

 4
63

(4oz a. i. /acre) or water containing 1% mineral oil surfactant was added by spray to all of the 

pots. The above-mentioned forbs and Bromus tectorum (n=10) were then planted after 2, 4, 8, 

and 16 weeks in a pattern (represented in Figure 5). The plants were watered daily and kept at 

optimal conditions in the greenhouse. Soils waiting to be planted were maintained under the 

same conditions and watered three times weekly.

The plants grew for 41 days when the height was recorded and the foliage was then harvested for 

aboveground biomass. The plant matter was dried at 40°C for 24 hours and weighed to obtain 

aboveground biomass. The soils were frozen at -20°C until belowground biomass could be 

processed. At the time of processing the live roots were removed from the soil, washed and dried 

in the oven at 40°C for 24 hours. After drying, the roots were weighed to obtain belowground 

biomass. The differences were then analyzed using a Two-Way ANOVA in SPSS.

.

Results and Discussion

Experiment 1: Impact of Herbicide Concentrations on Forb Performance

Burning significantly affected soil microbial activity (p < 0.05); recently burned soils had a 

greater content of labile C leading to greater resource availability for microbes and greater CO2 

respiration rates as compared to soils that were not burned (Figure 6). These data indicate that 

changes in the soil microbial community occurred following a fire and these changes may have 

altered decay rates of imazapic, hence its impact on forb performance. However, we found that 

imazapic application significantly suppressed biomass production for each of the species

p < 0.05, n = 10; Table 2), but that burning did not impact the response of forbs to imazapic 

applications.

B. t. S. g. A. m.

N=10 Plant Species

4 oz a. i.

/acre

Control

2 4 8 1

Imazapic

Application

Week Planted

Figure 5. Experimental design for herbicide application timing impacts on forb performance. Ima

zapic Co

n

c

entration
64

The magnitude of the response to imazapic application differed among species. We calculated 

the response as the percent change in biomass production relative to the control (i.e.no imazapic 

application); where a greater response signifies a greater sensitivity of the forb to imazapic. The 

average response of above and belowground biomass production was significantly different

(p < 05) across forb species (n = 20). Specifically, Astragalus filipes (A. f. ) was affected by the 

herbicide the least (n = 20).

Figure 6. Impact of fire on soil microbial activity. Values

are means + SE (n = 5). Different letters indicate

significant differences in CO2 respiration betweenburned

and unburned soils.

Experiment 2: Impact of Timing of Imazapic Application Relative to Seeding on Forb 

Performance.

Since fire had no impact on imazapic 

effects on forb performance, we did not 

include the burned soils in the second 

greenhouse experiment. Instead, in this 

experiment we focused on the timing of 

imazapic application to forbs relative to 

seeding. In addition, we included 

cheatgrass to ascertain if any impacts of 

imazapic on forb performance also impact 

cheatgrass growth. For each one of the 

forbs, biomass production in soils not 

receiving imazapic was greater when 

plants were seeded into soil at a later date 

(Table 3). This results from the multiple 

irrigation events that promoted soil organic

matter decomposition and the release of 

nutrients in soil over time. Our results 

further indicated that imazapic negatively

Figure 8. Visual differences of forb species containing

different concentrations of imazapic.

impacted biomass production of forbs and cheatgrass regardless of application timing (Table 4).

b

a
65

Interestingly, performance of Astragalus filipes was much less negatively impacted by imazapic 

across both experiments (Figure 8). A. filipes is a legume, whereas the others are not. These data 

indicate that legumes may be impacted less by imazapic applications than other plantsfunctional 

types.

Conclusions

Imazapic significantly impacts forb performance, and burning did not mediate the impact of 

herbicide on forb performance. This suggests that imazapic is not specific to cheatgrass and that 

the use of imazapic could impact species diversity and essential sage-grouse preferred forbs. This 

information should be taken into account in future restoration projects.

Ongoing research funded by the Forest Service will assess whether imazapic impacted N 

mineralization in the soil. This may explain why performance of the legume was not affected by 

imazapic to the extent that the other forb species were.
66

Table 2. Total standing biomass (shoots + roots) at harvest for Astragalusfilipies,

Achillea millefolium, and Sphaeralcea grossulariifolia. Values are means + SE (n =

10). Significant differences within treatments are shown by different letters above

the gaps (p<0. 05, N=10).
67

Table 3. Differences in the response to imazapic application among the three forbs. A. f.

(Astragalus filipes), A. m. (Achillea millefolium), S. g. (Sphaeralcea grossulariifolia).

Values are means + SE (n = 20). Different letters indicate significant differences inresponse

to imazapic among forbs.
68

0 oz/acre

4 oz/acre

a

a

a

b

b b

Table 4. Differences in the total biomass at different imazapic application times for Astragalus

filipes, Achillea millefolium, Sphaeralcea grossulariifolia, and Bromus tectorum. Values are means

+ SE (n = 10). Different letters indicate significant differences in response to imazapic application

between control and imazapic treatments.

8a. Astragalusfilipes 8c. Sp. haeralcea grossulariifolia

0.03

0.06

0.02 0.04

0.01 0.02

0.00 0.00

8b. Achilleamillefolium 8d.Bro.musTectorum

0.04 0.06

0.03

0.02

0.01

0.00 2 4 8 16

0.04

0.02

0.00

2 4 8 16

Weeks until seeding followingthe application of imazapic

Presentations

Johns, A.; de Graaff, M. A. Herbicide impacts on forb performance in degraded sagebrush steppe 

ecosystems. Society for Ecological Restoration Regional Conference, Northwest and Great Basin 

Chapters; 2014 October 6-10; Redmond, OR. Poster.

de Graaff, M. A. 2014. Herbicide impacts on forb performance in degraded sagebrush steppe 

ecosystems. Great Basin Native Plant Project Annual Meeting; 2014 March 17-18; Boise, ID.

Acknowledgements

This project was supported by the USDI Bureau of Land Management, the U.S. Forest Service 

Great Basin Native Plant Project, and by the National Science Foundation EPSCoR Program 

under award number EPS-0814387.

a

a

b b

a

a

a

a

b b b

b

Biomass (g)
69

Project Title Increasing the Availability and Integration of GreatBasin

Native Plant Project Tools for BLM Restoration 

Practitioners

Project Agreement No. 13-IA-11221632-161

Principal Investigators and Contact Information

Anne Halford, Idaho State Botanist/GBNPP Liaison 

DOI BLM Idaho State Office

1387 S. Vinnell Way 

Boise, ID 83709

208.373.3940

ahalford@blm.gov

Additional Collaborators

Francis Kilkenny, Research Biologist 

USDA Forest Service

Rocky Mountain Research Station 

322 E Front Street Suite 401

Boise, ID 83702

208.373.4376

ffkilkenny@fs.fed.us

Nancy Shaw, Research Botanist (Emeritus) 

USDA Forest Service

Rocky Mountain Research Station 

322 E. Front Street, Suite 401

208.373.4360

nancylshaw@fs.fed.us

Vicki Erickson, Regional Geneticist & Native Plant Program 

Manager

USDA Forest Service 

Pacific Northwest Region 

Pendleton, OR 97801

541.278.3715

verickson@fs.fed.us

Doug Kendig, Native Plant/Restoration Coordinator 

DOI BLM Medford District

3040 Biddle Road

Medford, OR 97504

541. 618.2285
70

Project Description

There is a critical need within the Bureau of Land Management (BLM) to step-up the integration 

and use of applied research tools and products that inform restoration of sagebrush-steppe 

systems. Key BLM and United States Forest Service (USFS) landscape planning efforts that are 

driving this immediate need include 1) the Greater Sage-Grouse Environmental Impact 

Statement (EIS) and commensurate plan amendments that will be required, and 2) the Fire and 

Invasive Assessment effort (FIAT) that identifies management strategies (Chambers et. al 2014) 

to reduce the habitat threats resulting from impacts of invasive annual grasses, wildfires and 

conifer expansion. In addition, in January 2015 Department of Interior Secretary, Sally Jewel, 

issued Secretarial Order 3336 (USDI 2015) that sets forth enhanced policies and strategies for 

preventing and suppressing rangeland fire and for restoring sagebrush landscapes impacted by 

fire across the West. One of the key sections of the order directs BLM to develop an enhanced 

fire prevention, suppression and restoration strategy. Sections germane to restoration include:

 Apply science and research to improve the identification and protection of resistant and 

resilient sagebrush-steppe landscapes and the development of biocontrols and other tools 

for cheatgrass control to improve capability for long-term restoration of sagebrush-steppe 

ecosystems.

 To the extent practicable, utilize locally-adapted seeds and native plant materials 

appropriate to the location, conditions, and management objectives for vegetation 

management and restoration activities, including strategic sourcing for acquiring,storing, 

and utilizing genetically appropriate seeds and other plant materials native to the 

sagebrush-steppe ecosystem.

For more than a decade, the Great Basin Native Plant Project has been at the forefront of 

addressing regionally relevant applied research questions that cover ecologically complex topics 

such as, genetics, climate change, native plant interactions, plant cultural practices, plant material 

development and restoration. To assist BLM in better integrating this knowledge and use of 

available tools we have been working with our partners and BLM Botanists and Natural 

Resource Specialists throughout Idaho, Oregon, Nevada, Utah and eastern California to work on 

acquisition and deployment of genetically appropriate native plant material for restoration.

Fundamental to this process is the increased coordination of Great Basin-wide seed collectionsto 

accelerate acquisition and production of seed stock for plant increase. We will be drawing on 

“lessons learned” from these efforts (Shaw et. al 2005; Youtie 2013). Via development of a 

“seed increase” technical team, we are integrating a systematic procedure to refine this work.

Our goal is to identify species, and/or type of plant materials (seeds, plant container stock) and 

average quantity of each needed in each provisional (Bower et. al 2014) or empirical seed zone 

when available for the next 10 years. The first priority is development and refinement ofspecific 

plant material needs by these seed zones for workhorse species that are, or could be, increased in 

the agricultural market. This can be further refined by prioritizing seed zones and plant 

community types where restoration is a priority.
71

A commensurate task will be to develop seed management plans for each species that will 

include the following information:

 Average quantity of each species/year/seed zone

 Appropriate production and storage requirements and location for each species, e.g.,

 speculative production or wildland collection

 contract production or wildland collection

 nursery stock production

 seed storage location/capacity

The technical team includes geneticists to help inform number of collections required,seed 

source pooling, and selection of sources in consideration of climate change, seed growers, 

agronomists, and nurserymen to review target species for production feasibility.

The final task will be to refine and provide the requisite contracting tools via already established 

BLM and USFS Oregon models (Erickson 2005; Ferriel 2015; Kendig 2015) including IDIQ

(Indefinite-Delivery, Indefinite-Quantity) contracts with growers to expand Great Basin-wide 

coordinated plant increase.

To date the BLM Vale and Burns Districts are working with Eastern Oregon Stewardship 

Services to pilot this model for development of Greater Sage-Grouse habitat seed mixes(Figure 

1). In addition, BLM and GBNPP have partnered with the Fire Science Delivery Exchange to 

develop a restoration webinar series titled “The Right Seed in the Right Place at the Right Time -

Tools for Sustainable Restoration.” Series topics include seed zones, seed production,

restoration equipment, seed mixes, climate, and use of plant container stock to augment wildland 

seedings.

The past, current, and future applied research that has been spearheaded by GBNPP researchers 

has set a critical foundation and example for the Draft National Seed Strategy, a multi-agency 

and partner effort that outlines the “coordinated federal, state, and non-governmental 

organizations’ response to the need to provide an adequate supply of appropriate seed and plant 

materials for ecosystem and land health conservation, restoration, and rehabilitation” (Draft 

BLM IM 2015). Via the GBNPP and on-going development of technical teams comprised of 

subject matter experts, an increased breadth, coordination, and deployment of native plant 

materials across the Great Basin can be achieved.
72

Figure 1. Vale BLM provisional seed zones for sage-grouse forb seed mix 

(Fritts 2015), based on Bower et al. (2014).
73

References

Bower, Andrew D.; St. Clair, J. Bradley; Erickson, Vicky. 2014. Generalized provisionalseed 

zones for native plants. Ecological Applications 24: 913–919.

Chambers, Jeanne C.; Pyke, David A.; Maestas, Jeremy D.; Pellant, Mike; Boyd, Chad S.; 

Campbell, Steven B.; Espinosa, Shawn; Havlina, Douglas W.; Mayer, Kenneth E.; Wuenschel, 

Amarina. 2014. Using resistance and resilience concepts to reduce impacts of invasive annual 

grasses and altered fire regimes on the sagebrush ecosystem and greater sage-grouse: a strategic 

multi-scale approach. Gen. Tech. Rep. RMRS-GTR-326. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 73 p.

Erikson, Vicky. 2005. Management of regional native seed production contract; 2005 

Accomplishments. [Available online at

http://www.fs.fed.us/wildflowers/Native_Plant_Materials/documents/npmreports/fy2005/R6/R6_

seed_contract.pdf. Accessed 2/11/2015].

Kendig, Doug. 2015. [Personal communication]. BLM Medford District Botanist. 

Ferriel, Roger. 2015. [Personal Communication]. BLM Baker City Field Office.

Fritts, Susan.2015. [Personal Communication]. BLM Vale District Botanist. 

Fritts, Susan. 2015. [Email ]. January 14. BLM Vale District Botanist.

Shaw, Nancy L.; Lambert, Scott M.; DeBolt, Ann M.; Pellant, Mike 2005. Increasing native forb 

seed supplies for the Great Basin. In: Dumroese, R. K.; Riley, L. E.; Landis, T. D. [tech. 

coords.]. 2005. National proceedings: Forest and Conservation Nursery Associations—2004; 

2004 July 12–15; Charleston, NC; and 2004 July 26–29; Medford, OR. Proc. RMRS-P-35. Fort 

Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 

p. 94-102.

USDI. 2015. Secretarial Order 3336 [Available at

http://www.doi.gov/news/download/upload/Final-Rangeland-SO-greater.pdf]. Accessed 

2/11/2015.

USDI BLM. 2015. Draft Interagency Seed Strategy. [Available at

http://web.blm.gov/internal/wo-500/directives/dir-15/im2015-043.html].

USDA FS.2015. Western Wildland Environmental Threat Assessment Center. Prineville, OR. 

[Available at http://www.fs.fed.us/wwetac]. Accessed 2/10/2015.

Youtie, Berta. 2014. Coordination of Great Basin Native Plant Project Seed Increase and Eastern 

Oregon/Northern Nevada Seed Collections, p. 161-168. In: Kilkenny, Francis; Shaw, Nancy; 

Gucker, Corey. 2014. Great Basin Native Plant Project: 2013 Progress Report. Boise, ID: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station.
74

Project Title Developing Protocols for Maximizing Establishment of 

Great Basin Legume Species

Project Agreement No. 11-IA-11221632-007

Principal Investigators and Contact Information

Douglas A. Johnson, Plant Physiologist 

USDA-Agricultural Research Service 

Forage and Range Research Lab

Utah State University 

Logan, UT 84322-6300

435.797.3067, FAX 435.797.3075

doug.johnson@ars.usda.gov

B. Shaun Bushman, Research Geneticist 

USDA-Agricultural Research Service 

Forage and Range Research Lab

Utah State University 

Logan, UT 84322-6300

435.797.2901, FAX 435.797.3075

shaun.bushman@ars.usda.gov

Project Description

Rapid, uniform germination and initial stand establishment of legumes are frequently limited by 

hard-seededness. As part of this project, we previously conducted greenhouse experiments that 

determined how depth of planting and seed scarification treatments (soaking for 5 minutes in 

concentrated sulfuric acid or mechanical scarification by 120-grit sandpaper) affected 

germination and emergence of western prairie clover (Dalea ornata), basalt milkvetch 

(Astragalus filipes), and Searls’ prairie clover (Dalea searlsiae). These studies showed 

substantial improvements in germination and seedling emergence of western prairie clover and 

Searls’ prairie clover, but only marginal improvements for basalt milkvetch (Bushman et al., 

under review). For basalt milkvetch, recent experiments where seeds were imbibed, scarified 

seeds in moist sand resulted in over 90% germination within 7 days afterseeding.

In a second study, we planted scarified and non-treated seeds of western prairie clover and basalt 

milkvetch at three dryland sites in Oregon. The purpose was to determine an optimal time and 

seed treatment for rangeland stand establishment of these species. Seeded plots were planted for 

3 years, with each planting including a fall and spring seeding. The fall 2011/spring 2012 

occurred at the USFS/OSU Powell Butte farm and the river bottoms at Clarno, Oregon. The fall 

2012/spring 2013 plantings occurred at the Powell Butte farm, an upland site near Clarno, and 

the OSU Malheur Experiment Station in Ontario, OR. The third planting occurred at those same 

three sites during fall 2014, and soon to be spring 2015. These plantings are in coordination with 

Matt Horning of the USFS and Clinton Shock of Oregon State University.
75

Results from the first year of data from these studies indicated that early spring plantings of 

scarified western prairie clover seed were the most successful (Figure 1) while scarified basalt 

milkvetch seed established best in fall seedings (Figure 2). The second year of seeding resulted 

in no establishment due to a lack of precipitation from January through June of 2013; however

both scarified and non-treated seed of basalt milkvetch established the following year at a low 

percentage.

Figure 1. Seedling emergence and establishment at Powell Butte and *Clarno river bottom 

locations after the fall 2011/spring 2012 planting. Data collected July 2012. Majestic (AS) is 

acid-scarified seed of Majestic D. ornata gemrplasm, and Majestic(N) is non-treated seed.

Figure 2. Seedling emergence and establishment at Powell Butte and Clarno river bottom 

locations after the fall 2011/spring 2012 planting. Data collected July 2012. NBR-1(AS) 

corresponds to acid scarified NBR-1 germplasm of basalt milkvetch. NBR-1(N) is non-treated 

seed of NBR-1.

#

g

erm of

2

5

0

#

g

erm of

2

5

0
76

Research with Utah trefoil (Lotus utahensis)

Seed collections were made of ecotypes of Utah trefoil (a Great Basin leguminousspecies) 

throughout its range of distribution in southern Utah, southern Nevada, and northern Arizona 

during 2012 and 2013. Forage samples are currently being analyzed for tannin content.

Replicated common gardens of 19 Utah trefoil collections were established at three sites in 

northern Utah and will be evaluated for various morphological and physiological characteristics 

during 2015. Jason Stettler (previous biologist with the Utah Division of Wildlife Resources at 

Ephraim, UT) is pursuing his M. S. degree on this project at Utah State University.

Publications

Bushman, B. S.; Johnson, D. A.; Connors, K. J.; Jones, T. A. Germination and seedling 

emergence of three western North American rangeland legumes. Under Review, Rangeland 

Ecology and Management.

Johnson, D. A.; Bushman, B. S.; Jones, T. A.; Bhattarai, K. 2013. Using common gardens and 

AFLP analyses to identify metapopulations of indigenous plant materials for rangeland 

revegetation in western USA. In: Michalk, D. L.; Millar, G. D.; Badgery, W. B.; Broadfoot, K. 

M., eds. Revitalizing grasslands to sustain our communities. Orange, Australia: New South 

Wales Department of Primary Industry: 371-372.

Cane, J. H.; Johnson, C. D.; Napoles, J. R.; Johnson, D. A.; Hammon, R. 2013. Seed-feeding 

beetles (Bruchinae, Curculionidae, Brentidae) from legumes (Dalea ornata, Astragalus filipes) 

and other forbs needed for restoring rangelands of the Intermountain West. Western North 

American Naturalist 73: 477-484.

Presentations

Johnson, D. A.; Connors, K. J.; Bushman, B. S.; Jones, T. A. 2014. Lotus utahensis: Southern Great 

Basin legume for possible use in rangeland revegetation. Society for Range Management Annual 

Meeting; 2014 February; Orlando, FL.

Management Applications and Seed Production Guidelines

Seed producers will benefit by knowing how to obtain stands of these North American legume 

species for seed production. This information will provide land managers with informationto 

diversify their rangeland revegetation projects.

Products

 Germplasm releases for basalt milkvetch, western prairie clover, and Searls’ prairie clover.

 Seed production guidelines and planting guides.
77

Project Title Native Forb Increase and Research at the Great Basin

Research Center

Project Agreement No. 12-JV-11221632-050 and 11-JV-11221632-009

Principal Investigators and Contact Information

Kevin Gunnell, Range Trend Project Leader 

Utah Division of Wildlife Resources

Great Basin Research Center 

494 West 100 South

Ephraim, Utah 84627

435.283.4441, FAX 435.283.2034

kevingunnell@utah.gov

Jason Stettler, Native Forb Biologist 

Utah Division of Wildlife Resources 

Great Basin Research Center

494 West 100 South

Ephraim, Utah 84627

435.283.4441, FAX 435.283.2034

jasonstettler@utah.gov

Project Description

Cushion buckwheat (Eriogonum ovalifolium) common garden study.

Objectives

Analyze seed production of multiple native collected accessions of cushion buckwheat 

(Eriogonum ovalifolium) and G1 production seed from those accessions to test if seed production 

is a heritable trait. Evaluate drought tolerance of the same accessions with a common garden 

planted on the Desert Experimental Range.

Methods

In the fall of 2013 a common garden of the top 11 producing accessions (Table 1) from a prior 

common garden study, along with the first generation daughter plants (G1) of the same 

accessions, were planted on our Ephraim, UT farm and the Desert Experimental Range (DER) 

located in Millard County, UT. Ten plants of each accession and generation were planted in 

randomized blocks replicated four times per site. Seedlings were planted into weed barrier fabric 

and thoroughly watered in at the time of transplanting to minimize the effects of transplanting.

No supplemental water was given after the initial transplant. Mortality, seed production,seed 

size, and seed viability will be monitored on the Ephraim study site, and mortality will be 

measured on the DER site.
78

Table 1. Cushion buckwheat (Eriogonum ovalifolium) germplasm sources

Lot # Site Ecoregion

EROV-U11-2002 Sand Pass 13k Lahontan Sagebrush Slopes

EROV-U13-2002 Toano 13q Carbonate Woodland Zone

EROV-U20-2005 Crittenden Res. 13r Central Nevada High Valleys

EROV-U23-2005 Crittenden Res. 80a Dissected High Lava Plateau

EROV-U27-2005 Newcastle 13c Sagebrush Basins and Slopes

EROV-U28-2005 Jungo Road 13z Upper Lahontan Basin

EROV-U29-2005 Cobre 80a Dissected High Lava Plateau

EROV-U30-2004 Hwy 26 mm 280-281 12g Eastern Snake River Basalt Plains

EROV-U7-2002 Pequop Foothills 13p Carbonate Sagebrush Valleys

EROV-U8-2002 North Battle Mountain 13m Upper Humboldt Plains

EROV-U9-2002 Adobe Hill 13m Upper Humboldt Plains

Future Plans

Measurements will continue to be taken through the 2015 growing season. Analysis and 

publication are anticipated in 2016.

Project Description

Lupine species iron deficiency chlorosis (IDC) study.

Objectives

Identify a practical and effective method of treating iron deficiency chlorosis (IDC) forimproved 

establishment and seed production in multiple lupine species.

Methods

In the fall of 2011 two common gardens (randomized complete block design) were planted at our 

Fountain Green and Snow Field Station research farms to study the effects of iron fertilizer 

treatments. The treatments of EDDHA chelated iron fertilizer were: seed applied iron, foliar 

applied iron, a combined treatment of seed and foliar applied iron, and a control group without

any fertilizer application. These were applied to three lupine species: longspur lupine (L. 

arbustus), hairy bigleaf lupine (L. prunophilus), and silky lupine (L. sericeus). Measurements of 

emergence, establishment, chlorophyll content, plant height, and seed production were collected 

in 2012 and 2013, and plant height and seed production were collected in 2014.

Results and Discussion

Results from 2012 and 2013 were published in Kilkenny et al. (2014). Preliminary results from 

the 2014 data collection are presented here. Seed was only harvested on the Snow Field Station 

in 2014 due to an insect infestation on the Fountain Green farm that destroyed the seed inside the 

pods. Seed yield of silky lupine was significantly lower in 2014 than prior years on the Snow 

Field Station. This too may have been due to predation from insects. There remained no 

significant difference in yield of silky lupine among the treatments* (Figure 1). There was no
79

significant difference in plant height between treatments or treatment*farm in 2014. Similar to 

seed yield, plant height was significantly smaller in 2014 than in 2013 (Figure 2). Preliminary 

results indicate that iron treatments had limited impact on seed production and vegetation 

response.

Figure 1. Seed yield of silky lupine (Lupinus sericeus) on now 

Field Station by treatment for 2013 and 2014.

Figure 2. Mean plant height of four lupine species on the 

Fountain Green (FG) and Snow Field Station (SFS) for 

2013 and 2014.
80

Future Plans

Final analysis and publication are anticipated in 2015.

Project Description

Wildland collection native seed increase.

Objectives

Increase seed from wildland collections to support further research and increase distribution of 

native seed to commercial growers. Develop and test methods to increase germination and 

establishment in agronomic settings.

Methods

Work will be done in conjunction with the Provo Shrub Sciences Lab wildland seed collection 

project. Selected species and accessions of wildland collected seed will be planted on Utah 

Division of Wildlife Resources (UDWR) farms in Fountain Green, UT or Ephraim, UT for the 

purpose of seed increase. Seed will be either direct seeded into fields or propagated in the 

greenhouse for plug transplanting depending on the amount of seed available.

Production fields will be maintained and harvested to maximize seed production of native 

species. When opportunities arise, development of propagation protocols and research for 

increased establishment and production will be conducted.

Results

Seed from 12 species with 58 accessions were harvested on the UDWR farms in 2014 (Table 2). 

Two species with 10 accessions were direct seeded into new increase beds in the fall of 2014, 

and a further 14 accessions of those same species will be propagated in the greenhouse and 

seedlings transplanted in the spring of 2015 (Table 3).

Table 2. Seeded species harvested from UDRW farms in 2014.

Code Species # of Accessions Location

ASFI Astragalus filipes 1 Ephraim

CRAC2 Crepis acuminata 3 Ft. Green

CRIN4 Crepis intermedia 3 Ft. Green

CRYPT Cryptantha sp. 1 Ft. Green

DAOR2 Dalea ornata 1 Ephraim

ENNUN Enceliopsis nudicaulis 3 Ft. Green

IPAG Ipomopsis aggregata 3 Ft. Green

LECI4 Leymus cinereus 31* Ephraim

LILE3 Linum lewisii 6 Ephraim

LUAR6 Lupinus arbustus 1 Ephraim

LUSES2 Lupinus sericeus 1 Ephraim

PEPA6 Penstemon pachyphyllus 4 Ft. Green

*Foundation source of UDWR Tetra Great Basin Wildrye selected release. 

Accessions are pooled when harvested.
81

Table 3. Species sown on UDWR farms in 2014-2015.

Code Species # of Accessions Location

HEMUN Heliomeris multiflora nevadensis 5 Ft. Green

PEPA6 Penstemon pachyphyllus 19 Ft. Green

Future Plans

Maintenance, harvest and distribution of seed from established increase plots will continue. 

Expansion of increase plots will proceed with the addition of new species and accessions as 

deemed necessary. Development of propagation protocols and research for increased 

establishment and production will be conducted as the opportunity arises.

Project Description

N-sulate® Fabric assessment on germination and establishment study.

Objectives

Determine the effects of N-sulate® fabric on germination and establishment of native forb 

species in a wildland setting.

Methods

This study was implemented in 2009 and 2010. Four sites were selected in Utah, two in Tooele 

County, one in Sanpete County, and one in Carbon County. Each study site consists of two years 

replication with five randomized complete blocks of treatments in each year. Treatments are two 

seed mixes (Table 4) and two mulch treatments covered with N-sulate® fabric, and an uncovered 

control. The study sites were prepared by mechanically removing sagebrush and other vegetation 

with two passes of a pipe harrow. The seed mixes were sown with rice hulls using a handoperated seed broadcaster at a rate of 20-25 seeds ft2

, depending on the species. The plots were 

then compacted with a Brillion packer wheel (Brillion Farm Equipment, Brillion, WI) to ensure 

good seed to soil contact. Germination and establishment data were recorded one, two, and five

years post treatment.

Results and Discussion

The five year data has not been collected for the 2010 planting yet, so only results from the first 

two years are reported here. When looking at total mean density of seeded species across all four 

study sites, we see differing responses over time. Germination was higher for species in both 

mixes in the covered treatment in the first growing season, but establishment of species by the 

second growing season of both seed mixes was much lower and there was no substantial 

difference of mean density in covered and uncovered treatments (Figure 3).

When response is looked at across study sites, germination is higher than the unseeded control 

for both mixes on all sites. Density is higher in the covered treatments for both mixes acrossthe 

treatment types, but this is most pronounced in the lower precipitation sites of Hatch Ranch and 

Lookout Pass (Figure 4).
82

Table 4. Forb island study seed mixes.

Seed Mix 1

USDA Plant Code Scientific Name Accession No. Seeded Rate (g)

BASA3 Balsamorhiza sagittata Basa-I-UT-R2 17

CLSE Cleome serrulata CSE-107 2. 8

HEBO Hedysarum boreale 2008. 0366 21. 3

LIPE2 Linum perenne ‘Appar’ NBS-RR8-APP-2 3. 12

LUAR3 Lupinus argenteus 122 63. 8

PEPA6 Penstemon pachyphyllus PEPA6-B5-2009 4. 3

POFE Poa fendleriana 2006. 04. 08 1. 1

SPGR2 Sphaeralcea grossulariifolia 287 2. 3

Seed Mix 2

USDA Plant Code Scientific Name Accession No. Seeded Rate (g)

AGGR2 Agoseris grandiflora AGGLG 28080 LPN 4. 5

AGHE2 Agoseris heterophylla SI-1 1. 9

ARMU Argemone munita ARMU-U1-2009 8. 7

HEMUN Heliomeris multiflora nevadensis HEMUN-B4-2009 1. 9

LONU2 Lomatium nudicaule Big Butte source 28. 4

NIAT Nicotiana attenuata Mud Springs source 0. 4

THMI5 Thelypodium milleflorum THMI5-P2-2009 0. 4
83

Figure 3. Total density across sites of seeded species for the first and 

second growing season.

Future Plans

Data collection will occur one more time in the summer of 2015 to assess the 2010 treatment for 

the full 5 years. After data collection is complete, analysis and publication is anticipated for 

2016.

Seed Mix 1

80

70

60

50

40

30

20

10

0

Uncovered Covered Uncovered Covered

SPGR2

LUAR3 

CLSE 

BASA3 

POFE 

HEBO 

PEPA6

LIPE2

1stGrowing Season 2ndGrowing Season

Seed Mix 2

80

70

60

50

40

30

20

10

0

HEMUN

THMI5

Agoseris 

NIAT 

ARMU

LONU2

Uncovered Covered Uncovered Covered

1stGrowing Season 2ndGrowing Season

Plants/m.sq.

Plants/m.sq.
84

Figure 4. Density of perennial species by treatment type and study location.

Project Description

Forb seeding method study using Truax drill, Lawson aerator, Dixie pipe harrow and Ely chain.

Objectives

Currently, seed mixes used for restoration generally do not include native forb species, mainly 

due to their high cost. Because these species have not been included in past restoration efforts 

there is a lack of information about planting and establishing native forbs in wildland settings. As 

these native forbs become more available on the market, the lower costs will facilitate land 

manager’s ability to include these species in seed mixes used for restoration. This creates a need 

to determine the best methods of establishing native forbs in wildland settings. The main 

objective of this study is to evaluate native forb establishment in Wyoming big sagebrush 

communities using four different pieces of planting equipment: Truax drill, Lawson aerator, 

Dixie pipe harrow, and Ely chain.

Methods

This study was established on two sites (referred to as North and South) in Tooele County, Utah. 

Treatments occurred in the fall of 2008 and 2009. The seed mix used for the study consisted of 

only native forbs (Table 5). Each study site consists of two years replication with two 

randomized complete blocks of disturbance treatments in each year. Cover was collected for all 

species and density for perennial species. Measurements were done in post treatment years one, 

two, and six.

Future Plans

Results from 2009 and 2010 were published in Shaw and Pellant (2011). Final sampling will 

occur in the summer of 2015. Final analysis and publication are anticipated in 2016.

1stGrowing Season by Site

60

50

40

30

20

10

0

Uncovered

Covered

Fountain Gordon Creek Hatch Ranch Lookout Pass

Green

D

e

nsity/

m. sq.

Mix 1

Mix 2 

C

ontrol 

Mix 1

Mix 2 

C

ontrol 

Mix 1

Mix 2 

C

ontrol 

Mix 1

Mix 2

C

ontrol
85

Table 5. Seed mix used on the project.

Species

Bulk 

lbs/acre)

PLS

lbs/acre Seeds/ft2

Munro Globemallow (Sphaeralcea munroana) 0.5 0.38 4.32

Blue Flax (Linum lewisii) 1.0 0.90 6.04

Northern Sweetvetch (Hedysarum boreale) 1.0 0.77 0.60

Silvery Lupine (Lupinus argenteus) 1.5 0.55 0.27

Palmer Penstemon (Penstemon palmeri) 0.5 0.39 4.86

Utah Milkvetch (Astragalus utahensis) 0.6 0.39 2.03

Arrowleaf Balsamroot (Balsamorhiza sagittata) 2.0 1.81 2.28

Firecracker Penstemon (Penstemon eatonii) 0.29 0.23 3.18

Tapertip Hawksbeard (Crepis acuminata) 0.16 0.10 0.48

Total 7.55 6.02 24.04

Presentations

Stettler, J. 2014. Lupine cultivation and ecological niche modeling. Great Basin Native Plant 

Project Annual Meeting; 2014 March 17-18; Boise, ID.

References

Kilkenny, Francis; Shaw, Nancy; Gucker, Corey. 2014. Great Basin Native Plant Project: 2013 

Progress Report. Boise, ID: U. S. Department of Agriculture Forest Service, Rocky Mountain 

Research Station: 74-93.

Shaw, Nancy; Pellant, Mike. 2011. Great Basin Native Plant Selection and Increase Project: 

2010 Progress Report. Boise, ID: U. S. Department of Agriculture Forest Service, Rocky 

Mountain Research Station: 167-172.
86

Project Title Plant Material Work at the Provo Shrub Sciences Lab

Project Agreement No. 13-IA-11221632-161

Principal Investigators and Contact Information

Scott Jensen, Botanist

Rocky Mountain Research Station

Grassland, Shrubland and Desert Ecosystem Program 

735 N. 500 E.

Provo, UT 84606-1865

801.356.5124, FAX 801.375.6968

sljensen@fs.fed.us

Project Descriptions

Wildland Seed Collection

Seed collection efforts in 2014 (Table 1) were focused on identifying and harvesting new 

populations of thickleaf penstemon (Penstemon pachyphyllus) and recollecting basin wildrye 

(Leymus cinereus) for an ongoing study. A cheatgrass (Bromus tectorum) die-off afforded an 

opportunity for a bulk collection of Nevada bluegrass (Poa secunda).

Table 1. Species and number of sources collected in 2013.

Family Name Common Name 2014 Collections

Poaceae Poa secunda Nevada bluegrass 1

Poaceae Leymus cinereus Basin wildrye 27

Scrophulariaceae Penstemon pachyphyllus thickleaf penstemon 17

Seed Distributions for Stock Seed and Commercial Seed Production

In cooperation with the Utah Division of Wildlife Resources and the Great Basin Research 

Center (GBRC), 19 sources of thickleaf penstemon from provisional seed zone 15 - 20 Deg. F./ 3

– 6 (Bower et al. 2014) and five sources of Nevada showy goldeneye (Heliomeris multiflora var. 

nevadensis) from provisional seed zone 15 - 20 Deg. F./ 6 – 12 (Bower et al. 2014) were either 

direct seeded or seeded into plugs for grow out and transplant at the Fountain Green farm facility 

(Table 2). Future seed harvests from these beds will serve as stock seed for these species/zone 

combinations for distribution to commercial producers.

A small number of private industry seed producers continue to show interest in commercial 

production of Great Basin native forbs and grasses. In 2014, eight species representing stock 

seed for 12 species/zone combinations were distributed to private commercial seed producers 

(Table 3).
87

Table 2. Species and sources planted for stock seed production at Fountain Green, Utah.

Species

#

Sources Method Provisional Seed Zone

Penstemon pachyphyllus 7 S 15 - 20 Deg. F. / 3 - 6

Penstemon pachyphyllus 12 P 15 - 20 Deg. F. / 3 - 6

Heliomeris multiflora var. nevadensis 3 S 15 - 20 Deg. F. / 6 - 12

Heliomeris multiflora var. nevadensis 2 P 15 - 20 Deg. F. / 6 - 12

Totals 2 24 2

S - Direct seed, P - Transplant plugs

Table 3. Lots distributed to private industry for commercial production.

Species

#

Sources Lots

Weight 

(lbs) Provisional Seed Zone

Heliomeris multiflora var. nevadensis 3 15 14.6 15 - 20 Deg. F. / 6 - 12

Heliomeris multiflora var. nevadensis 2 2 2.1 20 - 25 Deg. F. / 6 - 12

Lomatium nudicaule 1 1 6.25 15 - 20 Deg. F. / 6 - 12

Agoseris grandiflora 1 2 3.39 20 - 25 Deg. F. / 3 - 6

Erigeron speciosus 1 1 0.1 15 - 20 Deg. F. / 3 - 6

Cleome lutea 1 1 0.79 15 - 20 Deg. F. / 6 - 12

Sphaeralcea grossulariifolia 1 1 7.5 15 - 20 Deg. F. / 6 - 12

Sphaeralcea munroana 1 1 0.21 10 - 15 Deg. F. / 6 - 12

Leymus cinereus 9 9 9 10 - 15 Deg. F. / 6 – 12

Leymus cinereus 5 5 2 15 - 20 Deg. F. / 3 – 6

Leymus cinereus 8 8 11.47 15 - 20 Deg. F. / 6 - 12

Leymus cinereus 5 5 3.33 20 - 25 Deg. F. / 6 – 12

Totals 8 38 51 60.74 6
88

Evaluating Emergence from Increasing Planting Depth for 20 Native Forbs

Objectives

Evaluate the effect of planting depth on seedling emergence for 20 native forb species (Table 4) 

planted in loam-textured soils at three locations.

Methods

In Fall 2013 and 2014 each species was sown at four depths in a randomized block design with 

10 blocks per site. Seeding rates were adjusted for viability and purity to 25 PLS/ft. Five blocks 

were covered with N-sulate fabric and five were left uncovered. The study was replicated at three 

sites, Orovada Nevada; Wells, Nevada; and Fountain Green, Utah. Emergence and short term 

persistence data will be collected 2014 - 2016.

Table 4. Forbs seeded to evaluate planting depth on seedling emergence.

Family Name Common Name

Asteraceae Achillea millefolium Common yarrow

Asteraceae Agoseris grandiflora Bigflower agoseris

Asteraceae Agoseris heterophylla Annual agoseris

Caryophyllaceae Arenaria macradenia ssp. ferrisiae Ferris' sandwort

Fabaceae Astragalus filipes Basalt milk-vetch

Asteraceae Balsamorhiza hookeri Hooker's balsamroot

Asteraceae Enceliopsis nudicaulis Nakedstem sunray

Polygonaceae Eriogonum umbellatum Sulphur-flower buckwheat

Asteraceae Heliomeris multiflora var. nevadensis Nevada showy goldeneye

Polemoniaceae Ipomopsis aggregata Scarlet gilia

Apiaceae Lomatium nudicaule Barestem biscuitroot

Fabaceae Lupinus argenteus Silvery lupine

Fabaceae Lupinus prunophilus Hairy big leaf lupine

Asteraceae Machaeranthera canescens Hoary tansy-aster

ScrophulariaceaePenstemon acuminatus Sharpleaf penstemon

ScrophulariaceaePenstemon pachyphyllus Thickleaf penstemon

ScrophulariaceaePenstemon speciosus Royal penstemon

Malvaceae Sphaeralcea grossulariifolia Gooseberryleaf globemallow

Malvaceae Sphaeralcea munroana Munro's globemallow

Brassicaceae Stanleya pinnata var. integrifolia Golden prince's-plume

Evaluating Methods of Pairing Seed Sources to Restoration Sites Using Basin Wildrye (Leymus 

cinereus)

Objectives

Basin wildrye is a common grass native to western North America. Its distribution extends from 

New Mexico on the south, to Saskatchewan on the north, then west across all states and 

provinces to the Pacific Coast. Across its distribution is an enormous array of climactic and 

ecological variation that has likely led to the development of localized adaptions between 

spatially divergent populations. In recent years plant material development strategies have
89

emphasized methods that focus on local adaptation or matching adaptive variation to climate. 

The products of these genecological studies are maps that display similarities in plant 

performance metrics and climate parameters across a species distribution.

For a number of important restoration species genecological work is underway. However, this 

type of work may continue for decades and will likely never be completed for many species. 

Data must be developed on a species by species basis requiring several years of study adding to 

expenses. Where genecological work is lacking, surrogate approaches have been suggested.

In this study we evaluate generalized provisional seed zones (GPSZ) as a method of matching 

seed sources to restoration sites. Because this locally adapted approach of matching seed supplies 

to restoration sites is relatively new in large scale restoration efforts and in direct competition 

with the common cultivar model, we include basin wildrye cultivars in the trial. This permits a 

comparison of establishment success between primary and secondary restoration gene pool 

materials at sites with varying degrees of divergence from historic norms.

Methods

In excess of 105 known populations of basin wildrye were mapped by GPSZ. Approximately 

90% of these populations fell within four provisional seed zones and none of the poorly 

represented zones had a proportional occurrence of more than 5%. Next we located test sites in

each of the four well represented zones at Fountain Green, Utah; Spanish Fork, Utah; Nephi, 

Utah; and Orovada, Nevada. Seed from four basin wildrye cultivars Magnar, Trailhead, 

Continental and Tetra was procured through local vendors and 27 wildland sources representing 

four provisional seed zones were collected from native stands in 2013 and 2014. Individual 

sources were sampled for seed weight, viability, and ploidy; then seeded at 25 PLS/ft in a 

randomized block design with ten blocks per site. Five blocks were covered with N-sulate fabric 

and five blocks were left uncovered. Plots were seeded in 2013 and 2014. Emergence and 

persistence data will be collected 2014 - 2016.

References

Bower, A. D.; St. Clair, J. B.; Erickson, V. 2014. Generalized provisional seed zones for native 

plants. Ecological Applications 24(5): 913-919.
90

Project Title Aberdeen Plant Materials Center Report of Activities

Project Agreement No. 11-1A-11221632-004

Principal Investigators and Contact Information

Derek Tilley, PMC Manager

USDA NRCS, Aberdeen Plant Materials Center 

PO Box 296, Aberdeen, ID 83210

208.397.4133, FAX 208.397.3104

Derek.Tilley@id.usda.gov

Project Description

The Aberdeen Plant Materials Center (PMC) is involved with plant collection, selection and 

development, and seed and plant production for improving degraded or disturbed lands. The 

Aberdeen PMC also transfers garnered plant species production, establishment, and management 

knowledge to others.

Plant Guides

The Aberdeen PMC is gathering information on selected plant species to create Natural 

Resources Conservation Service (NRCS) plant guides. Plant guides offer the most recent 

information on plant establishment, management and seed and plant production information. 

General information for the species can also be found in our plant guides including information 

on potential uses, ethnobotanical significance, adaptation, pests, and potential problems. In 2014 

plant guides were completed or revised for Rocky Mountain Penstemon (Penstemon strictus) and 

Hoary tansyaster (Machaeranthera canescens). Plant guides are available at the PLANTS 

database, www.plants.usda.gov, and at the Aberdeen Plant Materials Center website: 

www.id.nrcs.usda.gov/programs/plant.html.

Plant Materials Development

Hoary tansyaster

Nine accessions of hoary tansyaster (Machaeranthera canescens) were evaluated from 2009 

through 2011 for establishment, plant growth, and seed production. Accession 9076670 had the 

best establishment and stands for 2009 and 2010. It also had the best rated vigor in 2010 and the 

tallest plants in the overall study. The population for 9076670 is located near the St. Anthony 

Sand Dunes in Fremont County, Idaho at 5,000 ft. elevation. The site has sandy soils and 

supports a bitterbrush, Indian ricegrass, rabbitbrush, and scurfpea plant community. The location 

receives between 10 and 15 inches of mean annual precipitation. In 2014, accession 9076670 

was officially released as Amethyst Germplasm hoary tansyaster.

G1 and G2 seed of Amethyst Germplasm hoary tansyaster will be maintained by the USDA 

Natural Resources Conservation Service, Aberdeen Plant Materials Center. Seed through the G5 

generation will be eligible for certification. G1 and G2 seed will be made available to 

commercial growers for distribution by the University of Idaho Foundation Seed Program and 

Utah Crop Improvement Association. Small quantities of seed will be provided to researchers by 

request to the Plant Materials Center.
91

Wyeth or whorled buckwheat

Thirty-nine accessions of Wyeth and sulphur-flower buckwheat (Eriogonum heracleoides and E. 

umbellatum) were compared in a common garden study from 2007 through 2011. Accession 

9076546 Wyeth buckwheat showed good establishment, seed production, and longevity 

compared with the other accessions.

Initial seed increase of accession 9076546 is ongoing. Release documentation is being 

developed, and official release will occur once the Aberdeen PMC has produced a sufficient 

amount of early generation seed.

Publications

St. John, Loren (editor). 2014. Plant Guide for Rocky Mountain penstemon (Penstemon strictus). 

Aberdeen, ID: U. S. Department of Agriculture, Natural Resources Conservation Service, 

Aberdeen Plant Materials Center. Revised January, 2014.

Tilley, Derek; Ogle, Dan; St. John, Loren. 2014. Plant Guide for hoary tansyaster 

(Machaeranthera canescens). Aberdeen, ID: U. S. Department of Agriculture, Natural Resources 

Conservation Service, Aberdeen Plant Materials Center. 4 p.

Tilley, Derek. 2014. Release Brochure for Amethyst Germplasm Hoary Tansyaster 

(Machaeranthera canescens). Aberdeen, ID: U. S. Department of Agriculture, Natural Resources 

Conservation Service, Aberdeen Plant Materials Center. 2 p.

Products

 Revised Plant Guides are available for Rocky Mountain penstemon and hoary tansyaster.

 Early generation certified seed of hoary tansyaster is in production, and allocations have 

been made to Utah Crop Improvement and Idaho Foundation Seed Program.

 Early generation certified seed of Wyeth buckwheat is being produced and will be 

available through Utah Crop Improvement and University of Idaho Foundation Seed 

Program when release is approved.
92

Project Title Bee Pollination and Breeding Biology Studies

Project Agreement No. 11-IA-11221632-010

Principal Investigators and Contact Information

James H. Cane, Research Entomologist 

USDA Agriculture Research Service 

Pollinating Insects Research Unit

Utah State University 

Logan, UT 84322-5310

435.797.3879, FAX 435.797.0461

Jim.Cane@ars.usda.gov

Project Description

Successful reproduction by flowering plants requires the confluence of several favorable factors. 

Some are intrinsic to the plant (e.g. age, nutritional status); others are external, either abiotic (e.g. 

rainfall, fire) or biotic (e.g. pollinators). The survival and performance of perennial sagebrushsteppe forbs after fire is an unknown with potentially great ramifications for pollinators(floral 

resources for reproduction). Seed choices (e.g. alfalfa) and management tactics may be needed 

to retain remnant pollinator communities while awaiting establishment of new seedlings of their 

floral hosts.

Methods

Native Bees in the Aftermath of Wildfire in the Great Basin

Opportunities are few to sample bee faunas in the weeks and months following large wildfires. 

In the aftermath of the huge Long Butte wildfire near Twin Falls (440 sq miles), we sampled 

native bee faunas at scattered remnant stands of sunflowers growing in ditches along gravel 

roads that served as fire lines some 7 miles from the burn perimeter. Comparable quantitative 

samples of bees were taken at coflowering sunflowers outside the burn. All but two of the bee 

species were non-social and soil nesting. These sunflower bee faunas largely survived the fire 

intact (but for one surface-nesting species). Females even continued to collect pollen for their 

nests. The fire had passed the sample sites at night, a time when female bees are in their nest 

(and thus safe underground). A manuscript has been submitted to the journal Biological 

Conservation entitled: “Direct effects of wildfire on a bee community: nesting habitus 

determines persistence.”

Fire Impact on Forb Survival

Changes in the frequency and intensity of fire in the sagebrush steppe have become key factors 

in degrading native plant communities in basin and foothill habitats. We know that many of the 

dominant Great Basin shrubs are consumed by fire and do not resprout well. Some forbs clearly 

survive some wildfires to thrive and bloom thereafter, but the observations are anecdotal and not 

related to fire attributes. Hence, we do not know which native forb species survive fires of a 

particular intensity.
93

Penstemon 

Eriogonum 

Astragalus 

Dalea 

Lomatium 

Sphaeralcea

To examine the individual plant’s response to wildfire, six common Great Basin perennial forb 

species were experimentally burned in 2012 at increasing intensities using a custom portable 

propane burner designed to mimic wildfire attributes. Six propane-fueled prescription treatments 

were used to mimic wildfire conditions, varying heat intensity and duration (Table 1). Survival 

data were collected along with a number of characteristics thought to be indicators important for 

bees in a post-fire environment (e.g. numbers of flowers, of flowering stalks and/or the lengths of 

racemes. Analyses are completed and manuscript writing is underway.

To reinforce our conclusions with field data, we sampled densities of perennial blooming 

Penstemon, Lupine, Lomatium and Balsamorhiza plants 8 months after the State Line Fire in the 

Samaria Mountains of southern Idaho. Obviously, these plants had to be survivors and not 

seedling recruits. Plants were larger, more vigorous and equally numerous within the fire 

perimeter compared with outside the edge in comparable soils, slopes and aspects. The general 

prediction from our forb burning treatment was upheld, that many Great Basin perennial forbs 

can and do survive wildfire to flower the following spring.

Table 1. Propane burn treatments used on native plants to mimic wildfires of increasing 

severity, and mortality of select burned forbs.

100

80

60

40

20

0

Increasing Fire Severity

Future plans

1) This year we expect to finish writing and submit manuscripts for the forb burning study and 

for the design, manufacture and use of the burner built for this application. A final manuscript 

will report analyses of the native bee faunas systematically sampled in and out of a 

chronosequence of large wildfires in the Great Basin sage steppe.

2) Manual pollination proved to be an intractable method for documenting the breeding biology 

of silverleaf phacelia (Phacelia hastata). We therefore grew plants from seed in conetainers. 

We have transplanted these to six separate locations in and around Logan. At each site, we 

transplanted one isolated P. hastata plant and, at a distance, a close-set trio of plants. The former 

should be primarily selfed by resident bees, whereas the trio should have considerable 

outcrossing. Mortality varied among sites. Dead plants were replaced, a few others flowered in

Mortality (percent)

Category Prescription Heat attained (

o

C)

Control No fuel applied Ambient

Very Low 7.5 psi, 25 sec 100

Low 7.5 psi, 50 sec 200

Medium 15 psi, 65 sec 300

High 15 psi, 175 sec 500

Very High 15 psi, 240 sec 600
94

this, their first year. In 2015, we will collect seeds per umbel of counted flowers to evaluate seed 

production with these naturally pollinated plants.

3) My colleagues and I in the Great basin Native Plant Project will be submitting a proposal to 

support a research project that will compare two methods for improved forb seedling 

establishment. We will enhance shallow soil moisture in early spring either by: 1) directed 

snowdrift accretion using custom snow fences, or 2) reduced spring evapotranspiration using 

floating row cover fabric. If successful, the technique could be used to establish replicate islands 

of mixed forbs at burn sites.

4) We will replicate our quantitative samples of native bees at wild stands of blue flax and 

western yarrow to evaluate a preliminary impression that these two workhorse forbs of sagesteppe restoration are relatively unattractive to native bees. Results will guide our 

recommendation about the value of these two easily grown forbs for bee conservation.

Publications

Cane, J. H.; Dunne, R. 2014. Generalist bees pollinate red-flowered Penstemon eatonii: refining 

the hummingbird pollination syndrome. American Midland Naturalist 171: 365-370.

Cane, J. H. 2014. The oligolectic bee Osmia brevis sonicates Penstemon flowers for pollen; a 

newly documented behavior for the Megachilidae. Apidologie 45: 675-684.

Presentations

Cane, J. Wild bees for Wyoming’s wild flowers. Wyoming Native Plant Society; 2014 June 21; 

Lander, WY. Invited keynote.

Cane, J. Floral guilds of native bees endure wildfire in the Great Basin. North America Congress 

for Conservation Biology; 2014, July 13-16; Missoula, MT.

Cane, J. Pollinators and agricultural seed production, consideration of pollinators in management 

and restoration planning. Society for Ecological Restoration Northwest/Great Basin Meeting; 

2014, October 6-11; Bend, OR. Invited symposium.

Management Applications and Seed Production Guidelines

A number of the common perennial forbs in sage steppe will survive late summer wildfires. 

Knowing what species were on a landscape prior to fire can guide forb seed selection for postfire seedings. Emphasis should be given to those forb species expected at the location but absent 

owing to earlier extirpation and not the recent fire.
95

Project Title Seed Production of Great Basin NativeForbs

Project No. 12-JV-11221632-066

Principal Investigators and Contact Information

Clinton C. Shock, Director/Professor 

Oregon State University

Malheur Experiment Station 

595 Onion Avenue

Ontario, OR 97914

541.889.2174, FAX 541.889.7831

clinton.shock@oregonstate.edu

Erik B. Feibert, Senior Faculty Research Assistant 

Oregon State University

Malheur Experiment Station 

595 Onion Avenue

Ontario, OR 97914

541.889.2174

erik.feibert@oregonstate.edu

Joel Felix, Associate Professor 

Oregon State University 

Malheur Experiment Station 

595 Onion Avenue

Ontario, OR 97914

541.889.2174, FAX 541.889.7831

joel.felix@oregonstate.edu

Nancy Shaw, Research Botanist (Emeritus) 

USFS Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4360, FAX 208.373.4391

nancylshaw@fs.fed.us

Francis Kilkenny, Research Biologist

USFS Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4376, FAX 208.373.4391

ffkilkenny@fs fed us
96

Project Description

A five part study regarding seed production of Great Basin native forbs using subsurfacedrip 

irrigation (SDI) for stable, efficient native forb seed production using small amounts of 

supplemental irrigation water, weed control, and seeding practices.

A. Direct Surface Seeding Strategies for Emergence of Native Plants in 2014

Clinton Shock, Erik Feibert, Alicia Rivera, and Lamont Saunders, Malheur Experiment Station, 

Oregon State University, Ontario, OR

Francis Kilkenny and Nancy Shaw, U.S. Forest Service, Rocky Mountain Research Station,

Boise, ID

Introduction

Seed of native plants is needed to restore rangelands of the Intermountain West. Reliable 

commercial seed production is desirable to make seed readily available. Direct seeding of native 

range plants has been generally problematic, especially for certain species. Fall planting is 

important for many species, because their seed requires a period of cold to break dormancy 

(vernalization). Fall planting of native seed has resulted in poor stands in some years at the 

Malheur Experiment Station. Loss of soil moisture, soil crusting, and bird damage are some 

detrimental factors hindering emergence of fall planted seed. Previous trials at the Malheur 

Experiment Station have examined seed pelleting, planting depth, and soil anti-crustants (Shock 

et al. 2010). Planting at depth with soil anti-crustant led to improved emergence compared to 

surface planting. Seed pelleting did not improve emergence. Despite some positive results, 

emergence was extremely poor for all treatments, due to soil crusting and bird damage.

In established native perennial fields at the Malheur Experiment Station and in rangeland areas 

we have observed prolific natural emergence from seed that falls on the soil surface and is 

covered by thin layers of organic debris. In 2011, 2012, and 2013, trials tested the effect of seven 

planting systems on surface planted seed (Table 1; Shock et al. 2012, 2013, 2014).

Row cover can be a protective barrier against soil desiccation and bird damage. Sawdust can 

mimic the protective effect of organic debris. Sand can help hold the seed in place. Seed 

treatment can protect the emerging seed from fungal pathogens that might cause seed 

decomposition or seedling damping off. Hydroseeding mulch could be a low cost replacement 

for row cover. The treatments did not test all possible combinations of factors, but tested 

combinations that might be likely to result in adequate stand establishment. In 2014, the seven 

planting systems were tested for emergence of 8 important species, 7 of which are native to 

Malheur County and surrounding rangelands and forestlands.

Materials and Methods

Seven species for which stand establishment has been problematic were chosen. An eighth 

species (Penstemon speciosus) was chosen as a check, because it has reliably produced good 

stands at Ontario. Seed weights for all species were determined. A portion of the seed was 

treated with a liquid mix of the fungicides Thiram® and Captan (10g Thiram, 10g Captan in 0.5 

liter of water). Seed weights of the treated seeds were determined after treatment. The seed

weights of untreated and treated seed were used to make seed packets containing approximately 

300 seeds each. The seed packets were assigned to one of seven treatments (Table 1). The trial 

was planted manually on November 26, 2013. The experimental design was a randomized
97

complete block with six replicates. Plots were 30-inch-wide by 5-ft-long beds. Two seed rows 

were planted on each bed.

Tetrazolium tests were conducted to determine seed viability of each species (Table 2). The 

tetrazolium results were used to correct the emergence data to emergence of viable seed.

After planting, the sawdust was applied in a narrow band over the seed row at 0.26 oz/ft of row 

(558 lb/acre). For the treatments receiving both sawdust and sand, the sand was applied at 0.65 

oz/ft of row (1,404 lb/acre) as a narrow band over the sawdust. Following planting, sawdust and 

sand applications, some of the beds were covered with row cover. The row cover (N-sulate, 

DeWitt Co. Inc., Sikeston, MO) covered four rows (two beds) and was applied with a 

mechanical plastic mulch layer.

For the hydro seeding mulch treatments, 12 lb of hydro seeding paper mulch (Premium Hydro 

seeding Mulch, Applegate Mulch, http://applegatemulch. com) was mixed in 50 gallons of water 

in a jet-agitated 50-gallon hydro seeder (Turbo Turf Technologies, Beaver Falls, PA). The

mulch was applied with the hydro seeder in a 3-cm band over the seed row.

On April 15 2014, the row cover was removed and emergence counts were made in each plot. 

Data were analyzed using analysis of variance (General Linear Models Procedure, NCSS, 

Kaysville, UT). Means separation was determined using a protected Fisher’s leastsignificant 

difference test at the 5 percent probability level, LSD (0. 05).

Results and Discussion

Overall, emergence in the trial was poor. December of 2013 and January of 2014 had 

precipitation that was about 50% lower than normal and could have contributed to the poor 

emergence. Emergence and stand of Phacelia linearis, Ligusticum porteri, and Ligusticum 

canbyi was poor at 10% or less (Table 3). Emergence and stand of Phacelia hastata, Heliomeris 

multiflora, and Penstemon speciosus was fair at 20% or less. There was no significant difference 

in stand between treatments for Phacelia linearis, Phacelia hastata, Heliomeris multiflora, and 

Ligusticum canbyi.

The row cover with sawdust and seed treatment resulted in higher stands than no row cover (bare 

ground) with sawdust and seed treatment for Machaeranthera canescens and Ligusticum porteri 

(Table 3). Sawdust added to the row cover and seed treatment only improved stands of 

Ligusticum porteri. Adding seed treatment to sawdust and row cover did not improve stands of 

any species. Adding sand to sawdust, seed treatment, and row cover increased stand for 

Penstemon speciosus.

The hydroseed mulch applicator is designed to have the seed mixed in the tank with the mulch 

and then the mixture is sprayed on the soil surface. Since it was not practical to mix the seed with 

the mulch, the application pressure had to be reduced enough to not displace the previously 

planted seed on the soil surface. The resulting mulch application was too thick and when dry 

resembled paper mâché, which was probably too hard of a barrier for seed emergence. Future 

adjustments of the thickness of the mulch mix and spray nozzle might allow for the application 

of mulch of the right consistency. Stand with hydroseed mulch and seed treatment waslower
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than with row cover and seed treatment for Machaeranthera canescens. For Chaenactis 

douglasii, stand with hydroseed mulch and seed treatment was not different from stand with row 

cover and seed treatment. Stand for the other species was too poor to draw conclusions.

In 2013 (Shock et al. 2014), the same trial was conducted with 15 species and included all of the 

species in the 2014 trial. In 2013, systems including row cover improved stand of 6 of the 

species including Chaenactis douglasii, Machaeranthera canescens, and Penstemon speciosus. 

In 2013, seed treatment only improved stand of Penstemon speciosus and reduced stand of 

Phacelia hastata. In 2013, sand improved stand of Chaenactis douglasii and Heliomeris 

multiflora.

Systems including row cover have been the most consistent factor for improving stand 

establishment over the years of trials at the Malheur Experiment Station (Shock et al. 2011, 

2012, and 2013). Seed treatment, sawdust, and sand are factors that for some species in some 

trials have shown value in improving stand, but their performance has not been consistent.

References

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2010. Emergence of native plantseeds 

in response to seed pelleting, planting depth, scarification, and soil anti-crusting treatment.

Oregon State University Malheur Experiment Station Annual Report 2009: 218-222.

Shock, C. C.; Feibert, E. B. G.; Parris, C.; Saunders, L. D.; Shaw, N. . 2011. Direct surface 

seeding strategies for establishment of Intermountain West native plants for seed production. 

Oregon State University Malheur Experiment Station Annual Report 2011, Ext/CrS 141: 130-

135.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Johnson, D.; Bushman, S. 2012. Direct surface 

seeding strategies for establishment of two native legumes of the Intermountain West. Oregon 

State University Malheur Experiment Station Annual Report 2012, Ext/CrS 144: 132-137.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2013. Direct surface seeding systems 

for successful establishment of native wildflowers. Oregon State University Malheur Experiment 

Station Annual Report 2013, Ext/CrS 149: 159-165.
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Table 1. Planting systems evaluated for emergence of eight native plant species in 2014. Mouse 

bait packs were scattered over the trial area. Malheur Experiment Station, Oregon State 

University, Ontario, OR.

No. Row cover Seed treatment* Sawdust Sand Mulch

1 yes yes yes no no

2 yes yes no no no

3 yes no yes no no

4 no yes yes no no

5 yes yes yes yes no

6 no yes no no yes

7 no no no no no

*Mixture of Captan and Thiram fungicides for prevention of seed decomposition and seedling dampingoff.

Table 2. Seed weights and tetrazolium test (seed viability) for seed submitted to emergence 

treatments in fall 2013 and evaluated in spring 2014. Malheur Experiment Station, Oregon State 

University, Ontario, OR.

Species Common name Untreated seed weight Tetrazolium test

seeds/g %

Chaenactis douglasii Douglas' dustymaiden 787 71.0

Machaeranthera canescens hoary tansyaster 2,321 84.0

Phacelia hastata silverleaf phacelia 1,471 98.0

Phacelia linearis threadleaf phacelia 2,843 98.0

Heliomeris multiflora showy goldeneye 1,700 83.0

Ligusticum porteri Porter's licorice-root 158 77.0

Ligusticum canbyi Canby's licorice-root 238 54.0

Penstemon speciosus showy penstemon 710 89.0
Table 3. Plant stands of eight native plant species on April 18, 2014 in response to seven planting systems used in 2013. Emergence 

for each species was corrected to the percent emergence of viable seed. Oregon State University, Malheur Experiment Station, 

Ontario, OR.

Species

Row cover, seed 

treatment, sawdust

Row cover, 

seed 

treatment

Row 

cover, 

sawdust

Seed 

treatment,

sawdust

Row cover, 

seed 

treatment, 

sawdust, sand

Seed 

treatment,

mulch

Untreated

check

LSD (0.

05)

------------------------------------------------------------% Stand------------------------------------------------------------------

Chaenactis douglasii 36.0 34.7 49.9 15.8 57.0 33.1 11.3 26.8

Machaeranthera canescens 67.3 63.0 76.4 28.4 70.2 8.6 26.4 31.6

Phacelia hastata 11.5 15.0 11.3 4.1 7.7 15.0 3.9 NS

Phacelia linearis 2.6 2.7 9.0 0.5 0.6 1.2 1.6 NS

Heliomeris multiflora 13.5 16.4 19.9 4.2 13.7 0.5 8.8 NS

Ligusticum porteri 1.1 0.4 0.9 0.0 0.9 0.2 0.0 0.7

Ligusticum canbyi 1.3 0.5 2.7 1.5 1.9 0.0 0.0 NS

Penstemon speciosus 5.4 1.7 1.9 1.9 12.2 1.9 0.7 6.1

100
101

B. Forb Response to Post-emergence Herbicides

Joel Felix, Joey Ishida, Erik B. G. Feibert, and Alicia Rivera, Malheur Experiment Station, 

Oregon State University, Ontario, OR

Introduction

Weed control methods are needed for the production of native wildflower seeds. Herbicides were 

tested for post emergence weed control. The methods and products reported here are not 

registered for use and do not construe recommendations. Growers should always carefully follow 

product labels.

Materials and Methods

Seed of 12 wildflower species were planted on the soil surface during fall 2013 (Table 1). After 

planting, a protective ground cover was immediately placed on each bed. The protective ground 

cover was removed on April 2, 2014.

Before planting, drip tape (T-Tape TSX 515-16-340) was buried at 12-inch depth midway 

between each pair of 30-inch rows. The flow rate for the drip tape was 0.34 gal/min/100ft at 8 psi 

with emitters spaced 16 inches apart, resulting in a water application rate of 0.066 inch/hour.

Treatments to evaluate forb species tolerance to various herbicides were applied post emergence 

on April 16 (early) or May 14, 2014 (late). The treatments were broadcast using a CO2 sprayer 

with 8002 nozzles at 30 psi applying 20 gal/acre. The study had a randomized complete block 

design with four replications. Each plot consisted of 12 single rows (one row/wildflowerspecies) 

and 7.33 ft wide. Herbicides were sprayed perpendicular to the rows, with all 12 species/plot 

sprayed in a single pass. Subjective plant damage ratings were taken in each plot on June 16, 

2014.

Results and Discussion

Eight species had good stand (Table 2). Lomatium dissectum (fernleaf biscuitroot), Phacelia 

linearis (threadleaf phacelia), Ligusticum porteri (Porter's licorice-root), and Ligusticum canbyi 

(Canby's licorice-root) did not emerge.

Evaluations on June 14 indicated very high injury for the plants that were sprayed on April 16, 

2014 (data not shown). All annual plant species were killed and perennial species were severely 

stunted. Evaluations indicated some level of herbicide tolerance for plants sprayed on May 14, 

2014 (Table 2). Sphaeralcea grossulariifolia, Eriogonum umbellatum, Penstemon speciosus, and 

Achillea millefolium suffered injury ranging from 24 to 95% across the herbicide treatments. The 

injury was characterized by stunting and reduced flowering. Injury for Machaeranthera 

canescens, Heliomeris multiflora, Chaenactis douglasii, and Phacelia hastata ranged from 4 to 

58%.

These results suggest that these wildflower species may be very sensitive to the herbicides 

evaluated, especially when applied early after removing the ground cover. However, it is

possible that directed sprays between rows using protective shields (though not tested in this 

study) could be an option for weed control in forbs. These treatments will be applied in the same 

plots in 2015 to evaluate the response of perennial species that will have well-established root 

systems.
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Table 1. Wildflower species planted in the fall of 2013 for the herbicide trial at the Malheur 

Experiment Station, Oregon State University, Ontario, OR.

Species Common name

1 Lomatium dissectum fernleaf biscuitroot

2 Sphaeralcea grossulariifolia gooseberryleaf globemallow

3 Eriogonum umbellatum sulphur-flower buckwheat

4 Penstemon speciosus royal penstemon, sagebrush penstemon

5 Achillea millefolium yarrow

6 Machaeranthera canescens hoary tansyaster

7 Heliomeris multiflora showy goldeneye

8 Chaenactis douglasii Douglas' dustymaiden

9 Phacelia hastata silverleaf phacelia

10 Phacelia linearis threadleaf phacelia

11 Ligusticum porteri Porter's licorice-root

12 Ligusticum canbyi Canby's licorice-root
Table 2. Response of eight forbs to post emergence herbicides for weed control in 2014. Malheur Experiment Station, Ontario, OR.

Treatment Rate Unit Product rate Sphaeralcea Eriogonum Penstemon

Injurya

Achillea Machaeranthera Heliomeris Chaenactis Phacelia

grossulariifolia umbellatum speciosus millefolium canescens multiflora douglasii hastata

---------------------------------------------------------- % -----------------------------------------------------

1 Untreated 0c 0b 0b 0c 0d 0a 0b 0b

2 Buctril 0.125 lb ai/a 0.5 pt/a 73a 30b 11b 24bc 21ab 11a 5b 58a

Prowl H2O 0.95 lb ai/a 2 pt/a

3 GoalTender 0.25 lb ai/a 0.5 pt/a 23bc 31b 39ab 24bc 14bc 6a 4b 48ab

Prowl H2O 0.95 lb ai/a 2 pt/a

4 Buctril 0.125 lb ai/a 0.5 pt/a 83a 31b 43ab 40ab 31a 11a 23ab 35ab

GoalTender 0.25 lb ai/a 0.5 pt/a

Prowl H2O 0.95 lb ai/a 2 pt/a

5 Valor 

(Chateau) 0.128 lb ai/a 4 oz/a 74a 21b 79a 31ab 8cd 11a 13b 35ab

Prowl H2O 0.95 lb ai/a 2 pt/a

6 Basagran 1.125 lb ai/a 2.25 pt/a 65ab 30b 53ab 58a 23ab 6a 8b 14ab

Prowl H2O 0.95 lb ai/a 2 pt/a

7 Linex 1 lb ai/a 2 pt/a 55ab 91a 73a 38ab 14bc 11a 40a 16ab

Prowl H2O 0.95 lb ai/a 2 pt/a

8 Raptor 0.0156lb ae/a 0.125pt/a 95a 33b 46ab 35ab 18bc 1a 5b 54a

Basagran 1.125 lb ai/a 2.25 pt/a

Prowl H2O 0.95 lb ai/a 2 pt/a

LSD (P=0.05) 49 33 53 28 13 14 23 48 a Means within a column followed by same letter do not significantly differ (P=0.05, LSD)

103
104

C. Irrigation Requirements for Native Perennial Wildflower Seed Production in a Semiarid Environment

Clinton C. Shock, Erik B. G. Feibert, Alicia Rivera, and Lamont D. Saunders, Malheur 

Experiment Station, Oregon State University, Ontario, OR, 2014

Nancy Shaw and Francis Kilkenny, U.S. Forest Service, Rocky Mountain Research Station, 

Boise, ID

Ram S. Sampangi, University of Idaho, Parma, ID

Introduction

Native wildflower seed is needed to restore rangelands of the Intermountain West. Commercial 

seed production is necessary to provide the quantity of seed needed for restoration efforts. A 

major limitation to economically viable commercial production of native wildflower (forb)seed 

is stable and consistent seed productivity over years.

In native rangelands, the natural variations in spring rainfall and soil moisture result in highly 

unpredictable water stress at flowering, seed set, and seed development, which for other seed 

crops is known to compromise seed yield and quality.

Native wildflower plants are not well adapted to croplands. They often are not competitive with 

crop weeds in cultivated fields. Poor competitiveness with weeds could also limit wildflower 

seed production. Both sprinkler and furrow irrigation could provide supplemental water for seed 

production, but these irrigation systems risk further encouraging weeds. Also, sprinkler and 

furrow irrigation can lead to the loss of plant stand and seed production due to fungal pathogens. 

By burying drip tapes at 12-inch depth and avoiding wetting the soil surface, we hoped to assure 

flowering and seed set without undue encouragement of weeds or opportunistic diseases. The 

trials reported here tested the effects of three low rates of irrigation on the seed yield of 13 native 

wildflower species.

Materials and Methods

Plant Establishment

Seed of seven Intermountain West wildflower species (the first seven species in Table 1) was 

received in late November in 2004 from the Rocky Mountain Research Station (Boise, ID). The 

plan was to plant the seed in the fall of 2004, but due to excessive rainfall in October, the ground 

preparation was not completed and planting was postponed to early 2005. To try to ensure 

germination, the seed was submitted to cold stratification. The seed was soaked overnight in 

distilled water on January 26, 2005, after which the water was drained and the seed soaked for 20 

min in a 10 percent by volume solution of 13 percent bleach in distilled water. The water was 

drained and the seed was placed in thin layers in plastic containers. The plastic containers had 

lids with holes drilled in them to allow air movement. These containers were placed in a cooler 

set at approximately 34°F. Every few days the seed was mixed and, if necessary, distilled water

added to maintain seed moisture. In late February, seed of Lomatium grayi and L. triternatum 

(see Table 1 for common names) had started to sprout.

In late February 2005, drip tape (T-Tape TSX 515-16-340) was buried at 12-inch depth between 

two 30-inch rows of a Nyssa silt loam with a pH of 8.3 and 1.1 percent organic matter. The drip

tape was buried in alternating inter-row spaces (5 ft apart). The flow rate for the drip tape was
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0. 34 gal/min/100ft at 8 psi with emitters spaced 16 inches apart, resulting in a water application 

rate of 0.066 inch/hour.

On March 3, seed of the first seven species in Table 1 was planted in 30-inch rows using a 

custom-made plot grain drill with disc openers. All seed was planted at 20-30 seeds/ft. of row. 

The Eriogonum umbellatum and the Penstemon spp. were planted at 0.25-inch depth and the 

Lomatium spp. at 0.5-inch depth. The trial was irrigated with a minisprinkler system (R10 Turbo 

Rotator, Nelson Irrigation Corp., Walla Walla, WA) for even stand establishment from March 4 

to April 29. Risers were spaced 25 ft apart along the flexible polyethylene hose laterals that were 

spaced 30 ft apart and the water application rate was 0.10 inch/hour. A total of 1.72 inches of 

water was applied with the minisprinkler system. Eriogonum umbellatum, Lomatium triternatum, 

and L. grayi started emerging on March 29. All other species except L. dissectum emerged by 

late April. Starting June 24, the field was irrigated with the drip system. A total of 3.73 inches of 

water was applied with the drip system from June 24 to July 7. The field was not irrigated further 

in 2005.

Plant stands for Eriogonum umbellatum, Penstemon spp., Lomatium triternatum, and L. grayi 

were uneven. Lomatium dissectum did not emerge. None of the species flowered in 2005. In 

early October, 2005, more seed was received from the Rocky Mountain Research Station for 

replanting. The empty lengths of row were replanted by hand in the E. umbellatum and 

Penstemon spp. plots. The Lomatium spp. plots had the entire row lengths replanted using the 

planter. The seed was replanted on October 26, 2005. In the spring of 2006, the plant stands of 

the replanted species were excellent, except for P. deustus.

On April 11, 2006, seed of three globemallow species (Sphaeralcea parvifolia, S.

grossulariifolia, S. coccinea), two prairie clover species (Dalea searlsiae, D. ornata), and basalt 

milkvetch (Astragalus filipes) was planted at 30 seeds/ft of row. The field was sprinkler irrigated 

until emergence. Emergence was poor. In late August of 2006, seed of the three globemallow 

species was harvested by hand. On November 9, 2006, the six wildflowers that were planted in 

2006 were mechanically flailed and on November 10, they were replanted. On November 11, the 

Penstemon deustus plots were also replanted at 30 seeds/ft of row.

Table 1. Wildflower species planted in the drip irrigation trials at the MalheurExperiment 

Station, Oregon State University, Ontario, OR.

Species Common names

Eriogonum umbellatum Sulphur-flower buckwheat

Penstemon acuminatus Sharpleaf penstemon, sand-dune penstemon

Penstemon deustus Scabland penstemon, hotrock penstemon

Penstemon speciosus Royal penstemon, sagebrush penstemon

Lomatium dissectum Fernleaf biscuitroot

Lomatium triternatum Nineleaf biscuitroot, nineleaf desert parsley

Lomatium grayi Gray’s biscuitroot, Gray’s lomatium

Sphaeralcea parvifolia Smallflower globemallow

Sphaeralcea grossulariifolia Gooseberryleaf globemallow

Sphaeralcea coccinea Scarlet globemallow, red globemallow
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Irrigation for Seed Production

In April, 2006 each planted strip of each wildflower species was divided into plots 30 ft long. 

Each plot contained four rows of each species. The experimental designs were randomized 

complete blocks with four replicates. The three treatments were a non-irrigated check, 1 inch of 

water applied per irrigation, and 2 inches of water applied per irrigation. Each treatment received 

4 irrigations that were applied approximately every 2 weeks starting with flowering of the 

wildflowers. The amount of water applied to each treatment was calculated by the length of time 

necessary to deliver 1 or 2 inches through the drip system. Irrigations were regulated with a 

controller and solenoid valves. After each irrigation the amount of water applied was read on a 

water meter and recorded to ensure correct water applications.

In March of 2007, the drip-irrigation system was modified to allow separate irrigation of the 

species due to different timings of flowering. The three Lomatium spp. were irrigated together 

and Penstemon deustus and P. speciosus were irrigated together, but separately from the others. 

Penstemon acuminatus and Eriogonum umbellatum were irrigated individually. In early April, 

2007 the three globemallow species, two prairie clover species, and basalt milkvetch were 

divided into plots with a drip-irrigation system to allow the same irrigation treatments that were 

received by the other wildflowers.

Irrigation dates are found in Table 2. In 2007, irrigation treatments were inadvertently continued 

after the fourth irrigation. Irrigation treatments for all species were continued until the last 

irrigation on June 24, 2007.

Soil volumetric water content was measured by neutron probe. The neutron probe was calibrated 

by taking soil samples and probe readings at 8-, 20-, and 32-inch depths during installation of the 

access tubes. The soil water content was determined volumetrically from the soil samples and 

regressed against the neutron probe readings separately for each soil depth. Regression equations 

were then used to transform the neutron probe readings into volumetric soil water content.

Flowering, Harvesting, and Seed Cleaning

Flowering dates for each species were recorded (Table 2). The Eriogonum umbellatum and 

Penstemon spp. plots produced seed in 2006, in part because they had emerged in the spring of 

2005. Each year, the middle two rows of each plot were harvested when seed of each species was 

mature (Table 2), using the methods listed in Table 3. The plant stand for P. deustus was too 

poor to result in reliable seed yield estimates. Replanting P. deustus in fall 2006 did not result in 

adequate plant stand in the spring of 2007.

Eriogonum umbellatum seeds did not separate from the flowering structures in the combine; the 

unthreshed seed was taken to the U. S. Forest Service Lucky Peak Nursery (Boise, ID) and run 

through a dewinger to separate seed. The seed was further cleaned in a small clipper seed 

cleaner.

Penstemon deustus seed pods were too hard to be opened in the combine; the unthreshed seed 

was precleaned in a small clipper seed cleaner and then seed pods were broken manually by 

rubbing the pods on a ribbed rubber mat. The seed was then cleaned again in the small clipper 

seed cleaner.
107

Penstemon acuminatus and P. speciosus were threshed in the combine and the seed was further 

cleaned using a small clipper seed cleaner.

Cultural Practices in 2006

On October 27, 2006, 50 lb. phosphorus (P)/acre and 2 lb. zinc (Zn)/acre were injected through 

the drip tape to all plots of Eriogonum umbellatum, Penstemon spp., and Lomatium spp. On 

November 11, 100 lb. nitrogen (N)/acre as urea was broadcast to all Lomatium spp. plots. On 

November 17, all plots of Eriogonum umbellatum, Penstemon spp. (except P. deustus), and 

Lomatium spp. had Prowl® at 1 lb. ai/acre broadcast on the soil surface. Irrigations for all species 

were initiated on May 19 and terminated on June 30. Harvesting and seed cleaning methodsfor 

each species are listed in Table 3.

Cultural Practices in 2007

Penstemon acuminatus and P. speciosus were sprayed with Aza-Direct® at 0.0062 lb. ai/acre on 

May 14 and 29 for lygus bug control. Irrigations for each species were initiated and terminated 

on different dates (Table 2). Harvesting and seed cleaning methods for each species are listed in 

Table 3. All plots of the Sphaeralcea spp. were flailed on November 8, 2007.

Cultural Practices in 2008

On November 9, 2007 and on April 15, 2008, Prowl® at 1 lb. ai/acre was broadcast on all plots

for weed control. Capture® 2EC at 0.1 lb. ai/acre was sprayed on all plots of Penstemon

acuminatus and P. speciosus on May 20 for lygus bug control. Irrigations for each species were 

initiated and terminated on different dates (Table 2). Harvesting and seed cleaning methods for 

each species are listed in Table 3.

Cultural Practices in 2009

On March18, Prowl® at 1 lb. ai/acre and Volunteer® at 8 oz/acre were broadcast on all plots for 

weed control. On April 9, 50 lb. N/acre and 10 lb. P/acre were applied through the drip irrigation 

system to the three Lomatium spp.

The flowering, irrigation timing, and harvest timing were recorded for each species (Table 2). 

Harvesting and seed cleaning methods for each species are listed in Table 3. On December 4, 

2009, Prowl at 1 lb. ai/acre was broadcast for weed control on all plots.

Cultural Practices in 2010

The flowering, irrigation, and harvest timing of the established wildflowers were recorded for 

each species (Table 2). Harvesting and seed cleaning methods for each species are listed in Table

3. On November 17, Prowl® at 1 lb. ai/acre was broadcast on all plots for weed control.

Cultural Practices in 2011

On May 3, 2011, 50 lb. N/acre was applied to all Lomatium spp. plots as Uran (urea ammonium 

nitrate) injected through the drip tape. The timing of flowering, irrigations, and harvests varied 

by species (Table 2). Harvesting and seed cleaning methods for each species are listed in Table

3. On November 9, Prowl® at 1 lb. ai/acre was broadcast on all plots for weed control.
108

Cultural Practices in 2012

The soil volumetric water content was very low in 2012 prior to the onset of irrigation for each 

species. Iron deficiency symptoms were prevalent in 2012. On April 13, 50 lb. N/acre, 10 lb.

P/acre, and 0.3 lb. iron (Fe)/acre was applied to all Lomatium spp. plots as liquid fertilizer 

injected through the drip tape. On November 7, Prowl® at 1 lb. ai/acre was broadcast on all plots 

of Eriogonum umbellatum, Lomatium grayi, Lomatium dissectum, and Lomatium triternatum for 

weed control.

Cultural Practices in 2013

On March 29, 20 lb. N/acre, 25 lb. P/acre, and 0.3 lb. Fe/acre were applied through the drip tape 

to all plots of Lomatium grayi, Lomatium dissectum, and Lomatium triternatum. On April 3, 

Select Max at 32 oz/acre was broadcast for grass weed control on all plots of Eriogonum 

umbellatum, Lomatium grayi, Lomatium dissectum, Lomatium triternatum, and Penstemon 

speciosus.

Cultural Practices in 2014

On February 26, Prowl® at 1 lb. ai/acre and Select Max at 32 oz/acre were broadcast on all plots 

of Eriogonum umbellatum, Lomatium grayi, Lomatium dissectum, and Lomatium triternatum for 

weed control. On April 2, 20 lb. N/acre, 25 lb. P/acre, and 0. 3 lb. Fe/acre were applied through 

the drip tape to all plots of Lomatium grayi, Lomatium dissectum, and Lomatium triternatum. On 

April 18, Orthene at 8 oz/acre was broadcast to all plots of Penstemon speciosus for lygus bug 

control. On April 29, 5 lb. Fe/acre was applied through the drip tape to all plots of Penstemon 

speciosus.
109

Table 2, continued on next pages. Native wildflower flowering, irrigation, and seed harvest 

dates by species in 2006-2014, Malheur Experiment Station, Oregon State University, Ontario, 

OR.

Flowering Irrigation

Species Start Peak End Start End Harvest

2006

Eriogonum umbellatum 19-May 20-Jul 19-May 30-Jun 3-Aug

Penstemon acuminatus 2-May 10-May 19-May 19-May 30-Jun 7-Jul

Penstemon deustus 10-May 19-May 30-May 19-May 30-Jun 4-Aug

Penstemon speciosus 10-May 19-May 30-May 19-May 30-Jun 13-Jul

Lomatium dissectum 19-May 30-Jun

Lomatium triternatum 19-May 30-Jun

Lomatium grayi 19-May 30-Jun

Sphaeralcea parvifolia

S. grossulariifolia

Sphaeralcea coccinea

2007

Eriogonum umbellatum 25-May 25-Jul 2-May 24-Jun 31-Jul

Penstemon acuminatus 19-Apr 25-May 19-Apr 24-Jun 9-Jul

Penstemon deustus 5-May 25-May 25-Jun 19-Apr 24-Jun

Penstemon speciosus 5-May 25-May 25-Jun 19-Apr 24-Jun 23-Jul

Lomatium dissectum 5-Apr 24-Jun

Lomatium triternatum 25-Apr 1-Jun 5-Apr 24-Jun 29-Jun, 16-Jul

Lomatium grayi 5-Apr 10-May 5-Apr 24-Jun 30-May, 29-Jun

Sphaeralcea parvifolia 5-May 25-May 16-May 24-Jun 20-Jun, 10-Jul, 13-Aug

S. grossulariifolia 5-May 25-May 16-May 24-Jun 20-Jun, 10-Jul, 13-Aug

Sphaeralcea coccinea 5-May 25-May 16-May 24-Jun 20-Jun, 10-Jul, 13-Aug

2008

Eriogonum umbellatum 5-Jun 19-Jun 20-Jul 15-May 24-Jun 24-Jul

Penstemon acuminatus 29-Apr 5-Jun 29-Apr 11-Jun 11-Jul

Penstemon deustus 5-May 20-Jun 29-Apr 11-Jun

Penstemon speciosus 5-May 20-Jun 29-Apr 11-Jun 17-Jul

Lomatium dissectum 10-Apr 29-May

Lomatium triternatum 25-Apr 5-Jun 10-Apr 29-May 3-Jul

Lomatium grayi 25-Mar 15-May 10-Apr 29-May 30-May, 19-Jun

Sphaeralcea parvifolia 5-May 15-Jun 15-May 24-Jun 21-Jul

S. grossulariifolia 5-May 15-Jun 15-May 24-Jun 21-Jul

Sphaeralcea coccinea 5-May 15-Jun 15-May 24-Jun 21-Jul
110

Table 2, continued. Native wildflower flowering, irrigation, and seed harvest dates by speciesin 

2006-2014. Malheur Experiment Station, Oregon State University, Ontario, OR.

Flowering Irrigation

Species start peak end start end Harvest

2009

Eriogonum umbellatum 31-May 15-Jul 19-May 24-Jun 28-Jul

Penstemon acuminatus 2-May 10-Jun 8-May 12-Jun 10-Jul

Penstemon deustus 19-May 24-Jun

Penstemon speciosus 14-May 20-Jun 19-May 24-Jun 10-Jul

Lomatium dissectum 10-Apr 7-May 20-Apr 28-May 16-Jun

Lomatium triternatum 10-Apr 7-May 1-Jun 20-Apr 28-May 26-Jun

Lomatium grayi 10-Mar 7-May 20-Apr 28-May 16-Jun

Sphaeralcea parvifolia 1-May 10-Jun 22-May 24-Jun 14-Jul

Sphaeralcea grossulariifolia 1-May 10-Jun 22-May 24-Jun 14-Jul

Sphaeralcea coccinea 1-May 10-Jun 22-May 24-Jun 14-Jul

2010

Eriogonum umbellatum 4-Jun 12-19 Jun 15-Jul 28-May 8-Jul 27-Jul

Penstemon speciosus 14-May 20-Jun 12-May 22-Jun 22-Jul

Lomatium dissectum 25-Apr 20-May 15-Apr 28-May 21-Jun

Lomatium triternatum 25-Apr 15-Jun 15-Apr 28-May 22-Jul

Lomatium grayi 15-Mar 15-May 15-Apr 28-May 22-Jun

Sphaeralcea parvifolia 10-May 4-Jun 25-Jun 28-May 8-Jul 20-Jul

Sphaeralcea grossulariifolia 10-May 4-Jun 25-Jun 28-May 8-Jul 20-Jul

Sphaeralcea coccinea 10-May 4-Jun 25-Jun 28-May 8-Jul 20-Jul

2011

Eriogonum umbellatum 8-Jun 30-Jun 20-Jul 20-May 5-Jul 1-Aug

Penstemon speciosus 25-May 30-May 30-Jun 20-May 5-Jul 29-Jul

Lomatium dissectum 8-Apr 25-Apr 10-May 21-Apr 7-Jun 20-Jun

Lomatium triternatum 30-Apr 23-May 15-Jun 21-Apr 7-Jun 26-Jul

Lomatium grayi 1-Apr 25-Apr 13-May 21-Apr 7-Jun 22-Jun

Sphaeralcea parvifolia 26-May 15-Jun 14-Jul 20-May 5-Jul 29-Jul

Sphaeralcea grossulariifolia 26-May 15-Jun 14-Jul 20-May 5-Jul 29-Jul

Sphaeralcea coccinea 26-May 15-Jun 14-Jul 20-May 5-Jul 29-Jul
111

Table 2, continued. Native wildflower flowering, irrigation, and seed harvest dates by speciesin 

2006-2014. Malheur Experiment Station, Oregon State University, Ontario, OR.

Flowering Irrigation

Species start peak end start end Harvest

2012

Eriogonum umbellatum 30-May 20-Jun 4-Jul 30-May 11-Jul 24-Jul

Penstemon speciosus 2-May 20-May 25-Jun 2-May 13-Jun 13-Jul

Lomatium dissectum 9-Apr 16-Apr 16-May 13-Apr 24-May 4-Jun

Lomatium triternatum 12-Apr 17-May 6-Jun 13-Apr 24-May 21-Jun

Lomatium grayi 15-Mar 25-Apr 16-May 13-Apr 24-May 14-Jun

2013

Eriogonum umbellatum 8-May 27-May 27-Jun 8-May 19-Jun 9-Jul

Penstemon speciosus 2-May 10-May 20-Jun 2-May 12-Jun 11-Jul

Lomatium dissectum 10-Apr 25-Apr 4-Apr 16-May 4-Jun

Lomatium triternatum 18-Apr 10-May 4-Apr 16-May 4-Jun

Lomatium grayi 15-Mar 30-Apr 4-Apr 16-May 10-Jun

2014

Eriogonum umbellatum 20-May 4-Jun 1-Jul 13-May 24-Jun 10-Jul

Penstemon speciosus 29-Apr 13-May 9-Jun 29-Apr 10-Jun 11-Jul

Lomatium dissectum 28-Mar 21-Apr 7-Apr 20-May 2-Jun

Lomatium triternatum 7-Apr 29-Apr May 2 7-Apr 20-May 4-Jun

Lomatium grayi 28-Mar May 2 7-Apr 20-May 10-Jun
112

Table 3. Native wildflower seed harvest and cleaning by species, Malheur Experiment Station, 

Oregon State University, Ontario, OR.

Species Number of 

harvests/year

Harvest method PrecleaningThreshing 

method

Cleaning

method

Eriogonum umbellatum 1 combinea none dewingerb mechanicalc

Penstemon acuminatus 1 combined none combine mechanicalc

Penstemon deustus 1 combinea mechanicalc hande mechanicalc

Penstemon speciosusf 1 combined none combine mechanicalc

Lomatium dissectum 1 hand hand none mechanicalc

Lomatium triternatum 1–2 hand hand none mechanicalc

Lomatium grayi 1–2 hand hand none mechanicalc

Sphaeralcea parvifolia 1–3 hand or combined none combine none

Sphaeralcea grossulariifolia 1–3 hand or combined none combine none

Sphaeralcea coccinea 1–3 hand or combined none combine none

Dalea searlsiae 0 or 2 hand none dewinger mechanicalc

Dalea ornate 0 or 2 hand none dewinger mechanicalc

a Wintersteiger Nurserymaster small-plot combine with dry bean concave.

b Specialized seed threshing machine at USDA Lucky Peak Nursery used in 2006. Thereafter an adjustablehanddriven corn grinder was used to thresh seed. c Clipper seed cleaner.

d Wintersteiger Nurserymaster small-plot combine with alfalfa seed concave. For the Sphaeralcea spp., flailingin

the fall of 2007 resulted in more compact growth and one combine harvest in 2008, 2009, and 2010. e Hard seed pods were broken by rubbing against a ribbed rubber mat. f Harvested by hand in 2007 and 2009 due to poor seed set.

Results and Discussion

Precipitation from January through June in 2014 was 5.1 inches, close to the average of 5.8 

inches (Table 4). The accumulated growing degree-days (50-86°F) from January through June in 

2014 were higher than average (Table 4, Figure 1 and 2).

Flowering and Seed Set

Penstemon acuminatus and P. speciosus had poor seed set in 2007, partly due to a heavy lygus 

bug infestation that was not adequately controlled by the applied insecticides. In the Treasure 

Valley, the first hatch of lygus bugs occurs when 250 degree-days (52°F base) are accumulated. 

Data collected by an AgriMet weather station adjacent to the field indicated that the first lygus 

bug hatch occurred on May 14, 2006; May 1, 2007; May 18, 2008; May 19, 2009; and May 29,

2010. The average (1995-2010) lygus bug hatch date was May 18. Penstemon acuminatus and P. 

speciosus start flowering in early May. The earlier lygus bug hatch in 2007 probably resulted in 

harmful levels of lygus bugs present during a larger part of the Penstemon spp. flowering period 

than normal. Poor seed set for P. acuminatus and P. speciosus in 2007 also was related to poor 

vegetative growth compared to 2006 and 2008. In 2009 all plots of P. acuminatus and P. 

speciosus again showed poor vegetative growth and seed set. Root rot affected all plots of P. 

acuminatus in 2009, killing all plants in two of the four plots of the wettest treatment (2 inches
113

per irrigation). Root rot affected the wetter plots of P. speciosus in 2009, but the stand partially 

recovered due to natural reseeding.

The three Sphaeralcea spp. showed a long flowering period (early May through September) in 

2007. Multiple manual harvests were necessary because the seed falls out of the capsules once 

they are mature. The flailing of the three Sphaeralcea spp. starting in the fall of 2007 was done 

annually to induce a more concentrated flowering, allowing only one mechanical harvest.

Precipitation in June of 2009 (2.27 inches) and 2010 (1.95 inches) was substantially higher than 

average (0.76 inches). Rust (Puccinia sherardiana) infected all three Sphaeralcea spp. in June of 

2009 and 2010, causing substantial leaf loss and reduced vegetative growth. Stand of all three 

Sphaeralcea spp. deteriorated in 2011 and the plots were disked out in 2012.

Eriogonum umbellatum

In 2006, seed yield of Eriogonum umbellatum increased with increasing water application, up to 

8 inches, the highest amount tested (Tables 5 and 7). In 2007-2009 seed yield showed a quadratic 

response to irrigation rate (Tables 6 and 7). Seed yields were maximized by 8.1 inches, 7.2 

inches, and 6.9 inches of water applied in 2007, 2008, and 2009, respectively. In 2010, there was 

no significant difference in yield between treatments. In 2011, seed yield was highest with no 

irrigation. The 2010 and 2011 seasons had unusually cool (Table 4, Figure 2) and wet weather 

(Figure 2). The accumulated precipitation in April through June of 2010 and 2011 was the 

highest over the years of the trial (Table 4). The relatively high seed yield of E. umbellatum in 

the non-irrigated treatment in 2010 and 2011 seemed to be related to the high spring 

precipitation. The negative effect of irrigation on seed yield in 2011 might have been related to 

the presence of rust. Irrigation could have exacerbated the rust and resulted in lower yields. In 

2012, seed yield of Eriogonum umbellatum increased with increasing water application, up to 8 

inches, the highest amount tested (Tables 6 and 7). In 2013 and 2014, seed yields showed 

quadratic responses to irrigation rate. Seed yield was maximized by 5.7 inches and by 5.3 inches 

of water applied in 2013 and 2014, respectively. Averaged over 9 years, seed yield of E. 

umbellatum showed a quadratic response to irrigation rate with the highest yield achieved with

6.9 inches of water applied.

Penstemon acuminatus

There was no significant difference in seed yield between irrigation treatments for P. acuminatus 

in 2006 (Table 5). Precipitation from March through June was 6.4 inches in 2006. The 64-yearaverage precipitation from March through June is 3.6 inches. The wet weather in 2006 could 

have attenuated the effects of the irrigation treatments. In 2007 seed yield showed a quadratic 

response to irrigation rate. Seed yields were maximized by 4.0 inches of water applied in 2007. 

In 2008, seed yield showed a linear response to applied water. In 2009 there was no significant 

difference in seed yield between treatments (Table 7). However, due to root rot affecting all plots 

in 2009, the seed yield results were compromised. By 2010, substantial lengths of row contained 

only dead plants. Measurements in each plot showed that plant death increased with increasing 

irrigation rate. The stand loss was 51.3, 63.9, and 88.5 percent for the 0-, 4-, and 8-inch irrigation 

treatments, respectively. The trial area was disked out in 2010. Following the 2005 planting, seed 

yields were substantial in 2006 and moderate in 2008. P. acuminatus performed as a short-lived 

perennial.
114

Penstemon speciosus

In 2006-2009 seed yield of P. speciosus showed a quadratic response to irrigation rate (Tables 5, 

6, and 7). Seed yields were maximized by 4.3, 4.2, 5.0, and 4.3 inches of water applied in 2006,

2007, 2008, and 2009, respectively. In 2010, 2011, and 2012 there was no difference in seed 

yield between treatments. Seed yield was low in 2007 due to lygus bug damage, as discussed 

previously. Seed yield in 2009 was low due to stand loss from root rot. The plant stand recovered 

somewhat in 2010 and 2011, due in part to natural reseeding, especially in the non-irrigated 

plots. In 2013, seed yield increased with increasing water application, up to 8 inches, the highest 

amount tested. In 2014 seed yield of P. speciosus showed a quadratic response to irrigation rate 

with the calculated optimum of 276 lb. /acre at 5.2 inches of irrigation water. Averaged over 9 

years, seed yield was maximized by 5.0 inches of water applied.

Lomatium triternatum

Lomatium triternatum showed a trend for increasing seed yield with increasing irrigation rate in 

2007 (Table 5). The highest irrigation rate resulted in significantly higher seed yield than the 

non-irrigated check in seven of the last 8 years. Seed yields of L. triternatum were substantially 

higher in 2008-2011 (Tables 6 and 7). In 2008–2011 seed yields of L. triternatum showed a 

quadratic response to irrigation rate. Seed yields were estimated to be maximized by 8.4, 5.4, 

7.8, and 4.1 inches of water applied in 2008, 2009, 2010, and 2011, respectively. In 2012, 

2013, and 2014 seed yield increased linearly with increasing water applied up to the highest 

amount of water applied, 8 inches. Over 8 years, seed yield of L. triternatum was estimated to 

be maximized at 1328 lb. /acre/year at 6.9 inches of applied water (Table 7).

Lomatium grayi

Lomatium grayi showed a trend for increasing seed yield with increasing irrigation rate in 2007 

(Table 5). The highest irrigation rate resulted in significantly higher seed yield than the nonirrigated check in six of the last eight years. Seed yields of L. grayi were substantially higher in 

2008 and 2009. In 2008, seed yields of L. grayi showed a quadratic response to irrigation rate. 

Seed yields were estimated to be maximized by 6.9 inches of water applied in 2008. In 2009, 

seed yield showed a linear response to irrigation rate (Tables 6 and 7). Seed yield with the 4-inch 

irrigation rate was significantly higher than in the non-irrigated check, but the 8-inch irrigation 

rate did not result in a significant increase above the 4-inch rate.

In 2010, seed yield was not responsive to irrigation, possibly caused by the unusually wetspring 

of 2010. A further complicating factor in 2010 that compromised seed yields was rodent damage. 

Extensive rodent (vole) damage occurred over the 2009-2010 winter. The affected areas were 

transplanted with 3-year-old L. grayi plants from an adjacent area in the spring of 2010. To 

reduce their attractiveness to voles, the plants were mowed after becoming dormant in early fall 

of 2010 and in each subsequent year. In 2011 seed yield again did not respond to irrigation. The 

spring of 2011 was unusually cool and wet. In 2012 seed yields of L. grayi showed a quadratic 

response to irrigation rate, with a maximum seed yield at 5.5 inches of applied water. In 2013 

seed yield increased linearly with increasing water application, up to 8 inches, the highest 

amount applied. In 2014 seed yields of L. grayi showed a quadratic response to irrigation rate, 

with a calculated maximum seed yield of 1477 lb. /acre at 5.3 inches of applied water. Over 8 

years, seed yield of L. grayi was estimated to be maximized at 827 lb. /acre/year by 5.4 inches of
115

applied water. Most appropriately, irrigation probably should be variable according to 

precipitation.

Lomatium dissectum

Lomatium dissectum had very little vegetative growth during 2006-2008, and produced only very 

few flowers in 2008. All the Lomatium species tested were affected by Alternaria fungus, but the 

infection was greatest on the L. dissectum selection planted in this trial. This infection delayed L. 

dissectum plant development. In 2009, vegetative growth and flowering for L. dissectum were 

improved.

Seed yield of L. dissectum showed a linear response to irrigation rate in 2009 (Figure 14). Seed 

yield with the 4-inch irrigation rate was significantly higher than with the non-irrigated check, 

but the 8-inch irrigation rate did not result in a significant increase above the 4-inch rate. In 2010 

and 2011 seed yields of L. dissectum showed a quadratic response to irrigation rate. Seed yields 

were estimated to be maximized by 5.4 and 5.1 inches of applied water in 2010 and 2011 

respectively. In 2012, seed yields of L. dissectum were not responsive to irrigation rate. In 2013 

and 2014, seed yield increased linearly with increasing irrigation rate up to 8 inches, the highest 

amount applied. Over the 5 years, seed yield showed a quadratic response to irrigation rate and 

was estimated to be maximized at 869 lb./acre/year by applying 6.1 inches of applied water.

Sphaeralcea spp.

In 2007-2011, there were no significant differences in seed yield among irrigation treatments

for the three Sphaeralcea spp. (Tables 5 and 6). Stand of the three Sphaeralcea species was 

poor in 2012 and the plantings were eliminated.

Conclusions

Subsurface drip irrigation systems were tested for native seed production because they have two 

potential strategic advantages: a) low water use, and b) the buried drip tape provides water to the 

plants at depth, precluding most irrigation induced stimulation of weed seed germination on the 

soil surface and keeping water away from native plant tissues that are not adapted to a wet 

environment.

Due to the arid environment, supplemental irrigation may often be required for successful 

flowering and seed set because soil water reserves may be exhausted before seed formation. The 

total irrigation requirements for these arid-land species were low and varied by species (Table 7). 

The Sphaeralcea spp. and Penstemon acuminatus did not respond to irrigation in these trials.

Natural rainfall was sufficient to maximize seed production in the absence of weed competition.

Lomatium dissectum required approximately 6 inches of irrigation. Lomatium grayi, L. 

triternatum, and Eriogonum umbellatum responded quadratically to irrigation with the optimum 

varying by year. Penstemon deustus had insufficient plant stands to reliably evaluate itsresponse 

to irrigation over multiple years.
116

Table 4. Early season precipitation and growing degree-days at the 

Malheur Experiment Station, Ontario, OR, 2006-2014.

Precipitation (inches) Growing degree-days (50-86°F)

Year Jan-June April-June Jan-June

2006 9.0 3.1 1120

2007 3.1 1.9 1208

2008 2.9 1.2 936

2009 5.8 3.9 1028

2010 8.3 4.3 779

2011 8.3 3.9 671

2012 5.8 2.3 979

2013 2.6 1.4 1118

2014 5.1 1.6 1109

70-year average 5.8 2.7 1010a

a

24-year average.
117

Figure 1. Growing degree days (50 – 86 oF) for 2006 through 2009 and 24-year 

average. Malheur Experiment Station, Oregon State University, 2014.

Figure 2. Growing degree days (50 – 86 o

F) for 2010 through 2014 and 24-year 

average. Malheur Experiment Station, Oregon State University, 2014.
118

Table 5. Native wildflower seed yield response to irrigation rate (inches/season) in 2006 through 

2011. Malheur Experiment Station, Oregon State University, Ontario, OR.

2006 2007 2008

Species 0 inch 4 inch 8 inch LSD

(0.05)

0 inch 4 inch 8 inch LSD

(0.05)

0 inch 4 inch 8 inch LSD

(0.05)

---------------------------------------------------------- lb/acre --------------------------------------------------------

Eriogonum umbellatuma 155.3 214.4 371.6 92.9 79.6 164.8 193.8 79.8 121.3 221.5 245.2 51.7

Penstemon acuminatusa 538.4 611.1 544.0 NS 19.3 50.1 19.1 25.5b 56.2 150.7 187.1 79.0

Penstemon deustusc 1246 1201 1069 NS 120.3 187.7 148.3 NS --- very poor stand ---

Penstemon speciosusa 163.5 346.2 213.6 134.3 2.5 9.3 5.3 4.7b 94.0 367.0 276.5 179.6

Lomatium dissectumd ---- no flowering ---- --- no flowering --- - very little flowering -

Lomatium triternatumd ---- no flowering ---- 2.3 17.5 26.7 16.9b 195.3 1061 1387 410.0

Lomatium grayid ---- no flowering ---- 36.1 88.3 131.9 77.7b 393.3 1287 1445 141.0

Sphaeralcea parvifoliae 1063 850.7 957.9 NS 436.2 569.1 544.7 NS

Sphaeralcea grossulariifoliae 442.6 324.8 351.9 NS 275.3 183.3 178.7 NS

Sphaeralcea coccineae 279.8 262.1 310.3 NS 298.7 304.1 205.2 NS

2009 2010 2011

Species 0 inch 4 inch 8 inch LSD

(0.05)

0 inch 4 inch 8 inch LSD

(0.05)

0 inch 4 inch 8 inch LSD

(0.05)

-------------------------------------------------------- lb/acre -------------------------------------------------------

Eriogonum umbellatuma 132.3 223 240.1 67.4 252.9 260.3 208.8 NS 248.7 136.9 121.0 90.9

Penstemon acuminatusa 20.7 12.5 11.6 NS -- Stand disked out --

Penstemon speciosusa 6.8 16.1 9.0 6.0b 147.2 74.3 69.7 NS 371.1 328.2 348.6 NS

Lomatium dissectumd 50.6 320.5 327.8 196.4b 265.8 543.8 499.6 199.6 567.5 1342.8 1113.8 180.9

Lomatium triternatumd 181.6 780.1 676.1 177 1637 2830 3195 309.4 1983 2625 2028 502.3b

Lomatium grayi

d 359.9 579.8 686.5 208.4 1036 1144 704.8 NS 570.3 572.7 347.6 NS

Sphaeralcea parvifoliae 285.9 406.1 433.3 NS 245.3 327.3 257.3 NS 81.6 142.5 141.2 NS

Sphaeralcea grossulariifoliae 270.7 298.9 327.0 NS 310.5 351 346.6 NS 224.0 261.9 148.1 NS

Sphaeralcea coccineae 332.2 172.1 263.3 NS 385.7 282.6 372.5 NS 89.6 199.6 60.5 NS

a planted March, 2005, areas of low stand replanted by hand in October 2005. b LSD (0.10). c

planted March, 2005, areas of low stand replanted by hand in October 2005 and whole area replanted in October 2006. Yields in 2006 are based 

on small areas with adequate stand. Yields in 2007 are based on whole area of very poor and uneven stand. d planted March, 2005, whole area replanted in October2005. e

planted spring 2006, whole area replanted in November 2006.
Table 6. Native wildflower seed yield in response to irrigation rate (inches/season) in 2012 - 2014 and 2- to 9-year yield averages. 

Malheur Experiment Station, Oregon State University, Ontario, OR.

Species

2012 2013 2014

0 inch 4 inch 8 inch

LSD

(0.05) 0 inch 4 inch 8 inch

LSD

(0.05) 0 inch 4 inch 8 inch

LSD

(0.05)

Eriogonum

umbellatuma 61.2 153.2 185.4 84.4 113.2 230.1 219.8 77.5 257. 0 441.8 402.7 82.9

Penstemon

speciosusa 103.8 141.1 99.1 NS 8.7 80.7 138.6 63.7 76.9 265.6 215.1 76.7

Lomatium

dissectumd 388.1 460.3 444.4 NS 527.8 959.8 1167 282.4 353.4 978.9 1368 353.9

Lomatium

triternatumd 238.7 603.0 733.2 323.9 153.7 734.4 1051 425.0 240.6 897.1 1497 157.0

Lomatium

grayid 231.9 404.4 377.3 107.4 596.7 933.4 1036 NS 533.1 1418 1241 672.0

2- to 9-year averages

0 inches 4 inches 8 inches

LSD

(0.05)

Eriogonum

umbellatuma 161.4 228.0 239.7 24.9

Penstemon

speciosusa 108.5 174.4 152.7 36.1

Lomatium

dissectumd 358.9 807.9 820.1 142.7

Lomatium

triternatumd 579.0 1193 1311 150.2

Lomatium

grayid 469.6 803.4 746.3 248.3

b

LSD (0.10)

119
120

Table 7, continued on next page. Regression analysis for native wildflower seed yield response 

to irrigation rate (inches/season) in 2006- 2014, and 2- to 9-year averages. For the quadratic 

equations, the amount of irrigation that resulted in maximum yield was calculated using the 

formula: -b/2c, where b is the linear parameter and c is the quadratic parameter. Malheur 

Experiment Station, Oregon State University, Ontario, OR.

Eriogonum umbellatum Maximum

yield

Water applied for 

Year intercept linear quadratic maximum yield R2 P

lb/acre inches/season

2006 137.9 27.8 0.68 0.01 360.3 8.0

2007 79.6 28.3 -1.8 0.69 0.05 194.0 8.1

2008 121.3 34.6 -2.4 0.73 0.01 246.0 7.2

2009 132.3 31.9 -2.3 0.60 0.05 242.9 6.9

2010 252.9 9.2 -1.8 0.08 NS

2011 232.7 -16.0 0.58 0.01 232.7 0.0

2012 71.2 15.5 0.61 0.01 195.2 8.0

2013 113.2 45.2 -4.0 0.62 0.05 241.3 5.7

2014 257.0 74.2 -7.0 0.76 0.01 453.7 5.3

Average 161.4 23.5 -1.7 0.82 0.001 241.9 6.9

Penstemon speciosus Maximum

yield

Water applied for 

Year intercept linear quadratic maximum yield R2 P

lb/acre inches/season

2006 163.5 85.1 -9.9 0.66 0.05 346.4 4.3

2007 2.5 3.2 -0.4 0.48 0.10 9.2 4.2

2008 94.1 113.7 -11.4 0.56 0.05 377.6 5.0

2009 6.8 4.4 -0.5 0.54 0.05 16.1 4.2

2010 147.2 29.8 -2.1 0.35 NS

2011 360.6 -2.8 0.01 NS

2012 103.8 19.3 -2.5 0.30 NS

2013 11.0 16.2 0.77 0.001 141.0 8.0

2014 76.9 77.1 -7.5 0.62 0.05 275.5 5.2

Average 108.5 27.5 -2.7 0.55 0.05 177.2 5.0 a not significant. There was no statistically significant difference in yield between the non-irrigatedplots

and the plots receiving 4 or 8 inches of water.
121

Table 7, continued. Regression analysis for native wildflower seed yield response to irrigation 

rate (inches/season) in 2006 - 2014, and 2- to 9-year averages. Malheur Experiment Station, 

Oregon State University, Ontario, OR.

Lomatium triternatum Maximum

yield

Water applied for maximum 

Year intercept linear quadratic yield R2 P

lb/acre inches/season

2007 3.3 3.1 0.52 0.01 27.7 8.0

2008 195.3 283.9 -16.9 0.77 0.01 1387.6 8.4

2009 181.6 237.4 -22.0 0.83 0.001 822.0 5.4

2010 1637.2 401.5 -25.9 0.83 0.001 3193.2 7.8

2011 1982.9 315.1 -38.7 0.45 0.10 2624.3 4.1

2012 277.8 61.8 0.49 0.05 772.2 8.0

2013 197.7 112.2 0.66 0.01 1095.0 8.0

2014 249.6 157.4 0.97 0.001 1508.6 8.0

Average 579.0 215.7 -15.5 0.84 0.001 1328.2 6.9

Lomatium dissectum Maximum

yield

Water applied for maximum 

Year intercept linear quadratic yield R2 P

lb/acre inches/season

2009 86.4 34.6 0.31 0.10 363.2 8.0

2010 265.8 109.8 -10.1 0.68 0.01 564.2 5.4

2011 567.5 319.3 -31.4 0.86 0.001 1379.2 5.1

2012 402.7 7.0 0.04 NSa

2013 565.3 79.9 0.65 0.01 1204.2 8.0

2014 392.7 126.9 0.82 0.001 1407.9 8.0

Average 358.9 166.9 -13.7 0.91 0.001 869.1 6.1

Lomatium grayi Maximum

yield

Water applied for maximum 

Year intercept linear quadratic yield R2 P

lb/acre inches/season

2007 37.5 12.0 0.26 0.10 133.5 8.0

2008 393.3 315.4 -23.0 0.93 0.001 1474.6 6.9

2009 378.7 40.8 0.38 0.05 705.1 8.0

2010 1035.7 95.3 -17.1 0.22 NS

2011 608.2 -27.8 0.07 NS

2012 231.9 68.1 -6.2 0.66 0.01 418.9 5.5

2013 635.6 55.0 0.25 0.10 1075.3 8.0

2014 533.1 354.0 -33.2 0.64 0.05 1477.1 5.3

Average 469.6 132.3 -12.2 0.55 0.05 827.9 5.4 a not significant. There was no statistically significant difference in yield between the non-irrigatedplots

and the plots receiving 4 or 8 inches of water.
122

Table 8. Amount of irrigation water for maximum native wildflower seed yield, years to 

seed set, and life span. A summary of multi-year research findings, Malheur Experiment 

Station, Oregon State University, Ontario, OR.

Years to first

Species Optimum amount of irrigation seed set Life span

inches/season from fall planting years

Eriogonum umbellatum 0 in wet years, 5 to 8 in dry years 1 8+

Lomatium dissectum 5 in wet years, 8 in dry years 4 8+

Lomatium grayi 0 in wet years, 5 to 8 in dry years 2 8+

Lomatium triternatum average of 6.9 inches over 8 years 2 8+

Penstemon acuminatus no response 1 3

Penstemon speciosus 0 in wet years, 4 in dry years 1 3

Sphaeralcea coccinea no response 1 5

Sphaeralcea grossulariifolia no response 1 5

Sphaeralcea parvifolia no response 1 5
123

D. Irrigation Requirements for Native Wildflower Seed Production for the Second Set of 

Species Planted in the Fall of 2009

Clinton C. Shock, Erik B. G. Feibert, Alicia Rivera, and Lamont D. Saunders, Malheur 

Experiment Station, Oregon State University, Ontario, OR, 2014

Nancy Shaw and Francis Kilkenny, U.S. Forest Service, Rocky Mountain Research Station, 

Boise, ID

Introduction

Native wildflower seed is needed to restore rangelands of the Intermountain West as described 

above. The trials reported here tested the effects of three low rates of irrigation on the seed yield 

of a second set of 11 native wildflower species (Table 1) started with initial plantings in 2009.

Materials and Methods

Plant Establishment

Each wildflower species was planted in four rows 30 inches apart (a 10-foot wide strip) about 

450 feet long on Nyssa silt loam at the OSU Malheur Experiment Station, Ontario, Oregon. The 

soil had a pH of 8.3 and 1.1 percent organic matter. In October 2012, two drip tapes 5 feet apart 

(T-Tape TSX 515-16-340) were buried at 12-inch depth to irrigate the four rows in the plot. Each 

drip tape irrigated two rows of plants. The flow rate for the drip tape was 0.34 gal/min/100ft at 8 

psi with emitters spaced 16 inches apart, resulting in a water application rate of 0.066 inch/hour.

On November 25, 2009 seed of nine perennial species (Table 1) was planted in 30-inch rows 

using a custom-made plot grain drill with disk openers. All seed was planted on the soil surface 

at 20-30 seeds/ft of row. After planting, sawdust was applied in a narrow band over the seed row 

at 0.26 oz/ft of row (558 lb/acre). Following planting and sawdust application, the beds were 

covered with row cover. The row cover (N-sulate, DeWitt Co., Inc., Sikeston, MO) covered four 

rows (two beds) and was applied with a mechanical plastic mulch layer. The field wasirrigated 

for 24 hours on December 2, 2009 due to very dry soil conditions.

Cultural Practices in 2010

After the newly planted wildflowers had emerged, the row cover was removed in April. The 

irrigation treatments were not applied to these wildflowers in 2010. Stands of Penstemon 

cyaneus, P. pachyphyllus, and Eriogonum heracleoides were not adequate for yield estimates.

Gaps in the rows were replanted by hand on November 5. The replanted seed was covered with a 

thin layer of a mixture of 50 percent sawdust and 50 percent hydro seeding mulch (Hydrostraw 

LLC, Manteno, IL) by volume. The mulch mixture was sprayed with water using a backpack 

sprayer. On November 18, 2010, seed of Cleome serrulata was planted as previously described.

Cultural Practices in 2011

Seed from the middle two rows in each plot of Penstemon deustus and Eriogonum heracleoides 

was harvested with a small plot combine. Seed from the middle two rows in each plot of the 

other species was harvested manually.
124

On November 11, 2011, seed of Cleome serrulata was planted as previously described. On 

December 5, 2011, seed of C. lutea was planted as previously described. The Cleome species are 

annuals so they were replanted in new strips of land.

Cultural Practices in 2012

Many areas of the wildflower seed production were suffering from severe iron deficiency early 

in the spring of 2012. On April 13, 2012, 50 lb. nitrogen/acre, 10 lb. phosphorus/acre, and 0.3 lb. 

iron (Fe)/acre was applied to all plots of Lomatium nudicaule, Cymopterus bipinnatus, 

Penstemon deustus, P. cyaneus, and P. pachyphyllus as liquid fertilizer injected through the drip 

tape. On April 23, 2012, 0.3 lb. Fe/acre was applied to all plots of P. deustus, P. cyaneus, P. 

pachyphyllus, Eriogonum heracleoides, Dalea searlsiae, D. ornata, and Astragalus filipes as 

liquid fertilizer injected through the drip tape.

Flea beetles were observed feeding on leaves of Cleome serrulata and C. lutea in April. On April 

29, all plots of C. serrulata and C. lutea were sprayed with Capture® at 5 oz/acre to control flea 

beetles. On June 11, C. serrulata was again sprayed with Capture at 5 oz/acre to control a reinfestation of flea beetles.

A substantial amount of plant death occurred in the Penstemon deustus plots during the winter 

and spring of 2011/2012. For P. deustus, only the undamaged parts in each plot were harvested. 

Seed of all species was harvested and cleaned manually. On October 26, dead P. deustus plants 

were removed and the empty row lengths were replanted by hand at 20-30 seeds/ft. of row. After 

planting, sawdust was applied in a narrow band over the seed row. Following planting and 

sawdust application, the beds were covered with row cover.

On November 1, seed of Cleome serrulata and Cleome lutea was planted on the soil surface at 

20-30 seeds/ft of row. After planting, sawdust was applied in a narrow band over the seed row. 

Following planting and sawdust application, the beds were covered with row cover.

Cultural Practices in 2013

On March 27, row cover was removed and bird netting placed over the Cleome serrulata and 

Cleome lutea plots to protect seedlings from bird feeding. The bird netting was placed over No. 9 

galvanized wire hoops. Bird netting was also placed over Cymopterus bipinnatus on March 29 to 

protect new shoots.

Seed of Penstemon deustus and Eriogonum heracleoides was harvested with a small plot 

combine. Seed of the other species was harvested manually. Weeds were controlled by hand 

weeding as necessary.

Seed of Cleome serrulata was planted on October 31 and seed of Cleome lutea was planted on 

November 15. Seed of both species was planted on the soil surface at 20-30 seeds/ft of row.

After planting, sawdust was applied in a narrow band over the seed row. Following planting and 

sawdust application, the beds were covered with row cover.
125

Cultural Practices in 2014

On March 27, row cover was removed and bird netting placed over the Cleome serrulata and 

Cleome lutea plots to protect seedlings from bird feeding. The bird netting was placed over No. 9 

galvanized wire hoops. Bird netting was also placed over Cymopterus bipinnatus and Lomatium 

nudicaule on March 29 to protect new shoots. On April 29, 0.3 lb Fe/acre was applied through 

the drip tape to all plots of Penstemon deustus, P. cyaneus, P. pachyphyllus, Eriogonum 

heracleoides, and Astragalus filipes.

Seed of Penstemon deustus and Eriogonum heracleoides was harvested with a small plot 

combine. Seed of the other species was harvested manually.

Table 1. Wildflower species planted in the fall of 2009 at the Malheur Experiment 

Station, Oregon State University, Ontario, OR. All species are perennial except Cleome 

serrulata and Cleome lutea.

a recently classified as Cymopterus nivalis S. Watson “snowline springparsley.”

Irrigation for Seed Production

In April, 2011, each strip of each wildflower species was divided into twelve 30-ft plots. Each 

plot contained four rows of each species. The experimental design for each species was a 

randomized complete block with four replicates. The three treatments were a non-irrigated 

check, 1 inch of water applied per irrigation, and 2 inches of water applied per irrigation. Each 

treatment received 4 irrigations that were applied approximately every 2 weeks starting with 

flowering of the wildflowers. The amount of water applied to each treatment was calculated by 

the length of time necessary to deliver 1 or 2 inches through the drip system. Irrigations were 

regulated with a controller and solenoid valves. After each irrigation the amount of water applied 

was read on a water meter and recorded to ensure correct water applications.

The drip-irrigation system was designed to allow separate irrigation of the species due to 

different timings of flowering and seed formation. The three Penstemon spp. were irrigated 

together and the two Dalea spp. were irrigated together. Eriogonum heracleoides and Astragalus 

filipes were irrigated individually. Flowering, irrigation, and harvest dates were recorded (Table 

2). Lomatium nudicaule flowered in 2012; irrigation treatments were applied and seed was 

harvested. Cymopterus bipinnatus had not flowered as of 2012, but differential irrigation 

treatments were applied to Cymopterus bipinnatus starting in 2012. In 2014, after the 4 biweekly irrigations ended, Cleome serrulata and Cleome lutea received an additional 3 bi-weekly

Species Common names Growth Habit

Penstemon deustus Scabland penstemon, hotrock penstemon Perennial

Penstemon cyaneus Blue penstemon Perennial

Penstemon pachyphyllus Thickleaf beardtongue Perennial

Eriogonum heracleoides Parsnipflower buckwheat Perennial

Dalea searlsiae Searls’ prairie clover Perennial

Dalea ornata Western prairie clover, Blue Mountain prairie clover Perennial

Astragalus filipes Basalt milkvetch Perennial

Cleome serrulata 

Cleome lutea 

Lomatium nudicaule

Cymopterus bipinnatus

a

Rocky Mountain beeplant 

Yellow beeplant

Barestem biscuitroot, Barestem lomatium 

Hayden's cymopterus

Annual 

Annual 

Perennial 

Perennial
126

irrigations starting on August 12 in an attempt to extend the flowering and seed production 

period. On August 12, 50 lb N/acre, 30 lb P/acre, and 0.2 lb Fe/acre were applied through the 

drip tape to all plots of Cleome serrulata and Cleome lutea.

Table 2, continued on next page. Native wildflower flowering, irrigation, and seed harvest dates by 

species. Malheur Experiment Station, Oregon State University, Ontario, OR.

Flowering Irrigation

Species start peak end start end Harvest

2011

Penstemon cyaneus 23-May 15-Jun 8-Jul 13-May 23-Jun 18-Jul

Penstemon pachyphyllus 10-May 30-May 20-Jun 13-May 23-Jun 15-Jul

Penstemon deustus 23-May 20-Jun 14-Jul 13-May 23-Jun 16-Aug

Eriogonum heracleoides 26-May 10-Jun 8-Jul 27-May 6-Jul 1-Aug

Dalea searlsiae 8-Jun 20-Jun 20-Jul 27-May 6-Jul 21-Jul

Dalea ornata 8-Jun 20-Jun 20-Jul 27-May 6-Jul 22-Jul

Astragalus filipes 20-May 26-May 30-Jun 13-May 23-Jun 18-Jul

Cleome serrulata 25-Jun 30-Jul 15-Aug 21-Jun 2-Aug 26-Sep

Lomatium nudicaule No flowering

Cymopterus bipinnatus No flowering

2012

Penstemon cyaneus 16-May 30-May 10-Jun 27-Apr 7-Jun 27-Jun

Penstemon pachyphyllus 23-Apr 2-May 10-Jun 27-Apr 7-Jun 26-Jun

Penstemon deustus 16-May 30-May 4-Jul 27-Apr 7-Jun 7-Aug

Eriogonum heracleoides 23-May 30-May 25-Jun 11-May 21-Jun 16-Jul

Dalea searlsiae 23-May 10-Jun 30-Jun 11-May 21-Jun 10-Jul

Dalea ornata 23-May 10-Jun 30-Jun 11-May 21-Jun 11-Jul

Astragalus filipes 28-Apr 23-May 19-Jun 11-May 21-Jun 5-Jul

Cleome serrulata 12-Jun 30-Jun 30-Jul 13-Jun 25-Jul 24-Jul to 30-Aug

Cleome lutea 16-May 15-Jun 30-Jul 2-May 13-Jun 12-Jul to 30-Aug

Lomatium nudicaule 12-Apr 1-May 30-May 18-Apr 30-May 22-Jun

Cymopterus bipinnatus No flowering
127

Table 2, continued. Native wildflower flowering, irrigation, and seed harvest dates by species. Malheur 

Experiment Station, Oregon State University, Ontario, OR.

Flowering Irrigation

Species start peak end start end Harvest

2013

Penstemon cyaneus 3-May 21-May 5-Jun 24-Apr 5-Jun 11-Jul

Penstemon pachyphyllus 26-Apr 21-May 24-Apr 5-Jun 8-Jul

Penstemon deustus 3-May 18-May 15-Jun 24-Apr 5-Jun

Eriogonum heracleoides 29-Apr 13-May 10-Jun 24-Apr 5-Jun 1-Jul

Dalea searlsiae 13-May 15-Jun 8-May 19-Jun 29-Jun

Dalea ornata 13-May 21-May 15-Jun 8-May 19-Jun 28-Jun

Astragalus filipes 3-May 10-May 25-May 8-May 19-Jun 28-Jun

Lomatium nudicaule 11-Apr 20-May 12-Apr 22-May 10-Jun

Cymopterus bipinnatus 5-Apr 15-May 12-Apr 22-May 10-Jun

2014

Penstemon cyaneus 5-May 13-May 8-Jun 29-Apr 10-Jun 14-Jul

Penstemon pachyphyllus 22-Apr 5-May 4-Jun 29-Apr 10-Jun 13-Jul

Penstemon deustus 10-May 20-May 19-Jun 29-Apr 10-Jun 21-Jul

Eriogonum heracleoides 1-May 20-May 12-Jun 29-Apr 10-Jun 3-Jul

Dalea searlsiae 15-May 4-Jun 24-Jun 6-May 17-Jun 1-Jul

Dalea ornata 15-May 4-Jun 24-Jun 6-May 17-Jun 1-Jul

Astragalus filipes 5-May 13-May 28-May 29-Apr 10-Jun 24-Jun

Cleome serrulata 4-Jun 24-Jun 22-Jul 20-May 1-Jul 11-Jul to 30-Jul

Cleome lutea 29-Apr 4-Jun 22-Jul 23-Apr 3-Jun 23-Jun to 30-Jul

Lomatium nudicaule 7-Apr 13-May 7-Apr 20-May 16-Jun

Cymopterus bipinnatus 7-Apr 29-Apr 7-Apr 20-May 16-Jun

Results and Discussion

Precipitation from January through July was higher than average in 2011, close to average in 

2012 and 2014, and lower than average in 2013 (Figure 1). The accumulation of growing degree 

days (50 to 86oF) was below average in 2011, close to average in 2012, and higher than average 

in 2013 and 2014 (Figure 2).

In 2012, a flea beetle infestation on the two Cleome species occurred in April and was controlled 

by insecticide application. Flea beetle feeding occurred earlier in 2013 than in 2012. Upon 

removal of the row cover in March of 2013, the flea beetle damage to the seedlings of Cleome 

serrulata and C. lutea was extensive resulting in full stand loss. Flea beetles were not observed 

on either Cleome species in 2014. Harvested seed pods of Dalea ornata, D. searlsiae, and
128

Astragalus filipes had extensive damage from feeding by seed beetles in 2013 and 2014, 

indicating that control measures during and following flowering will be necessary to maintain 

seed yields.

Penstemon cyaneus

Seed yields did not respond to irrigation in 2011 and 2014 (Tables 3 and 4). In 2012, seed yields 

increased with increasing irrigation up to the highest tested rate of 8 inches. In 2013, seed yields 

showed a quadratic response to irrigation with a maximum seed yield at 4 inches of water 

applied.

Penstemon pachyphyllus

Seed yields did not respond to irrigation in 2011, 2012, and 2014. In 2013, seed yieldsincreased 

with increasing irrigation up to the highest tested of 8 inches.

Eriogonum heracleoides

Seed yields did not respond to irrigation in 2011, 2012, and 2014. In 2013, seed yields showed a 

quadratic response to irrigation with a maximum seed yield at 4.9 inches of water applied.

Dalea searlsiae

In 2011, seed yields were highest with no irrigation. In 2012 and 2014, seed yields showed a 

quadratic response to irrigation. Maximum seed yields were achieved with 6.6 and 8.9 inches of 

water applied in 2012 and 2014, respectively. In 2013, seed yields increased with increasing 

irrigation rate up to the highest tested of 8 inches. Averaged over the 4 years, seed yield showed 

a quadratic response to irrigation with a maximum seed yield at 7 inches of water applied.

Dalea ornata

Seed yields did not respond to irrigation in 2011. Seed yields showed a quadratic response to 

irrigation in 2012, 2013, and 2014 with a maximum seed yield at 7.7, 8.1, and 6.9 inches of 

water applied, respectively. Averaged over the 4 years, seed yield showed a quadratic response 

to irrigation with a maximum seed yield at 6.7 inches of water applied.

Astragalus filipes

Seed yields only responded to irrigation in 2013, when 4 inches of applied water was among the 

irrigation rates resulting in the highest yield (Table 3).

Lomatium nudicaule

Seed yields did not respond to irrigation in the 3 years of testing.

Cymopterus bipinnatus

Seed yields did not respond to irrigation in 2013. In 2014, seed yields increased with increasing 

irrigation rate up to the highest tested of 8 inches.

Cleome serrulata

In 2011, seed yields increased with increasing irrigation up to the highest tested of 8 inches. Seed 

yields did not respond to irrigation in 2012 or 2014. There was no plant stand in 2013 due to 

early severe flea beetle damage. The additional irrigations applied starting on August 12 in 2014

did result in an extension/resumption of flowering, but seed harvested in mid-October was not 

mature.
129

Cleome lutea

Seed yields did not respond to irrigation in 2012 or 2014. There was no plant stand in 2013. 

Early attention to flea beetle control is essential for Cleome seed production. The additional 

irrigations applied starting on August 12 did not result in an extension or resumption of 

flowering.

Figure 1. Cumulative annual and 66-year-average precipitation from January through 

July at the Malheur Experiment Station, Oregon State University, Ontario, OR.

Figure 2. Cumulative growing degree-days (50-86°F ) for selected years and 24-year 

average at the Malheur Experiment Station, Oregon State University, Ontario, OR.
130

Table 3. Native wildflower seed yield response to irrigation rate (inches/season). Malheur Experiment 

Station, Oregon State University, Ontario, OR.

2011 2012

Species 0 inches 4 inches 8 inches LSD (0.05) 0 inches 4 inches 8 inches LSD (0.05)

--------- lb/acre --------- --------- lb/acre ---------

Penstemon cyaneus 857.2 821.4 909.4 NSa 343.3 474.6 581.2 202.6b

Penstemon pachyphyllus 569.9 337.6 482.2 NS 280.5 215.0 253.7 NS

Penstemon deustus 637.6 477.8 452.6 NS 308.7 291.8 299.7 NS

Eriogonum heracleoides 55.2 71.6 49.0 NS 252.3 316.8 266.4 NS

Dalea searlsiae 262.7 231.2 196.3 50.1 175.5 288.8 303.0 93.6

Dalea ornata 451.9 410.8 351.7 NS 145.1 365.1 431.4 189.3

Astragalus filipes 87.0 98.4 74.0 NS 22.7 12.6 16.1 NS

Lomatium nudicaule 53.8 123.8 61.1 NS

Cymopterus bipinnatus

Cleome serrulata 446.5 499.3 593.6 100.9b 184.3 162.9 194.7 NS

Cleome lutea 111.7 83.7 111.4 NS

2013 2014

Species 0 inches 4 inches 8 inches LSD (0. 05) 0 inches 4 inches 8 inches LSD (0. 05)

--------- lb/acre --------- --------- lb/acre ---------

Penstemon cyaneus 221.7 399.4 229.2 74.4 213.9 215.4 215.1 NS

Penstemon pachyphyllus 159.4 196.8 249.7 83.6 291.7 238.6 282.1 NS

Penstemon deustus 356.4 504.8 463.2 NS

Eriogonum heracleoides 287.4 516.9 431.7 103.2 297.6 345.2 270.8 NS

Dalea searlsiae 14.8 31.7 44.4 6.1 60.0 181.4 232.2 72.9

Dalea ornata 28.6 104.6 130.4 38.8 119.4 422.9 476.3 144.1

Astragalus filipes 8.5 9.8 6.1 2.7b 56.6 79.3 71.9 NS

Lomatium nudicaule 357.6 499.1 544.0 NS 701.3 655.6 590.9 NS

Cymopterus bipinnatus 194.2 274.5 350.6 NS 1236 1934 2769 844.7

Cleome serrulata 66.3 80.0 91.3 NS

Cleome lutea 207.1 221.7 181.7 NS

Average 

Species 0 inches 4 inches 8 inches LSD (0.05)

--------- lb/acre ---------

Penstemon cyaneus 409.0 477.7 483.7 NS

Penstemon pachyphyllus 325.4 247.0 316.9 NS

Penstemon deustus 433.7 418.3 396.0 NS

Eriogonum heracleoides 223.1 312.6 254.4 NS

Dalea searlsiae 128.3 183.3 194.0 39.9

Dalea ornata 186.3 325.8 347.4 78.3

Astragalus filipes 43.7 41.7 42.0 NS

Lomatium nudicaule 370.9 426.2 398.7 NS

Cymopterus bipinnatus 715.2 1104 1217 NS

Cleome serrulata 211.3 221.4 263.1 NS

Cleome lutea 159.4 152.7 146.6 NS a not significant, b LSD (0.10)
131

Table 4, continued on next page. Regression parameters for native wildflower seed yield in 

response to irrigation rate (inches/season). For the quadratic equations, the amount of irrigation that 

resulted in maximum yield was calculated using the formula: -b/2c, where b is the linear parameter 

and c is the quadratic parameter. Malheur Experiment Station, Oregon State University, Ontario, 

OR.

Penstemon cyaneus

Year intercept linear quadratic R2 P Maximum yield Water applied for maximum yield

lb/acre inches/season

2011 836.6 6.5 0.01 NSa

2012 855.8 29.7 0.84 0.001 1093.6 8.0

2013 221.7 87.9 -10.9 0.63 0.05 399.4 4.0

2014 214.2 0.1 0.01 NS

Average 419.5 9.3 0.17 NS

Penstemon deustus

Year intercept linear quadratic R2 P Maximum yield Water applied for maximum yield

lb/acre inches/season

2011 615.2 -23.1 0.35 0.05 615.2 0

2012 304.6 -1.1 0.01 NS

2014 356.4 60.8 -5.9 0.26 NS

Average 452.7 -4.1 0.05 NS

Penstemon pachyphyllus

Year intercept linear quadratic R2 P Maximum yield Water applied for maximum yield

lb/acre inches/season

2011 507.1 -11.0 0.04 NS

2012 263.1 -3.3 0.01 NS

2013 156.8 11.3 0.33 0.10 247.2 8.0

2014 275.6 -1.2 0.01 NS

Average 300.7 -1.1 0.01 NS

Eriogonum heracleoides

Year intercept linear quadratic R2 P Maximum yield Water applied for maximum yield

lb/acre inches/season

2011 61.7 -0.8 0.01 NS

2012 271.5 1.8 0.01 NS

2013 287.4 96.7 -9.8 0.64 0.05 525.1 4.9

2014 297.6 27.2 -3.8 0.08 NS

Average 223.1 40.8 -4.6 0.56 0.05 313.4 4.4 a

not significant. There was no statistically significant difference in yield between the non-irrigated plots andthe

plots receiving 4 or 8 inches of water.
132

Table 4, continued. Regression parameters for native wildflower seed yield in response to 

irrigation rate (inches/season). Malheur Experiment Station, Oregon State University, Ontario, OR.

Dalea searlsiae Maximum Water applied for

Year intercept linear quadratic R2 P yield maximum yield

lb/acre inches/season

2011 263.3 -8.3 0.49 0.05 263.3 0

2012 175.5 40.7 -3.1 0.62 0.05 309.3 6.6

2013 15.5 3.7 0.54 0.01 45.1 8.0

2014 60.0 39.1 -2.2 0.79 0.001 234.0 8.9

Average 128.3 19.3 -1.4 0.61 0.05 195.5 7.0

Dalea ornata Maximum Water applied for

Year intercept linear quadratic R2 P yield maximum yield

2011 454.9 -12.5 0.11 NS

2012 145.1 74.2 -4.8 0.65 0.01 431.8 7.7

2013 28.6 25.3 -1.6 0.88 0.001 130.4 8.1

2014 119.4 107.1 -7.8 0.87 0.001 486.6 6.9

Average 186.3 49.6 -3.7 0.76 0.01 353.4 6.7

Astragalus filipes Maximum Water applied for

Year intercept linear quadratic R2 P yield maximum yield

lb/acre inches/season

2011 90.6 -1.6 0.01 NS

2012 19.7 -0.8 0.04 NS

2013 8.5 0.9 -0.2 0.25 NS

2014 56.6 9.4 -0.9 0.08 NS

Average 43.3 -0.2 0.01 NS

Lomatium nudicaule Maximum Water applied for

a

not significant. There was no statistically significant difference in yield between the non-irrigated plots andthe

plots receiving 4 or 8 inches of water.

Year intercept linear quadratic R2 P yield maximum yield

lb/acre inches/season

2012 75.9 0.9 0.01 NS

2013 373.7 23.3 0.1 NS

2014 704.5 -13.8 0.08 NS

Average 384.7 3.5 0.01 NS
133

Table 4, continued. Regression parameters for native wildflower seed yield in response to 

irrigation rate (inches/season). Malheur Experiment Station, Oregon State University, Ontario, OR.

Cymopterus bipinnatus Maximum Water applied for maximum

Year intercept linear quadratic R2 P yield yield

2013 194.9 19.6 0.07 NSa

lb/acre inches/season

2014 1214.6 190.6 0.41 0.05 2739.7 8

Average 761.2 62 8 0.11 NS

Cleome serrulata Maximum Water applied for maximum

Year intercept linear quadratic R2 P yield yield

2011 439.6 18.4 0.35 0. 05

lb/acre

586.7

inches/season

8

2012 175.4 1.3 0.01 NS

2014 66.7 3.1 0.16 NS

Average 206.0 6.5 0.33 NS

Cleome lutea Maximum Water applied for maximum

Year intercept linear quadratic R2 P yield yield

2012 102.4 -0.031 0.01 NS

lb/acre inches/season

2014 207.1 10.4 -1.7 0.20 NS

Average 159.3 -1.6 0.04 NS a

not significant. There was no statistically significant difference in yield between the non-irrigatedplots 

and the plots receiving 4 or 8 inches of water.
134

E. Irrigation Requirements of Native Wildflower Species for Seed Production for theThird 

Set of Species Started in the Fall of 2012

Clinton C. Shock, Erik B. G. Feibert, Alicia Rivera, and Lamont D. Saunders, Malheur 

Experiment Station, Oregon State University, Ontario, OR, 2014

Nancy Shaw and Francis Kilkenny, U.S. Forest Service, Rocky Mountain Research Station, 

Boise, ID

Introduction

Native wildflower seed is needed to restore rangelands of the Intermountain West. Commercial 

seed production is necessary to provide the quantity of seed needed for restoration efforts as 

described above. The trials reported here tested the effects of three low rates of irrigation on the 

seed yield on a third set seven native wildflower species (Table 1) planted first in the fall of 

2012.

Materials and Methods

Plant Establishment

Each wildflower species was planted on 60-inch beds in rows 450 feet long on Nyssa silt loam at 

the OSU Malheur Experiment Station, Ontario, Oregon. The soil had a pH of 8.3 and 1.1 percent 

organic matter. In October 2012, drip tapes (T-Tape TSX 515-16-340) were buried at 12-inch 

depth in the center of each bed to irrigate the rows in the plot. The flow rate for the drip tape was

0.34 gal/min/100 ft. at 8 psi with emitters spaced 16 inches apart, resulting in a water application 

rate of 0.066 inch/hour.

On October 30, 2012 seed of 11 species (Table 1) was planted in either 15-inch or 30-inch rows 

using a custom-made plot grain drill with disk openers. All seed was planted on the soil surface 

at 20-30 seeds/ft. of row. After planting, sawdust was applied in a narrow band over the seed row 

at 0.26 oz/ft of row (558 lb/acre). Following planting and sawdust application, the beds were 

covered with row cover. The row cover (N-sulate, DeWitt Co., Inc., Sikeston, MO) covered four 

rows (two beds) and was applied with a mechanical plastic mulch layer.

Cultural Practices in 2013

Starting on March 29, the row cover was removed. Immediately following the removal of the 

row cover, bird netting was placed over the seedlings to prevent bird feeding on young seedlings 

and new shoots. The bird netting was placed over No. 9 galvanized wire hoops. During seedling 

emergence, wild bird seed was placed several hundred feet from the trial to attract quail away 

from the trials. Bird netting was removed in early May. On July 26 all plots of Machaeranthera 

canescens were sprayed with Capture at 19 oz/acre (0.3 lb ai/acre) for aphid control. On October 

31 seed of Phacelia linearis was planted as previously described.

Due to poor stand, seed of Chaenactis douglasii was replanted on November 1 as previously 

described. Stand of Nicotiana attenuata was extremely poor and seed was unavailable for 

replanting.

Cultural Practices in 2014

Starting on March 27, the row cover was removed from Phacelia linearis and Chaenactis 

douglasii beds. Immediately following the removal of the row cover, bird netting was placed

over the seedlings to prevent bird feeding on young seedlings and new shoots. Bird netting was 

removed in early May.
135

Table 1. Wildflower species planted in the fall of 2012 at the Malheur Experiment Station, 

Oregon State University, Ontario, OR.

Species Common name Longevity Row spacing (inches)

Chaenactis douglasii Douglas' dustymaiden perennial 30

Machaeranthera canescens hoary tansyaster perennial 30

Phacelia hastata silverleaf phacelia perennial 15

Phacelia linearis threadleaf phacelia annual 15

Enceliopsis nudicaulis nakedstem sunray perennial 30

Heliomeris multiflora showy goldeneye perennial 30

Ipomopsis aggregata scarlet gilia perennial 15

Nicotiana attenuata coyote tobacco perennial 30

Thelypodium milleflorum manyflower thelypody biennial 30

Ligusticum porteri Porter's licorice-root perennial 30

Ligusticum canbyi Canby's licorice-root perennial 30

Irrigation for Seed Production

In March 2013, the strip of each wildflower species was divided into twelve 30-ft long plots. 

Each plot contained four rows of each species. The experimental design for each species was a 

randomized complete block with four replicates. The three treatments were a non-irrigated 

check, 1 inch of water per irrigation, and 2 inches of water per irrigation. Each treatment 

received four irrigations that were applied approximately every 2 weeks starting with flowering 

of the wildflowers. The amount of water applied to each treatment was calculated by the length 

of time necessary to deliver 1 or 2 inches through the drip system. Irrigations were regulated 

with a controller and solenoid valves.

The drip-irrigation system was designed to allow separate irrigation of each species due to 

different timings of flowering and seed formation. All species were irrigated separately except 

the two Phacelia spp. and the two Ligusticum spp. Flowering, irrigation, and harvest dates were 

recorded (Table 2) with the exception of Nicotiana attenuata which did not germinate and the 

Ligusticum spp. which did not flower.

Results and Discussion

Precipitation from January through July in 2013 (2.6 inches) was below and in 2014 (5.1 inches) 

was close to the 66-year average of 6.1 inches (Figure 1). The accumulation of growing degree 

days (50 to 86 o

F) was higher than average in 2013 and 2014 (Figure 2).

Stands of Chaenactis douglasii, Ligusticum porteri, and Ligusticum canbyi were poor and 

uneven, not permitting evaluation of irrigation responses. Thelypodium milleflorum is a biennial

and set seed in 2014. Ipomopsis aggregata had very little flowering in 2013, but flowered and

set seed in 2014.

In 2013, seed yields of Machaeranthera canescens showed a quadratic response to irrigation 

with a maximum seed yield at 2.4 inches of water applied (Tables 3 and 4). In 2014, seed yields 
136

of Machaeranthera canescens were not responsive to irrigation. Averaged over the 2 years,seed 

yields of Machaeranthera canescens showed a quadratic response to irrigation with a maximum 

seed yield at 4.1 inches of water applied.

Seed yields of Phacelia hastata showed a quadratic response to irrigation with a maximum seed 

yield at 5.4 and 7.5 inches of water applied in 2013 and 2014, respectively (Tables 3 and 4).

Seed yields of Phacelia linearis showed a quadratic response to irrigation in 2013 with a 

maximum seed yield at 6.2 of water applied. In 2014, seed yields of Phacelia linearis were not 

responsive to irrigation.

In 2013, seed yield of Enceliopsis nudicaulis was very low and was not responsive to irrigation. 

In 2014, seed yield of Enceliopsis nudicaulis showed a quadratic response to irrigation with a 

maximum seed yield at 5.4 inches of water applied.

Seed yield of Heliomeris multiflora increased with increasing irrigation rate up to thehighest 

tested of 8 inches in 2013 and 2014 (Tables 3 and 4).

In 2014, seed yield of Thelypodium milleflorum increased with increasing irrigation rate up to the 

highest tested of 8 inches. In 2014, seed yields of Ipomopsis aggregata were greatest with 4 

inches of applied water.
137

Figure 1. Cumulative annual and 66-year-average precipitation from 

January through July at the Malheur Experiment Station, Oregon State 

University, Ontario, OR

Figure 2. Cumulative growing degree-days (50-86°F ) for selected years 

and 24-year average at the Malheur Experiment Station, Oregon State 

University, Ontario, OR
Table 2. Native wildflower flowering, irrigation, and seed harvest dates in 2013 and 2014 by species. Malheur Experiment Station, 

Oregon State University, Ontario, OR.

Flowering Irrigation

Species start peak end start end Harvest

2013

Chaenactis douglasii 23-May 30-Jun 15-Jul 22-May 3-Jul 2-Jul, 22-Jul

Machaeranthera canescens 13-Aug 1-Oct 17-Jul 28-Aug 2-Oct

Phacelia hastata 17-May 30-Jul 22-May 3-Jul 30-Jul (0 in), 7-Aug, 19-Aug (8 in)

Phacelia linearis 3-May 16-May 15-Jun 2-May 12-Jun 2-Jul

Enceliopsis nudicaulis 30-Jun 15-Sep 3-Jul 14-Aug 8-Aug to 30-Aug

Heliomeris multiflora 15-Jul 30-Aug 5-Jun 17-Jun 8-Aug, 15-Aug, 28-Aug

Ipomopsis aggregata 31-Jul very little flowering 31-Jul 11-Sep

Thelypodium milleflorum No flowering

2014

Chaenactis douglasii 20-May 15-Jul 13-May 24-Jun

Machaeranthera canescens 20-Aug 17-Sep 22-Jul 2-Sep 8-Oct

Phacelia hastata 5-May 10-Jul 29-Apr 10-Jun 14-Jul

Phacelia linearis 5-May 4-Jun 1-Jul 1-May 10-Jun 7-Jul

Enceliopsis nudicaulis 5-May 1-Jul 30-Jul 6-May 17-Jun 14-Jul to 30-Aug

Heliomeris multiflora 20-May 20-Jun 30-Aug 13-May 24-Jun 15-Jul to 15-Aug

Ipomopsis aggregata 22-Apr 13-May 30-Jul 23-Apr 3-Jun 20-Jun

Thelypodium milleflorum 22-Apr 5-May 10-Jun 23-Apr 3-Jun 2-Jul

138
139

Table 3. Native wildflower seed yield response to season long irrigation rate (inches). Malheur 

Experiment Station, Oregon State University, Ontario, OR.

2013 2014

Species 0 inches 4 inches 8 inches

LSD 

(0.05)

0

inches

4

inches

8

inches

LSD 

(0.05)

------------------------------------------ lb/acre ------------------------------------------

Machaeranthera canescens 206.1 215.0 124.3 73.6 326.9 476.6 420.4 NS

Phacelia hastata 35.3 102.7 91.2 35.7 87.7 305.7 366.4 130.3

Phacelia linearis 121.4 306.2 314.2 96.0 131.9 172.9 127.2 NS

Enceliopsis nudicaulis 2.3 6.8 5.9 NS 1.5 34.6 29.1 20.7

Heliomeris multiflora 28.7 57.6 96.9 NS 154.6 200.9 271.7 107.3a

Thelypodium milleflorum 44.0 65.8 152.8 95.7a

Ipomopsis aggregata 47.1 60.9 63.6 9.0

Average

Species 0 inches 4 inches 8 inches

LSD (0.

05)

------------------ lb/acre ------------------

Machaeranthera canescens 266.5 345.8 272.3 NS

Phacelia hastata 61.5 204.2 228.8 53.0

Phacelia linearis 126.7 239.5 220.7 87.2

Enceliopsis nudicaulis 1.9 26.6 17.5 17.1a

Heliomeris multiflora 91.7 129.2 184.3 60.4 a

LSD (0.10)
140

Table 4. Regression parameters for native wildflower seed yield response to irrigation rate 

(inches/season). For the quadratic equations, the amount of irrigation that resulted in maximum yield 

was calculated using the formula: -b/2c, where b is the linear parameter and c is the quadratic parameter. 

Malheur Experiment Station, Oregon State University, Ontario, OR.

Maximum

yield

Water applied

for maximum

a

not significant. There was no statistically significant difference in yield between the forbs that were not irrigated

and those receiving 4 or 8 inches of water.

Species Year intercept linear quadratic R2 P yield

lb/acre inches/season

Machaeranthera canescens 2013 206.1 14.7 -3.1 0.54 0.05 223.4 2.4

2014 326.9 63.2 -6.4 0.34 NSa

Average 266.5 38.9 -4.8 0.41 0.1 345.8 4.1

Phacelia hastata 2013 35.3 26.7 -2.5 0.66 0.01 107.7 5.4

2014 87.7 74.2 -4.9 0.76 0.01 367.4 7.5

Average 61.5 50.5 -3.7 0.88 0.001 233.8 6.8

Phacelia linearis 2013 121.4 68.3 -5.5 0.69 0.01 332.5 6.2

2014 131.9 21.1 -2.7 0.11 NS

Average 126.7 44.7 -4.1 0.48 0.1 247.9 5.4

Enceliopsis nudicaulis 2013 3.1 0.4 0.16 NS

2014 1.5 13.1 -1.2 0.6 0.05 37.1 5.4

Average 1.9 10.4 -1.1 0.46 0.1 27.5 4.9

Heliomeris multiflora 2013 27 8.5 0.38 0.05 95.1 8

2014 150.5 14.6 0.27 0.1 267.6 8

Average 88.7 11.6 0.51 0.01 181.4 8

Thelypodium milleflorum 33.1 13.6 0.38 0.05 141.9 8

Ipomopsis aggregata 48.5 2.1 0.23 NS
141

Publications

Refereed Journal Articles

Shock, C. C.; Feibert, E. B. G.; Shaw, N.; Shock, M.; Saunders, L. D. 2014. Irrigation to enhance 

native seed production for Great Basin restoration. Natural Areas Journal 35(1): 74-82.

Annual Reports

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2014. Direct surface seeding systems 

for successful establishment of native wildflowers, p. 159-165. In: Shock, C. C. (ed.). Oregon 

State University Agricultural Experiment Station, Malheur Experiment Station Annual Report 

2013, Department of Crop and Soil Science Ext/CrS 149.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N.; Sampangi, R. S. 2014. Irrigation 

requirements for native perennial wildflower seed production in a semi-arid environment. P. 166-

183. In: Shock, C. C. (ed.). Oregon State University Agricultural Experiment Station, Malheur 

Experiment Station Annual Report 2013, Department of Crop and Soil Science Ext/CrS 149.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2014. Irrigation requirements for 

native wildflower seed production for perennial and annual species planted in the fall of 2009. p 

184-192. In: Shock, C. C. (ed.). Oregon State University Agricultural Experiment Station, 

Malheur Experiment Station Annual Report 2013, Department of Crop and Soil Science Ext/CrS 

149.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2014. Irrigation requirements for 

annual native wildflower species for seed production, fall 2012 planting. p. 193-197. In: Shock,

C. C. (ed.). Oregon State University Agricultural Experiment Station, Malheur Experiment 

Station Annual Report 2013, Department of Crop and Soil Science Ext/CrS 149.

Shock, C. C.; Feibert, E. B. G.; Saunders, L. D.; Shaw, N. 2014. Tolerance of native wildflower 

seedlings to preemergence and postemergence herbicides. p. 198-206. In: Shock, C. C. (ed.).

Oregon State University Agricultural Experiment Station, Malheur Experiment Station Annual 

Report 2013, Department of Crop and Soil Science Ext/CrS 149.

Presentations

Shock, C. C.; Feibert, E. B. G.; Shaw, N. 2014. Four native wildflower species differ in their 

seed yield response to irrigation. American Society of Horticultural Science annual meeting; 

2014, July 30; Orlando, FL.

Shock, C.; Feibert, E. B. G.; Shaw, N. L.; Kilkenny, F. F. 2014 Successful seed production of 

native Great Basin wildflowers. Society for Ecological Restoration Regional Conference 

Northwest and Great Basin Chapters; 2014 October 6-10; Redmond, OR.

Shock, C.; Feibert, E.; Saunders, M.; Shaw, N. 2014. Wildflower seed yield in response to

irrigation in a dry year. Great Basin Native Plant Project Annual Meeting; 2014 March 17-18; 

Boise, ID.
142

Field days

Feibert, E. B. G.; Shock, C. C. 2014. Establishing native wildflower seed production plantings. 

2014. Native Wildflower Seed Production Field Day, Oregon State University Malheur 

Experiment Station; 2014, May 15; Ontario, OR.

Felix, J.; Shock, C. C. 2014. Controlling weeds in native wildflower seed production plantings. 

Native Wildflower Seed Production Field Day, Oregon State University Malheur Experiment 

Station; 2014, May 15; Ontario, OR.

Shock, C. C.; Feibert, E. B. G. 2014. Plants that use little water. Native Wildflower Seed 

Production Field Day, Oregon State University Malheur Experiment Station; 2014, May 15; 

Ontario, OR.

Shock, C. C.; Feibert, E. B. G. 2014. Irrigation systems for native wildflower seed production. 

Plants that use little water. Native Wildflower Seed Production Field Day, Oregon State 

University Malheur Experiment Station; 2014, May 15; Ontario, OR.
Management Applications and/or Seed Production Guidelines

Irrigation water requirements for native wildflower seed production

Clint Shock and Erik Feibert, 31January 2015

Wildflower species irrigated with drip systems at the Malheur Experiment Station, Oregon State University, Ontario, OR.

Species Common names Flowering dates* Irrigation water

needs, inches/year

Years to first

harvest

Stand life,

years

Seed yield, lb/ac

per harvest

Eriogonum umbellatum Sulphur-flower buckwheat L. May to July 0-8 1 8+ 100-350

Eriogonum heracleoides Parsnip-flowered buckwheat May to June 0-4 2 4+ 50-400

Penstemon acuminatus Sharpleaf penstemon L. April to E. June <4 2 3 100-600

Penstemon deustus Scabland or hotrock penstemon May to E. July 0-4 1 5+ 200-600

Penstemon speciosus Royal penstemon May to June 0-4 2 3 25-350

Penstemon cyaneus Blue penstemon May to June 0-8 1 3 200-800

Penstemon pachyphyllus Thickleaf beardtongue L. April to E. June <4 2 3 200-500

Lomatium dissectum Fernleaf biscuitroot April to E. May 5-8 4 8+ 300-1200

Lomatium triternatum Nineleaf biscuitroot April to E. June 4-8 2 8+ 700-2000

Lomatium grayi Gray’s biscuitroot L. March to E. May 0-5 2 8+ 300-1200

Lomatium nudicaule Bare-stem desert parsley April to May 0-8 3 4+ 500-600

Cymopterus bipinnatus Hayden's cymopterus April to May 4-8 3 4+ 300-2500

Sphaeralcea parvifolia Smallflower globemallow May to June <4 1 4-5 100-500

Sphaeralcea grossulariifolia Gooseberryleaf globemallow May to June <4 1 4-5 150-400

Sphaeralcea coccinea Scarlet globemallow May to June <4 1 4-5 100-300

Dalea searlsiae Searls’ prairie clover May to June 0-8 2 4+ 150-300

Dalea ornata Western prairie clover May to June 0-8 2 4+ 150-450

Astragalus filipes Basalt milkvetch L. April to June <4 2 4+ 10-100

Cleome serrulata Rocky mountain beeplant June to August 0-8 1 1 100-500

Cleome lutea Yellow beeplant May to July <4 1 1 100-200

Machaeranthera canescens hoary tansyaster Mid-Aug to L. Sept. 4 1 2+ 200-400

Phacelia hastata silverleaf phacelia May to July 4-8 1 2+ 100-300

Phacelia linearis threadleaf phacelia E. May to L. June 4 1 1 150-300

Enceliopsis nudicaulis nakedstem sunray May to August 4 1 2+ ?

Heliomeris multiflora showy goldeneye L. May to August 8 1 2+ 50-250

Ipomopsis aggregata scarlet gilia L. April to July 4 2 2+ 50-60

Thelypodium milleflorum manyflower thelypody L. April-mid June 8 2 2+ ?

*Varies with the year and location. E = Early, L = Late

143
144

The preceding report describes irrigation and plant establishment practices that can be 

immediately implemented by seed growers. Multi-year summaries of species seed yield 

performance in response to irrigation are in the table above.

Additional Products

 Seed produced from these plantings was used to establish commercial seed production 

fields.

 A field tour for growers and the public was conducted 15 May 2014; a tour 

of the seed production trials was incorporated into the annual Malheur 

Experiment Station Field Day activities 9 July 2014.

 Continued improvement of native plant database on the internet (at the moment off line):

 http://www.malag.aes.oregonstate.edu/wildflowers/

Acknowledgements

The authors are thankful for support of several grants from the BLM and US Forest Service 

through the Great Basin Native Plant Project and support of the Oregon State University, the 

Malheur County extension service district, and Hatch funds.
145

Project Title Coordination of Great Basin Native Plant Project Plant

Materials Development, Seed Increase, and Use

Project Agreement No. 12-JV-11221632-044

Principal Investigators and Contact Information

Berta Youtie, Owner

Eastern Oregon Stewardship Services

P. O. Box 606 

Prineville, OR 97754 

541.447.8166

byoutie@crestviewcable.com

Project Description

Introduction

In 2014, Eastern Oregon Stewardship Services (EOSS) concentrated on assisting with the 

bluebunch wheatgrass common garden reciprocal transplant study. This study will test the 

hypotheses that local plant material is best adapted for restoration purposes and whetherseed 

zones perform well in delineating regions of local adaptation.

Reciprocal transplant studies, where plants from multiple populations are planted in a set ofsites 

that represent local and non-local climates, are useful for evaluating the effectiveness ofseed 

transfer guidelines and seed zones. Seed zones and guidelines ensure that adapted plant material 

is reintroduced after fires, and may be important for long-term resilience of native grass 

populations.

Objectives

 Prepare and plant bluebunch wheatgrass (Pseudoroegneria spicata) common 

garden study sites in recently burned areas in identified seed zones for reciprocal 

transplant studies.

 Expand the Great Basin Native Plant Project (GBNPP) Native Seed Growers 

Information Network.

 Increase native seed collections for GBNPP in Eastern Oregon and Northern 

Nevada according to research needs and gaps identified by provisional seed zone

maps.

Methods

Seed collections were made in 2013, plugs were grown in greenhouses and bluebunch plugs 

were planted at test sites in fall 2014 (Figure 1). In the Northern Great Basin/Snake River Plain, 

eight or more source treatments were planted within eight test sites with locations based on 

developed seed zones. Another eight sites were planted in the Columbia Plateau/Blue Mt. 

regions.
146

Figure 1. Quinn #1 site in Santa Rosa Mountains in Northern Nevada seed zone 6a.

Many of the test sites had sagebrush removed and were then tilled as part of seed bed 

preparation. Bluebunch wheatgrass plugs were planted into numerically identified study blocks 

consisting of several seed zones in all 16 sites, September through December, 2014 (Figures 2, 3, 

4, 5 and 6). Two bluebunch cultivars, Anatone and Goldar, were also planted at select study 

locations. Two rows of boundary plants were planted as protective borders for the bluebunch 

plugs at each site. After planting, plugs were watered (Figure 7) and plant identification numbers 

were recorded for future data collection (Figure 8).
147

Figure 2. Laying out plots in the Santa Rosa’s. Figure 3. Francis Kilkenny sorting bluebunch plugsfor

planting.

Figure 4. Chris and Jameson digging holes Figure 5. Berta planting bluebunch wheatgrass

for planting grass plugs. plugs.

Figure 6. Crew planting plugs in the Steens. Figure 7. Watering plugs after planting.
148

Figure 8. Brad St. Clair recording plug identification data after planting at Central Ferry, WA.

Results

EOSS assisted with planting six of the study sites (Table 1).

Table 1. EOSS Assistance – Planting Sites

SITES DATES ACTIVITIES

Central Ferry WA September 22-23 Travel and Planting

Stevens Ridge WA September 24-25 Planting and Travel

Quinn #1 NV September 29- October 1 Travel and Planting

Quinn #2 NV October 2-3 Planting and Travel

Steens Mt. OR October 15-17 Travel and Planting

Crooked River Grasslands, OR October 18 Delivering Plants

Management Applications and/or Seed Production Guidelines

Previous work on bluebunch (St. Clair et al. 2013) will allow for selecting populations that are 

more or less adapted to specific climatic variables, as well as allow testing the effectiveness of 

the prospective seed zones.

References

St. Clair, J. B.; Kilkenny, F. F.; Johnson, R. C.; Shaw, N. L.; Weaver, G. 2013. Genetic variation 

in adaptive traits and seed transfer zones for Pseudoroegneria spicata (bluebunch wheatgrass) in 

northwestern Unites States. Evolutionary Applications 6(6): 933-948.
149

Project Title Use of Native Annual Forbs and Early Seral Speciesin

Seeding Mixtures for Improved Success in Great Basin 

Restoration

Project No. 11-IA-11221632-011

Principal Investigators and Contact Information

Keirith A. Snyder, Research Ecologist 

USDA-Agricultural Research Service 

Great Basin Rangelands Research Unit 

920 Valley Road

Reno, NV 89512

775.784.6057 x245, FAX 775.784.1712

keirith.snyder@ars.usda.gov

Shauna M. Uselman, Research Scientist/Temporary Academic 

Faculty

Department of Natural Resources and Environmental Science 

University of Nevada-Reno

Reno, NV 89557

775.784.6057 x237, FAX 775.784.1712

s.uselman@sbcglobal.net

Collaborators

Sara E. Duke, Statistician

USDA-Agricultural Research Service 

Southern Plains Area,

College Station, TX 77845 

979.260.9320, FAX 979.260.9377

sara.duke@ars.usda.gov

Elizabeth A. Leger, Associate Professor

Department of Natural Resources and Environmental Science 

University of Nevada

Reno, NV 89557

775.784.4111, FAX 775.784.4789

eleger@cabnr.unr.edu
150

Project Description

Use of native annual and early seral species in Great Basin rangeland reseeding efforts may 

increase invasion resistance and facilitate succession to desired vegetation, thus improving 

restoration/rehabilitation success. Early serals may be similar to exotic annual grasses in growth 

and resource acquisition strategies. Because they occupy a similar ecological niche, due to 

functional trait similarities such as growth rates and resource acquisition strategies, theory 

predicts that early serals would compete more strongly against invasives like cheatgrass (Bromus 

tectorum) and medusahead (Taeniatherum caput-medusae). As invasion and success of exotic 

grasses appears to be associated with soil type, competitive interactions may also depend on soil 

type.

Objectives

In this study, we used a common garden approach to evaluate the performance of two exotic 

annual grasses (either cheatgrass or medusahead) and mixtures of native species growing in two 

soil types of contrasting texture. We assessed seedling emergence and early survival, as well as 

biomass and reproductive output during the first growing season, because early life stages are 

critical to restoration success. Our first objective was to assess the relative performance of two 

native seed mixes (a novel seed mix composed of early seral species versus a representative 

traditional seed mix composed of late seral species) when seeded in the presence of an exotic 

annual grass. We hypothesized that the early seral seed mix would be more successful in early 

life stages in comparison to the late seral seed mix when growing in the presence of an exotic 

annual grass. Our second objective was to test the potential suppressive effect of the early versus 

late seral seed mix on the early life stages of exotic annual grasses. Here we hypothesized that 

the early seral seed mix would have a greater suppressive effect on the exotic annual grassesin 

comparison to the late seral seed mix, and that this effect would be strongest in the soil type to 

which the exotic is presumed to be least well adapted. Our third objective was to examine the 

performance of exotics and natives in two different soil types contrasting in texture (i.e., a clay 

loam vs. a sandy loam). We hypothesized that the exotics would be more successful when 

growing in the soil type to which the exotic is presumed to be most well adapted. Specifically, 

we predicted that medusahead would be most successful on the clay loam, while cheatgrass 

would be most successful on the sandy loam. For the natives, we hypothesized that performance 

would differ by soil type when growing with exotics due to soil preference by the exotics.

Methods

We used a common garden approach to assess exotic and native plant performance in two soil 

types under the same climatic conditions. Three experiments were performed, using completely 

randomized designs. Two parallel 2x3 factorial experiments, one with each exotic annual grass 

(either cheatgrass or medusahead), were designed to assess the performance of the early seral 

native seed mix versus the late seral seed mix when growing in the presence of an exotic annual 

grass, as well as the relative performance of these two exotic annuals when growing with and 

without natives. In these experiments, two levels of soil type (clay loam and sandy loam) were 

crossed with three levels of seed mix (early seral mix, late seral mix, and no mix). A third 

experiment was designed to evaluate the performance of natives when growing in the absence of 

exotics. In this ‘No Exotic’ experiment, levels of soil type (clay loam and sandy loam) were 

crossed with two levels of seed mix (early seral mix and late seral mix).
151

Figure 1. Photo of one of the replicates in 

the medusahead experiment where several 

individuals of the exotic and the early seral 

natives have emerged, taken during 

emergence monitoring approximately 3 

weeks after seeding.

Each seed mix consisted of two forbs, two perennial grasses, and one shrub. A novel early seral 

mix was composed of the following species: bristly fiddleneck (Amsinckia tessellata), Veatch’s 

blazingstar (Mentzelia veatchiana), squirreltail (Elymus elymoides), Sandberg bluegrass (Poa 

secunda) and rubber rabbitbrush (Ericameria nauseosa), and these seeds were collected from 

multiple wild populations. A late seral mix, representative of traditional seeding mixes used in 

past restoration efforts, was composed of the following species: Palmer’s penstemon (Penstemon 

palmer), goosberryleaf globemallow (Sphaeralcea grossulariifolia), Snake River wheatgrass 

(Elymus wawawaiensis), Indian ricegrass (Achnatherum hymenoides), and Wyoming big 

sagebrush (Artemisia tridentata spp. wyomingensis). These seeds were commercially purchased 

from Comstock Seeds (Gardnerville, NV). A representative clay loam soil (clayey, smectitic, 

mesic, Lithic Argixeroll) and a representative sandy loam soil (fine-loamy, mixed, superactive, 

mesic Durinodic Xeric Haplargid) were collected from multiple field locations, homogenized by 

soil type, and filled into 41-cm deep treepots.

The pots were then sunk into the ground at the 

University of Nevada Agricultural Experiment 

Station in Reno, NV. In each pot, seeds were 

sown by hand into randomized locations 

within each pot (15x15 cm2 surface area) 

using a 20-location fixed grid (Oct. 2010). In 

the ‘No Exotic’ experiment, we used two 

species combinations: (1) early seral native 

species only (10 seeds, consisting of two of

each species) and (2) late seral native species 

only (10 seeds, consisting of two of each 

species). In the two parallel experiments with 

exotic grasses, we used three species 

combinations: (1) exotic species only (10 

seeds), (2) exotic and early seral native 

species together (10+10 seeds), and (3) exotic 

and late seral native species together (10+10 

seeds).

Emergence and seedling survival were monitored biweekly from first emergence (November 

2010) through spring of the following year (May 2011) (Figure 1). We also measured 

aboveground biomass production, seed output, and final establishment though fall 2011. Logistic 

regression analysis was performed on the emergence and survival of exotics and natives in each 

experiment. ANOVA was used to analyze establishment, biomass and seed production. Models 

included seed mix and soil type as factors. For the natives models, functional group (grass, forb, 

shrub) was also included as a factor. Additionally, ANOVA was used to assess treatment 

differences in timing of emergence, both for exotics and for natives, using the same model 

structure described above. Analyses were performed using SAS 9.3 or JMP 9.0 statistical 

analysis software (SAS Institute, Inc., Cary, NC).

Results

Emergence and Survival

We found that early seral species generally outperformed late seral species when growing with 
152

exotic annual grasses, for emergence probabilities (P < 0.0001), survival probabilities (P <

0.05, with medusahead), and earlier emergence timing (P < 0.0001), though results differed

among functional groups and soil types. When growing with cheatgrass, early and late seral

native survival differed in sandy loam but not clay loam (P < 0.01). Within each seed mix, 

native grasses exhibited the highest emergence probabilities of the functional groups (P <

0.0001). In both the cheatgrass and medusahead experiments, we found that the presence of 

native species did not have a suppressive effect on the exotics. In contrast, cheatgrass had a 

higher probability of survival when it was growing with either of the native seed mixes (P <

0.05). A suppressive effect of the natives on the exotics is more likely to be expressed in later 

life stages. In terms of soil preference, we found that cheatgrass performed better in sandy 

loam, as predicted, both for emergence and survival (P < 0.0001).

Contrary to our prediction, medusahead did not have a preference for the clay loam; rather it 

demonstrated superior performance in both soils. Emergence of all native functional groups was 

generally higher on sandy loam when growing with exotics (P < 0.01). We also found that native

grasses had higher survival in sandy loam while forbs and shrubs had higher survival in clay loam 

when growing with exotics (P < 0.0001).

Establishment, Biomass and Reproductive Output in the Medusahead Experiment Establishment 

of medusahead was not affected by seed mix, but was significantly higher in sandy loam than clay

loam(P = 0.003). Medusahead establishment was very high in all treatments and ranged from 83% 

to 93%. When growing with medusahead, establishment of early seral natives was higher than that 

of late seral natives for grasses and forbs but not for shrubs (P = 0.02). Native grasses generally 

exhibited higher establishment than forbs and shrubs. Establishment of grasses was higher in 

sandy loam than clay loam, while establishment of forbs and shrubs was not significantly affected 

by soil type. Both biomass and seed production of medusahead were significantly suppressed by 

the early seral seed mix relative to when the exotic was growing alone, although this effect was 

relatively small. Medusahead biomass and seed production were more strongly reduced in the 

sandy loam (21% and 22%, respectively) than in the clay loam (13% and 14%, respectively). In 

contrast, biomass and seed production of medusahead were not suppressed by the late seral seed 

mix. When growing with medusahead, native aboveground biomass production and seed 

production varied by seed mix, soil type, and functional group (seed mix by soil type by 

functional group interaction P = 0.02).

Both biomass and seed production of early native forbs was higher in clay loam than sandy loam: 

all other native treatment group means were zero or near zero.

Conclusions

Given their generally higher performance, the early seral natives may have a greater chance of 

persisting in the presence of cheatgrass or medusahead in comparison to the late seral natives. 

Success of seeded natives at the seedling stage is likely critical in determining the eventual 

success of native reseeding efforts. Our findings suggest that use of native annuals and early 

serals in reseeding efforts may result in greater density of natives in communities where exotics 

are present, though additional studies will be needed to assess whether these species are able to 

persist in the long-term. Differences in native emergence and survival on the two different soil 

types when growing with exotics suggest that use of early serals in restoration seedings may 

result in greater success in more coarse-textured soils, particularly when native grasses comprise 

a large proportion of the native seed mix. However, when biomass and reproductive output were 
153

measured, the early seral forb, bristly fiddleneck, had a suppressive effect on medusahead on 

both soil types. We also found that performance of medusahead is comparable in the two soil 

types. To date, medusahead invasion has been most closely associated with fine-textured soils, 

but it has been observed on certain sites with more coarse-textured soils (Dahl and Tisdale 1975). 

Our results support these observations and suggest that medusahead is capable of expanding its 

current range in the Intermountain West.

Management Applications and Seed Production Guidelines

This research lends support to the use of early seral natives to improve success rates of rangeland 

restoration/rehabilitation reseedings in the Great Basin. Given their greater performance when 

growing with exotics during early life stages, early seral species may have a greater chance of 

persisting in communities where exotic annual grasses are present. If they persist, and possibly 

suppress exotics once they mature, their establishment may facilitate succession to a more native 

dominant community of desired later seral vegetation.

Our research findings on emergence and survival suggest that it would be advisable to rely more 

on native grasses in Great Basin restoration seedings, based on the relative performance of the 

different native functional groups. Although the forbs in this study did not establish as well asthe 

native grasses, certain species such as bristly fiddleneck show promise for use in rangeland 

reseedings. Bristly fiddleneck had lower establishment, but much greater biomass and seed 

production which produced a suppressive effect on medusahead biomass (reduced by 16%) and 

reproductive output (17%) across both soil types. Further research to examine whether the use of 

increased seeding density of bristly fiddleneck, and/or whether greater diversity of species in the 

seeding mix would enhance exotic suppression, is warranted. Native perennial grasses, especially 

early seral species, were able to establish in the presence of exotic annual grasses in greater 

numbers than native forbs or shrubs, demonstrating their importance in restoration, though they 

did not appear to have a suppressive effect on medusahead during the first growing season.

Additional research is needed to assess the impact of native perennial grasses seeded with exotic 

annual grasses over a longer time frame to determine whether these species may suppressthe 

exotics during later stages of community development. Our findings suggest that efforts to find 

additional novel candidate species for seed mixtures may be best focused on early successional

species, similar to bristly fiddleneck, to improve restoration/rehabilitation outcomes in disturbed 

rangeland ecosystems.

The strong performance of medusahead in both soil types indicate that it is able to expand its 

range beyond fine-textured soils. Therefore, greater efforts to prevent its further spread in the 

Great Basin should be of high priority.
154

References

Dahl, B. E.; Tisdale, E. W. 1975. Environmental factors related to medusahead distribution. 

Journal of Range Management 28: 463-468.

Presentations

Uselman, S. M.; Snyder, K. A.; Leger, E. A.; Duke, S. E. Using annual forbs and early seral 

species in seeding mixtures for improved success in Great Basin restoration. National Native 

Seed Conference; April 8-10, 2013; Santa Fe, NM. Invited talk).

Publications

Uselman, S. M.; Snyder, K. A.; Leger, E. A.; Duke, S. E. 2014. First-year establishment, 

biomass and seed production of early vs. late seral natives in two medusahead (Taeniatherum 

caput medusae) invaded soils. Invasive Plant Science and Management. 7: 291-302. Selected for 

an early press release by the journal.

Uselman, S. M.; Snyder, K. A.; Leger, E. A.; Duke, S. E. Emergence and survival of early vs. 

late seral species in Great Basin restoration during early life stages in two different soil types. In 

Revision: Applied Vegetation Science.

Uselman, S. M.; Snyder, K. A.; Leger, E. A. Comparison of early vs. late seral native biomass 

and seed production when growing with Bromus tectorum. In Preparation.

Products

Outreach: Brochures for the public at the Nevada Agricultural Experiment Station Field Day, 

College of Agriculture, Biotechnology, and Natural Resources, University of Nevada-Reno; 

2013, September 14; Reno, Nevada.

Status of the Current SCA: Completed.
155

Project Title Restoration of Native Understory Plants in Degraded

Sagebrush Steppe Ecosystems

Project Agreement No. L11PG00279 BLM-USGS InteragencyAgreement

Principal Investigators and Contact Information

David A. Pyke, Research Ecologist

U.S. Geological Survey 

3200 SW Jefferson Way 

Corvallis, OR 97331 

541.750.7334

david_a_pyke@usgs.gov

Kari Veblen, Assistant Professor 

Utah State University

5230 Old Main Hill 

Logan, UT 84322

435.797.3970

kari.veblen@usu.edu

Project Description

Objectives

In the Great Basin, efforts to restore native sagebrush steppe plant communities typically begin 

after sagebrush shrubs have been killed (e.g. after wildfires) or mechanically removed to 

facilitate traditional revegetation approaches such as drill seeding. However, it may be feasible to 

begin restoration of native herbaceous plants while the sagebrush canopy is still intact. Our 

overall goal is to evaluate whether native bunchgrass and forb establishment from seeds and 

seedlings differs between sagebrush understories and interspaces in degraded sagebrush steppe 

ecosystems. We are addressing the following questions:

1) Does native bunchgrass and forb establishment differ in sagebrush understories vs. 

interspace gaps?

2) Do these patterns differ for seeds vs. seedlings?

Methods

Three native bunchgrasses: bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Love) 

Thurber’s needlegrass (Achnatherum thurberianum) and squirreltail (Elymus elymoides [Raf.] 

Swezey), and two native forbs: largeflower hawksbeard (Crepis occidentalis Nutt.) and Munro’s 

globemallow (Sphaeralcea munroana [Douglas] Spach), are the focal species for this study. At 

each of the three sites, we established approximately 50 different seeding locations per 

microhabitat (sagebrush understory, interspace) for bluebunch wheatgrass, squirreltail, 

globemallow, and largeflower hawksbeard. We repeated this procedure for seedling plantings 

for bluebunch wheatgrass, squirreltail, globemallow, and a smaller sample of largeflower 

hawksbeard and Thurber’s needlegrass. Seeding occurred in fall 2012, while planting of 

seedlings occurred in spring 2013. We recorded survival, average height, and number of tillers
156

for seeds in spring/summer 2013 and 2014. For seedlings we measured survival (live, dead), 

average height, and basal width in 2014. We also recorded overall site characteristics (e.g. shrub 

densities, plant cover) in 2012 and 2014.

Results and Discussion

Preliminary results suggest that restoration plantings of bluebunch wheatgrass and squirreltail 

from seed showed high (61-74%) establishment success across both canopy and interspace 

microsites after 7 months, but declined markedly after the first year (3-11%). Plantings of 

seedlings also showed high (81-99%) initial establishment rates (after 7 months); though 

establishment of squirreltail, appeared to be greater in interspace than canopy microsites at the 

driest site. Interestingly, this is in contrast to distributions of mature, naturally-established 

squirreltail plants (assessed over 29 sites, including our three project sites) which occur in higher 

densities in canopy microsites, particularly at drier sites. Together these results illustrate how 

plant responses to canopy vs. gap microsites may differ according to species, life stage, and both 

short- and long-term moisture conditions.

Future Plans

We will continue to analyze data, as well as re-assess seeding and seedling parameters in 

spring/summer 2015. We also will begin writing up the data for publication

Management Applications

This was a pilot project so lessons learned and application contain cautions associated with 

studies with few associated with only limited restoration attempts over time and space. However, 

our study does show that perennial grass associations of mature grasses and seedlings with 

sagebrush canopies appears to depend on species life stage (seedling or transplant) and moisture 

conditions. As more information on these relationships are developed, it appears that reestablishing perennial grasses into shrub-dominated communities is possible without waiting for 

fire to remove shrubs. This should allow these communities to recover resiliency so that when 

future fires occur, native perennial herbaceous plants will fill the void and resist annual grass 

invasions.

Presentations

Holthuijzen, M. F; Veblen, K. E.; Pyke, D. A.; Monaco, T. A. Grass-shrub spatial pattern 

response to a moisture gradient in Great Basin sagebrush communities. Ecological Society of 

America meeting; 2014, August 10-15; Sacramento, CA.

Veblen, K. E.; Holthuijzen, M. F.; Wirth, T. A.; DeCrappeo, N.; Pyke, D. A. Plant responses to 

rainfall and microsite in sagebrush communities: keys to restoration success in the Great Basin? 

Ecological Society of America meeting; 2013, August 4-9; Minneapolis, MN.
157

Project Title Science Delivery for the Great Basin Native Plant Project

Project Agreement No. 14-JV-11221632-018

Principal Investigators and Contact Information

Corey Gucker, Outreach Coordinator 

Corey Gucker Company

416 N. Garden St. 

Boise, Idaho 83706

208.373.4342

coreylgucker@fs.fed.us

Project Description

Science integration and science application are integral to the success of the Great Basin Native 

Plant Project (GBNPP). Completed studies in the areas of plant material development, cultural 

practices for seed production, and restoration ecology and technology provide a constantly 

increasing body of knowledge that is published and distributed in many ways: published in 

journals, theses and dissertations, reports, presented through posters, brochures, flyers, 

disseminated through e-mail and meeting networking. Sharing and synthesizing thisinformation 

is essential to rapid and appropriate application in the selection of appropriate plant materialsfor 

planting mixes, commercial seed production, and ecological restoration.

Objectives

The objectives of this project are to promote timely, useful, and user-friendly information to 

GBNPP cooperators and collaborators to advance successful Great Basin restoration efforts.

Results and Discussion

In the second year of this project, procedures and products remain similar to those reported for 

2014. Accomplishments related to the identification and sharing of GBNPP scientific 

developments include:

 Maintaining contact with Great Basin Native Plant Program (GBNPP) cooperators 

through regular correspondence, event alerts, and communication of project due dates.

 Updating and producing hard copy posters and brochures for Great Basin events and 

meetings.

 Regularly updating the GBNPP website with meeting announcements, new publications, 

and updated cooperator information.

 Helping to develop the agenda and secure presenters for the annual GBNPP meeting.

 Tracking and reporting GBNPP accomplishments.
158

Project Title Evaluating Strategies for Increasing Plant Diversity in 

Crested Wheatgrass Seedings

Project Agreement No. 10-CR-11221632-117

Principle Investigators and Contact Information

Kent McAdoo, Assoc. Professor, Natural Resources Specialist 

University Nevada Cooperative Extension

701 Walnut Street

Elko, NV 89801

775.738.1251, FAX 775.753.7843

mcadook@unce.unr.edu

John Swanson, Rangeland Ecologist

College of Agriculture, Biotechnology, and Natural Resources 

University of Nevada

Reno, NV 89512

775.784.1449, FAX 775.784.4583

jswanson@cabnr.unr.edu

Additional Collaborators

Peter Murphy, Biologist

College of Agriculture, Biotechnology, and Natural Resources 

University of Nevada- Reno

Reno, NV 89512

775.784.1449, FAX 775.784.4583

Nancy Shaw, Research Botanist 

USDA Forest Service

Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4360, FAX 208.373.4391

nancylshaw@fs.fed.us

Project Description

Introduction

A previous study by Cox and Anderson (2004) suggested that plant diversity in disturbed, 

formerly sagebrush (Artemisia tridentata)-perennial grass communities, could be increased 

through “assisted succession” by 1) “capturing” the disturbed site from weeds with crested 

wheatgrass (Agropyron cristatum or A. desertorum), 2) reducing crested wheatgrass with 

mechanical or chemical (herbicide) treatments, and 3) seeding the sites with adapted native 

species. Further, Pellant and Lysne (2005) viewed crested wheatgrass monocultures as “bridge” 

plant communities that could replace weed-dominated lands for potential future rehabilitation to 

more diverse plant communities. Recently, research by Hulet et al. (2010) in Utah and Fansler
159

and Mangold (2011) in Oregon has tested this hypothesis, with mixed results. Our research is an 

extension of this work in northern Nevada.

Objectives

Specifically, our objectives were to:

1) Evaluate various treatments for reducing crested wheatgrass in near-monoculture 

rangelands.

2) Determine the effect of crested wheatgrass control methods on establishment ofseeded 

native species.

Methods

Study Site

The study site was located 24 km southeast of Elko, NV, in the Owyhee High Plateau Major 

Land Resource Area (MLRA). The area (1615 m elevation, Loamy 8-10 PZ) had a surface soil 

texture of sandy loam underlain by loam (Orovada Puett association) and wasformerly 

dominated by a Wyoming big sagebrush (Artemisia tridentata spp. wyomingensis) plant 

community. The area was seeded to crested wheatgrass during the 1970s. Located within the 

boundaries of South Fork State Park, the site was fenced to exclude livestock grazing and had the 

necessary cultural resources clearance from the Nevada State Historic Protection Office (SHPO) 

to allow the soil disturbance activity associated with this research.

Treatment Implementation

We conducted two study trials, approximately 0. 9 km apart (center to center), each 10 ha in size 

and installed as a randomized block split-plot design with five blocks. For each of the five

blocks within a trial location, five 0.4-ha plots were divided in half and randomly selected for

either seeding or no seeding. Within each block, the randomly assigned 0.4-ha main plots were

either left “undisturbed” (U) or received a treatment as detailed below. Non-seeded split plots

were included to simulate the outcome of suppression treatments if the seeding treatment failed.

For Trial 1 (north blocks), “disked only” (D) plots were 3-way disked during November 2007. In 

May 2008 “spring-applied glyphosate” (G) and “combined disking + glyphosate” (DG) plots 

were sprayed with glyphosate. In early October 2008, “combined spring + fall-applied herbicide 

plots” (GG) were sprayed again with glyphosate. For Trial 2 (south blocks), another treatment 

suite was implemented within the same general study area during 2010 and using the same 

randomized block, split-plot design. However, because of our poor results with disking and 

similarly poor results reported in a similar study (Fansler and Mangold 2011), we omitted 

disking and included additional herbicide treatments. Specifically, crested wheatgrass 

suppression treatments for Trial 2 consisted of imazapic (I), chlorsulfuron + sulfometuron methyl 

(CS), glyphosate (G), and half-rate glyphosate (G/2).

For each of the trials, sub-plots selected for seeding were seeded at NRCS-recommended rates 

by personnel from the NRCS Aberdeen Plant Materials Center with a Truax Rough Rider 

minimum-till drill during late October 2008 for Trial 1, and late October 2010 for Trial 2. For 

small-seeded species, seed tubes were pulled so that seed fell on the soil surface; drill disks were 

raised, no furrows made, and a brillion-type cultipacker was attached to the rear of the drill to 

press broadcasted seeds into the soil surface. The seed mixture used is identified in Table 1.
160

Table 1. Seeded species and pure live seed (PLS) seeding rates for South Fork study area 

treatments, planted in late October 2008 for Trial 1 and late October 2010 for Trial 2.

Species Cultivar Seed Size Seeding Rate 

(kg PLS /ha)

Indian ricegrass 

(Achnatherum hymenoides)

‘Nezpar’ Large

2.24

Bottlebrush squirreltail 

(Elymus elymoides)

‘Toe Jam Creek’ Large

2.24

Needle-and-thread grass 

(Stipa comata)

Large

2.24

Basin wildrye

(Leymus cinereus)

‘Magnar’ Large

2.24

Bluebunch wheatgrass 

(Psuedoroegneria spicata)

‘Secar’ Large

1.12

Sandberg bluegrass 

(Poa secunda)

Small

0.84

Munro globemallow 

(Sphaeralcea munroana)

Large

0.56

Lewis flax 

(Linum lewisii)

‘Appar’ Large

0.84

Western yarrow 

(Achillea millefolium)

Small

0.22

Wyoming big sagebrush 

(Artemisia tridentata 

wyomingensis)

Small

0.22

Spiny hopsage 

(Grayia spinosa)

Small

0.56

Weather Data and Vegetation Sampling

We compiled precipitation data for South Fork State Park from the National Weather Service. 

During the summers of 2009, 2010, 2011, and 2013 for Trial 1 and 2012 and 2013 for Trial 2, we 

used a stratified random sampling design to measure vegetation parameters: cover and density of 

crested wheatgrass, cover of annual weeds, and density of seeded species. We established five 18

m transects in each subplot, 10 m apart. Along each transect, we measured density and canopy 

cover on ten 0.25-m

2 frames placed at 2 m intervals along each transect. Density of crested 

wheatgrass and seeded species was determined by counting each individual plant within the 

sampling frame; cover was estimated based on modified Daubenmire cover classes (Daubenmire 

1959) as follows: 1) 0-5%, 2) 6-25%, 3) 26-50%, 4) 51-75%, and 5) 76-100%. To determine 

percent cover, we calculated the midpoint of the cover classes and averaged across subplots.

For Trial 1, because of difficulty distinguishing species among seeded perennial grassseedlings 

during the first growing season, we grouped all into seeded perennial grasses, but the seeded forb 

and shrub species were readily identified and were recorded by species. Because of the high

mortality of seeded species that we observed between the first and second growing season for Trial 
161

1, we delayed initial data collection for Trial 2 until the second growing season after seeding.

Statistical Methods

We assessed the effect of treatment, seeding, and year on the density and cover of crested 

wheatgrass using linear mixed models (Mixed procedure, SAS v. 9.4). Trials 1 and 2 were analyzed 

separately because they involved different treatments and sampling years. The fixed effects were 

treatment, seeding, and year and all two-way interactions. The three-way interaction term 

(treatment*seeding*year) was dropped due to lack of significance and poorer model fit when it was 

included (higher AICC). Because of the split-plot design (i.e., seeding was applied to half of each 

treated block), block and treatment-by-block were treated as random effects. Year was a repeated 

effect implemented with unstructured covariance (SAS “type = un”), which produced models with 

better fit (lower AICC) than did compound-symmetric or first-order autoregressive covariance 

models (types “cs” and “ar (1)”). For each crested wheatgrass response, we compared treatment 

means pairwise for both the Trial 1 and Trial 2 blocks, using the Dunn-Sidak method to control for 

multiple comparisons (i.e., maintain test-wide Type 1 error at = 0.05). Cover percentage was

arcsine-squareroot transformed and density was log- transformed

We used the same method as for crested wheatgrass to assess weed cover by species and total 

weed cover. As for crested wheatgrass, weed cover percentages were arcsine-squareroot 

transformed prior to analysis.

For seeded species, we assessed the effect of treatment, year, and treatment-by-year on logtransformed density using repeated-measures analysis of variance. Each species was analyzed 

separately, and we ran analyses for total seeded species, perennials, and forbs. For each response, 

we compared treatment means pairwise for both Trial 1 and Trial 2 blocks, again using the

Dunn-Sidak method to control for multiple comparisons. In the species-specific analyses, a large 

number of zeroes in the Trial 1 blocks for seven species and for all seeded species in the Trial 2 

blocks, led to non-normal residuals, and therefore suspect statistical conclusions.

Results

Precipitation

Precipitation by water year and crop year is summarized in Table 2. Crop year precipitation 

(October through June) varied from a low of 55% of normal (2012) to a high of 124% of normal 

(2011).

Effect of Treatment, Seeding, and Year on Crested Wheatgrass

We found that mechanical treatment (disking) was less successful at reducing crested wheatgrass 

than was herbicide. Among herbicides, G treatment best reduced crested wheatgrass cover 

compared to U. In Trial 1, DG, G, and GG reduced crested wheatgrass cover significantly lower 

than D and U (Figure 1, left). In Trial 2, G and G/2 treatments had lower mean crested wheatgrass 

cover values than the other herbicides (CS and I) and then U (Figure 1, right).

However, crested wheatgrass density responded differently to G application in the two trials. In

Trial 1, G treatment reduced crested wheatgrass density compared to U (Figure 2, left), but in Trial 

2, the two G treatments approximately tripled crested wheatgrass density compared to controls 

(Figure 2, right). The other tested herbicides, CS and I, produced crested wheatgrass cover and 

density means similar to U (Figure 1-2, right).
162

Crested wheatgrass cover increased significantly by year across treatments in Trial 1 (2009 –

2013), while decreasing in Trial 2, but only in the non-G treatments (Figure 1). The consistent 

effect of year on crested wheatgrass across treatments in Trial 1 meant no interaction with 

treatment, while there was a marginal treatment-by-year interaction in Trial 2 (P = 0.102).

Crested wheatgrass density also changed significantly by year, experiencing a spike in 2011 in 

Trial 1, and in 2012 in Trial 2 (Figure 2). The proportional changes in density by year differed by 

treatment in both trials, producing year-by-treatment interactions.

Table 2. Precipitation (mm) by quarter for water year and crop year, 2008-2013, South Fork 

State Park.

Water Yr Oct-Dec Jan-Mar Apr-Jun Jun-Sep Water Yr 

Total

Crop Yr (Oct-Jun) 

Total % of Normal

2008-2009 73 67 133 29 302 274 119

2009-2010 55 57 69 31 212 182 79

2010-2011 140 46 101 15 302 287 124

2011-2012 25 71 31 16 143 127 55

2012-2013 48 51 35 11 145 135 58
163

Figure 2.

Figure 1.
164

Effect of Treatment, Seeding, and Year on Weedy Species

By effectively reducing the competition from crested wheatgrass cover, G treatmentstimulated 

the growth of weedy species. We detected at least five species of weeds in the Trial 1 blocks and 

four in the Trial 2 blocks. Mean weed cover values ranged from 0 – 7% in Trial 1 and 0 – 0.8% 

in Trial 2, with halogeton (Halogeton glomeratus) and exotic mustards (Sisymbrium altissimum

and Descurainia pinnata) typically the most abundant species. Cheatgrass (Bromus tectorum) 

occurred at extremely low densities in both trial areas (mean cover < 0.1% across treatments).

In the Trial 1 blocks, we found significantly higher total weed cover across years in the DG and 

GG treatments than in the D and U treatments (Figure 3, left). Mean weed cover in the G 

treatment fell between these two groups. In the Trial 2 blocks, weed cover was again higher in 

the G treatment than in U, as well as the CS and I herbicide treatments (Figure 3, right). Mean 

weed cover resulting from the G/2 treatment fell between these two groups. In Trial 1, weed 

cover peaked in 2011 (a relatively wet year), and dropped off thereafter, resulting in an effect of 

year (p<0.0001). Moreover, the peak in 2011 was only pronounced in the glyphosate treatments 

(DG, G, GG), resulting in a year-by-treatment interaction (p<0.0001). In Trial 2, an effect of year 

and year-by-treatment (i.e., year in G) was also detected, but of much smaller magnitude (Figure 

3, right).

Effects of Treatment and Year on Seeded Native Species

Herbicide, but not mechanical treatment, increased the density of seeded native species in both 

trials. We detected 10 of the seeded species in the Trial 1 blocks: Wyoming big sagebrush, all 6 

perennial grass species, and all 3 forb species. Seven species were detected in the Trial 2 blocks, 

including Wyoming big sagebrush, four perennial grass species, and two forb species. We found

no evidence of the seeded spiny hopsage in either of the trials.

Across years and treatments, total densities of seeded species ranged from 0-15.7 plants/m2 in

the Trial 1 blocks (grasses: 0-15.4, forbs: 0-1.5) and from 0-1.1 plant/m2 in the Trial 2 blocks 

(grasses: 0-0.8, forbs: 0-0.2). In Trial 1, the most abundant grasses across years were Indian 

ricegrass and bottlebrush squirreltail, and the most abundant forb was Munro globemallow. 

However, by 2013, Indian ricegrass was undetected in the Trial 1 blocks, leaving basin wildrye, 

bottlebrush squirreltail, and Munro globemallow as the most abundant seeded species. Average 

densities of seeded species in the Trial 2 blocks were much lower than in the Trial 1 blocks 

(Figure 4), yet by 2013 the same three species were the most abundant, but only consistently 

present in the G treatment.

Glyphosate treatments, but not disking or other herbicides, increased the density ofseeded 

species in both Trial 1 and Trial 2 by effectively reducing the competitive cover of crested 

wheatgrass. In Trial 1, across years, the mean density of seeded species was significantly higher 

in the glyphosate treatments (DG, G, and GG) than in U (Figure 4, left). In the D treatment, the 

mean total seeded species density, across years, was higher than in the U treatment and lower 

than in the glyphosate treatments (DG, G, and GG), but not significantly when accounting for 

multiple comparisons. In Trial 2, again the G treatment produced higher mean densities of 

seeded species than the other herbicides (CS and I) and U, but the lower dose glyphosate (G/2) 

did not result in significantly higher densities when accounting for multiple comparisons(Figure 

4, right). There were much larger transient effects of sampling year in Trial 1 (note extreme

decreases in seeded species density by year in Figure 4, left), and the year effect was more 
165

pronounced in the D and U treatments, leading to a year-by-treatment interaction.

Figure 4.

Figure 3.
166

Management Applications

The following management applications were derived from thisstudy:

 Disking treatments are ineffective in reducing crested wheatgrass cover and actually 

increase crested wheatgrass density in some cases.

 Imazapic and chlorsulfuron + sulfometuron methyl are ineffective at suppressing 

crested wheatgrass.

 Glyphosate treatments can be effective in reducing crested wheatgrass cover, but 

effectiveness diminishes within a few years as crested wheatgrass rebounds in treated 

areas. Re-treatment may be necessary to reduce competition.

 Reducing crested wheatgrass opens a window for weed invasion.

 After initial establishment of seeded native species, survival may decline steadily in 

successive years as crested wheatgrass rebounds.

Presentations

McAdoo, J. K.; Boyd, C.; Sheley, R. 2014. Transplanting Wyoming big sagebrush in grassdominated sites. Collaborative Restoration Conference, Sponsored by the Great Basin and 

Northwest Chapters, Society for Ecological Restoration Regional Conference; 2014 October 6-

10; Redmond, OR. Abstract published.

McAdoo, K.; Swanson, J.; Shaw, N. 2014. Reestablishing native plant diversity in crested

wheatgrass seedings in northern Nevada. Great Basin Native Plant Annual Meeting; 2014 March 

17-18; Boise, ID.

Publications

A journal manuscript summarizing this research is in preparation and targeted for submission to 

the journal, Ecological Restoration.
167

References

Cox, R. D.; Anderson, V. J. 2004. Increasing native diversity of cheatgrass-dominated rangeland 

through assisted succession. Journal of Range Management 44: 543-549.

Daubenmire, R. F. 1959. Canopy coverage method of vegetation analysis. Northwest Science 33: 

43-64.

Fansler, V. A.; Mangold, J. M. 2011. Restoring native plants to crested wheatgrass stands. 

Restoration Ecology 19: 16-23.

Hulet, A.; Roundy, B. A.; Jessop, B. 2010. Crested wheatgrass control and native plant 

establishment in Utah. Rangeland Ecology & Management 63: 450-460.

Pellant, M.; Lysne, C. R. 2005. Strategies to enhance plant structure and diversity in crested 

wheatgrass seedings. In: Shaw, N. L.; Pellant, M.; Monsen, S. B. (compilers). Proceedings:sagegrouse habitat restoration symposium. Fort Collins, CO, USA: USDA Forest Service RMRS-P38. p. 81-92.
168

Project Title The Ecology of Great Basin Native Forb Establishment

Project Agreement No. 13-JV-11221632-102

Principal Investigators and Contact Information

Jeremy James, Rangeland Specialists

Sierra Foothills Research and Extension Center 

University of California

8279 Scott Forbes Road 

Browns Valley CA

530.639.8803, FAX 530.639.8800

jjjames@ucanr.edu

Project Description

Objectives

Forbs are a key functional group in sage steppe systems yet forb seed is expensive and many forb 

seedings fail, greatly limiting the ability of practitioners to increase native forb density in 

degraded systems (Eiswerth et al. 2009; Fansler and Mangold 2011; Hardegree et al. 2011). Forb 

recruitment is likely limited by a complex set of interacting biotic and abiotic factorsincluding 

competition, drought, and soil physical properties (Humphrey and Schupp 1999; Seabloom et al.

2003). The broad objective of this research is to examine the degree to which crested wheatgrass 

competition and seedbed environmental conditions (soil texture as well as moisture and 

temperature dynamics) influence recruitment of key native forb species. Restoration research has 

traditionally examined the intensity of ecological processes such as competition (Funk et al.

2008). However, there is little effort placed on understanding the relative importance of these 

processes for determining plant community structure, which is ultimately the information 

practitioners want to know. Understanding the relative importance of various ecological 

processes in influencing forb establishment will provide the foundation for improving the ability 

of land managers to capitalize on the array of forb plant material available through the Great 

Basin Native Plant Project (GBNPP) and improve restoration outcomes. This line of work also 

will help us understand how to integrate available plant material with other potential restoration 

practices such as invasive plant control, seedbed preparation, and selection of appropriate spatial 

and temporal windows to conduct management activities to improve restoration outcomes (Boyd

and Davies 2012).

Methods

This project consisted of two phases. The first phase assessed general patterns of survival of 

native grass and forb seed following post fire restoration practices on BLM rangeland. The final 

results of this phase have been published (James et al. 2011) and this final report focuses on the 

methods and results of the second phase, quantifying ecological processes and conditions 

limiting native forb establishment.
169

In the second study phase we seeded two key native forb species across nine sites (Figure 1) in 

the Great Basin that differ in soil texture, annual precipitation and a number of other factorsthat 

influence seed bed abiotic conditions. Sites were seeded in fall 2012 and fall 2013 in 1m x 1m 

plots. Germinated seed and emerged seedlings were tracked through fall, winter, spring and 

summer.

The forbs used for this study include western yarrow (Achillea millefolium var. occidentales) 

and biscuitroot (Lomatium dissectum). Forbs were seeded at the recommended seeding depth

and a rate of 600 PLS m-2 in each plot. Ten plots were seeded for each species at each site and 

halfof these plots also were seeded with crested wheatgrass (Agropyron desertorum) at 600 PLS

m-2.

Crested wheatgrass often is included in restoration species as a bet hedging strategy for 

managers. Depending on drill and seed mixtures these seeds may or may not be planted in the 

same rows as forbs. This crested wheatgrass seeding treatment allows us to quantify the degree 

to which these seeding practices may influence native forb establishment. Forb and crested

wheatgrass seeds (50 seeds bag-1

) were also sown in germination bags (James et al. 2011) in the 

fall of the planting year to quantify germination rates and express seedling emergence rates as a 

function of seeds that germinated in the field (James et al. 2011). Plots were monitored monthly 

in spring for demographic transitions key in seedling recruitment including emergence and 

survival during the first spring and then monthly through the growing season.

We quantified seedbed microenvironmental conditions using the Simultaneous Heat and Water 

model (SHAW model) (Flerchinger and Hardegree 2004; Hardegree et al. 2013). We installed 

soil water and temperature sensors at three depths at each site as well as a micrometeorological 

Figure 1. Location of the nine study sites for phase 2.
170

station to collect the necessary input parameters for the model. Output of the model is a

favorability index (Hardegree et al. 2013) that indicates the relative favorability of the seedbed

to support the temperature and moisture conditions favorable for germination and seedling

growth. Higher numbers indicate more favorable seedbed conditions. Soil texture samples were

collected at multiple depths to develop soil water content/ soil water potential relationships for 

each site.

Results and Discussion

Germination for the native forb species differed by site (p < 0.01) and broadly corresponded to 

mean annual precipitation with wet sites showing higher germination (Figure 2). Crested 

wheatgrass showed higher and less variable germination than the native forbs and high 

germination (> 85%) across all sites. Seeding crested wheatgrass with forbs, however, had no 

effect on native forb germination rates (Figure 3). Emergence was categorically low for both 

forbs regardless of seedbed favorability across the nine sites and, in addition to germination, 

represented a major barrier in the recruitment process. Even in the best case scenario with high 

germination (LODI, wet silt loam site with 80% germination) less than 2% of the germinated 

seeds emerged and survived through the first growing season. This bottleneck also was key 

limitation to crested wheatgrass recruitment where across sites less than 2% of crested 

wheatgrass emerged and survived. Even with relatively high crested wheatgrass densities

(Figure 3, lower panel), seeding crested wheatgrass with forbs had little effect on forb 

recruitment (p > 0.5). The 2012-2013 and 2013-2014 water years (Oct. 1 to Sept. 30) were 

about 30% to 50% below normal across sites with exceptionally dry winter.

Collectively, these results suggest both germination and emergence are major recruitment 

barriers for these two key forb species and that biotic interactions with crested wheatgrass, at 

least at the recruitment stage, are not a major factor limiting forb recruitment. Biscuitroot

appears to have slower seedling growth rates than yarrow, as well as longer cold stratification 

ratesthat may ultimately drive large differences in recruitment between these two species

(Sheley andBates 2008; Scholten et al. 2009). High recruitment rates by yarrow have been 

observed in a number of studies (Sheley and Half 2006; Juran et al. 2008). In the current study, 

even in sites with favorable germination, recruitment never exceeded four plants per square 

meter, and in most cases was less than one plant per square meter. Because seeded yarrow 

tended to initiate germination during late winter, it is likely that the severe winter droughts 

observed across both study years were a major factor contributing to these low recruitment 

rates. Much less is known about how biscuitroot performs in large scale restoration efforts but 

the long cold stratification requirements may have precluded some of the planted seed from 

germinating and emerging.

These plots will be followed over the next winter and early spring to determine if more

favorable subsequent winter time conditions increase recruitment.

The germination and emergence patterns observed in the forb species greatly contrast with 

what is commonly observed among seeded grasses (James et al. 2011). Namely, forbs 

germination and emergence during the first growing season following seeding appear to be 

significant barriers to recruitment. With many grasses, in contrast, as observed here with 

crested wheatgrass, germination is often very high and most grass mortality is associated with 

death of germinated seed prior to emerging above the soil surface. With the forbs examined 

here, low germination, in addition to death of germinated seed, contributed to low recruitment. 
171

Dry Sandy loam

Dry sandy loam

Mid precip silt loam

If managers are trying to target specific time periods for native forb recruitment, it would be 

helpful to examine how large-scale deployment of treatments to break dormancy or accelerated 

rates of seedling development influence restoration outcomes (Menz et al. 2013). An additional 

line of work that could be useful would be to evaluate how seed treatments that alleviate some 

of the winter or spring time environmental stresses could be used to increase germination rates 

and/or speed rates of seedling development (Madsen et al. 2012a, b).

80

80

60

60

40

40

20

20

80

80

60

60

40

40

20

20

80 AGDE ACMI LODI SPCO

60

40

20

AGDE ACMI LODI SPCO

Figure. 2. Field germination of the four study species across the study sites and 

competition treatments (No crested wheatgrass planted = No comp, crested 

wheatgrass planted = Comp). Sites were sown fall 2012 and characterized by mean 

annual precipitation and soil texture from published ecological site descriptions.

Species codes AGDE, ACMI, LODI, and SPCO indicate Agropyron desertorum, 

Achillea millefolium var. occidentales, Lomatium dissectum, and Sphaeralcea 

coccinea, respectively. Germination was quantified using germination bags planted 

with 50 PLS bag-1. Sphaeralcea coccinea seeds were scarified with sulfuric acid 

prior to planting.

Wet silt loam

Wet sandy loam

Germi

n

ati

o

n (%)

No comp

Comp

Germi

n

ati

o

n (%)

No comp

Comp
172

Management Applications and/or Seed

Production Guidelines

Under the conditions of this study, seeding 

crested wheatgrass with forbs had no short term 

negative effects on forb recruitment. As a result, 

managers may be able to simultaneously seed 

forbs and grasses without these two contrasting 

functional groups negatively affecting forb 

recruitment. At sufficient densities there may be 

negative effects of grasses on forbs later as 

plants increase in size but during the initial

phases of restoration this does not seem to be a 

main barrier. For forbs, abiotic limitations that 

either constrain germination or kill germinated 

seed appear to contribute up to 98% of forb 

mortality. Decreasing the time between sowing 

and when seedlings emerge in spring may 

mitigate this mortality to some degree. For plant 

material selection and development, these

results indicate that it is extremely important to 

understand and quantify traits related to 

germination and emergence and that selection 

on these traits may be one way to accelerate our 

ability to increase native forb establishment in 

restoration efforts. Over the long run, seed

coating technologies that buffer seeds from 

environmental stress or accelerate development 

may also be helpful in overcoming these

barriers.

A. desertorum (plants m-2

) A. mellifolium (plants m-2

) L.

dissectum (plants m-2

)

+ A. desertorum

No neighbors 3

2

R2=-0.05

1

R2=-0.11

0

3

+ A. desertorum

No neighbors

2

1 R2

=-0.01

R2=-0.05

0

120

100

80 R2

=-0.07

60

40

20

2.0 2.5 3.0 3.5 4.0

Seedbed favorability index

Figure. 3. Seedling establishment of Lomatium 

dissectum (top panel), Achillea millefolium var. 

occidentales (middle panel), and Agropyron 

desertorum (lower panel) at the end of the first 

growing season in relation to site favorability 

quantified using the SHAW model. Open 

circles and dashed lines are for forbs planted 

with no crested wheatgrass, closed circles and 

solid lines are for forbs planted with crested 

wheatgrass.
173

Presentations

James, J. 2012. Increasing native plant recruitment in dryland systems. University of California; 

2012 October; Berkeley, CA.

James, J. 2012. Restoring native plant diversity in the Great Basin. Sage grouse working group; 

2012 April; Susanville, CA.

Gornish, E. S.; James, J. J. 2014. Demographic assessment of a native bunchgrass speciesfor 

rangeland restoration in the Great Basin. Ecological Society of America Annual Meeting; 2014 

August; Sacramento, CA. http://esa.org/am/

Publications

James, J. J.; Boyd, C. S.; Svejcar, T. 2013. Seed and seedling ecology research to enhance 

restoration outcomes. Rangeland Ecology & Management 66: 115-116.

Svejcar, T.; James, J.; Hardegree, S.; Sheley, R. 2014. Incorporating plant mortality and 

recruitment into rangeland management and assessment. Rangeland Ecology & Management

67: 603-613.

Additional Products

 My 2013 Journal of Applied Ecology paper was selected as Editor’s choice (1 of 6

for 2013) which included a podcast release

 Selected to serve on international dryland restoration advisory committee through Kings 

Park and Botanic Garden, University of Western Australia providing advice on research 

project development and implementation as well as lead and contribute to peerreviewed manuscripts

 Enhancing native plant establishment on Great Basin Rangeland (BLM Agreement

# L14AC00014) 2013-2018

References

Boyd, C. S.; Davies, K. W. 2012. Spatial variability in cost and success of revegetation in a 

Wyoming Big Sagebrush community. Environmental Management 50: 441-450.

Eiswerth, M. E.; Krauter, K.; Swanson, S. R.; Zielinski, M. 2009. Post-fire seeding on Wyoming 

big sagebrush ecological sites: Regression analyses of seeded nonnative and native species 

densities. Journal of Environmental Management 90: 1320-1325.

Fansler, V. A.; Mangold, J. M. 2011. Restoring native plants to crested wheatgrass stands. 

Restoration Ecology 19: 16-23.

Flerchinger, G. N.; Hardegree, S. P. 2004. Modelling near-surface soil temperature and moisture 

for germination response predictions of post-wildfire seedbeds. Journal of Arid Environments 

59: 369-385.
174

Funk, J. L.; Cleland, E. E.; Suding, K. N.; Zavaleta, E. S. 2008. Restoration through reassembly: 

Plant traits and invasion resistance. Trends in Ecology & Evolution 23: 695-703.

Hardegree, S. P.; Jones, T. A.; Roundy, B. A.; Shaw, N. L.; Monaco, T. A. 2011. Assessment of 

range planting as a conservation practice. In: Briske, D. D. (ed.). Conservation benefits of 

rangeland practices: assessment, recommendations, and knowledge gaps. Allen Press, Lawrence, 

KS: 171-212.

Hardegree, S. P. et al. 2013. Hydrothermal assessment of temporal variability in seedbed 

microclimate. Rangeland Ecology & Management 66: 127-135.

Humphrey, D. L.; Schupp, E. W. 1999. Temporal patterns of seedling emergence and early 

survival of Great Basin perennial plant species. Great Basin Naturalist 59: 35-49.

James, J. J.; Svejcar, T. J.; Rinella, M. J. 2011. Demographic processes limiting seedling 

recruitment in arid grassland restoration. Journal of Applied Ecology 48: 961-969.

Juran, C.; Roundy, B. A.; Davis, J. N. (2008) Wildfire rehabilitation success with and without 

chaining on the Henry Mountains, Utah. In: Kitchen, S. G.; Pendleton, R. L.; Monaco, T. A.; 

Vernon, J. (eds.). Shrublands under fire: disturbance and recovery in a changing world. Ft.

Collins, CO: U.S. Department of Agriculture, Forest Service: 91-106.

Madsen, M. D.; Davies, K. W.; Williams, C. J.; Svejcar, T. J. 2012a. Agglomerating seedsto 

enhance native seedling emergence and growth. Journal of Applied Ecology 49: 431-438.

Madsen, M. D.; Kostka, S. J.; Inouye, A. L.; Zvirzdin, D. L. 2012b. Postfire restoration of soil 

hydrology and wildland vegetation using surfactant seed coating technology. Rangeland Ecology

& Management 65: 253-259.

Menz, M. H. M.; Dixon, K. W.; Hobbs, R. J. 2013. Hurdles and opportunities for landscape-scale 

restoration. Science 339: 526-527.

Scholten, M.; Donahue, J.; Shaw, N. L.; Serpe, M. D. 2009. Environmental regulation of 

dormancy loss in seeds of Lomatium dissectum (Apiaceae). Annals of Botany 103: 1091-1101.

Seabloom, E. W.; Harpole, W. S.; Reichman, O. J.; Tilman, D. 2003. Invasion, competitive 

dominance, and resource use by exotic and native California grassland species. Proceedings of 

the National Academy of Sciences of the United States of America 100: 13384-13389.

Sheley, R. L.; Bates, J. D. 2008. Restoring western juniper- (Juniperus occidentalis) infested 

rangeland after prescribed fire. Weed Science 56: 469-476.

Sheley, R. L.; Half, M. L. 2006. Enhancing native forb establishment and persistence using a rich 

seed mixture. Restoration Ecology 14: 627-635.
175

Project Title Presence of Native Vegetation in CrestedWheatgrass

Seedings and the Influence of Crested Wheatgrass on 

Native Vegetation

Project Agreement No. 11-IA-11221632-003

Principal Investigators and Contact Information

Kirk Davies, Rangeland Scientist 

USDA Agricultural Research Service

Range and Meadow Forage Management Research 

67826-A Hwy 205

Burns, Oregon 97720

541.573.4074, FAX 541.573.3042

Kirk.Davies@oregonstate.edu

Aleta Nafus, Graduate Research Assistant 

Department of Rangeland Ecology and Management 

Oregon State University

202 Strand Agricultural Hall 

Corvallis, Oregon 97331

541.737.3341, FAX 541.737.0504

Aleta.Nafus@oregonstate.edu

Project Description

Objectives

The objectives of our study were to:

1. Determine vegetation successional trajectories of native perennial bunchgrasses that were 

co-planted with crested wheatgrass and whether they persist in the community.

2. Determine the variability in native vegetation presence in stands of crested wheatgrass 

and determine what factors (management, site/environmental characteristic, etc.) are 

associated with native plant recruitment in crested wheatgrass (Agropyron cristatum) 

plant communities.

Methods

We sampled 120 crested wheatgrass plant communities in southeastern Oregon to try to 

determine which factors promote native plant recruitment in crested wheatgrass plant 

communities between May through August 2012, and June through September 2013. Sites were 

selected with input from BLM Rangeland Management Specialists to represent a wide range of 

management practices such as timing (spring or fall) and intensity (distance to water/fence lines) 

of grazing, a range of time since seeding, and a variety of environmental factors (elevation, 

aspect and slope; proximity to untreated areas).
176

Slope, aspect, elevation and UTM location were recorded at each plot.

To sample vegetation, 60x50m plots were used to sample each site. In each plot, four 50m 

transects were deployed at 20m intervals along the 60m transect. Along each transect, 

herbaceous vegetation basal cover and density were visually estimated by species inside

40x50cm frames located at 3m intervals on each transect line (starting at 3m and ending at 45m), 

resulting in 15 frames per transect and 75 frames per plot. Percentage ground cover (bare ground, 

rock, litter and crypto-biotic crust) were measured in the 40x50cm frames. Shrub canopy cover 

by species was measured by line intercept. Canopy gaps less than 15cm were included in the 

canopy cover measurements. Shrub density was determined by counting all rooted individualsin 

five, 2x50m belt transects at each plot.

Soil samples were collected at three locations in each plot with a 24 inch hole located at the 

center of the plot and at 15m NW and SE of the center of the plot. From each of these holessoil 

samples were taken from 0-15cm, 0-8 inches, and 8-24 inches. Depth to hardpan was recorded if 

it fell within the 24 inches. The values for each of the samples per plot were averaged into a 

single plot value. Soil texture for the 0-8 inch and 8-24 inch samples was measured using the 

hydrometer method. The 0-15cm samples were used to obtain soil pH, nitrogen, carbon and total 

organic matter for each site.

Actual use data for sites was collected whenever available and for the maximum time period 

available. Disturbance and seeding history are collected for each site whenever possible.

Results and Discussion

Annual forbs occurred in 93%, annual grasses in 89%, Sandberg bluegrass in 85%, shrubs in 

81%, large native perennial forbs in 75%, and large native perennial bunchgrasses (LNPB) in 

59%, of the 121 sites sampled. All sites contained at least one native species. Average 

herbaceous perennial species diversity was 0.6 + 0.05. Total species richness, including shrub 

and herbaceous species, was between 2 and 37 species with an average of 13.2 + 0.8. Squirreltail 

(Elymus elymoides [Raf.] Swezey) was the most common LNPB, occurring on 35% of the sites. 

Wyoming big sagebrush was the most common shrub and occurred on 63% of the sites. LNPB 

and Wyoming big sagebrush occurred together on 49% of the sites while LNPB, sagebrush and 

perennial forbs occurred together on 39% of the sites.

Basal cover and density varied widely among and within herbaceous functional groups (Table 1 

and 2, respectively).

Plant community composition of crested wheatgrass stands was quite variable. Though nearmonoculture areas of crested wheatgrass existed, some stands had large quantities of native 

vegetation, especially shrubs. Correlations between environmental variables and vegetation 

characteristics suggest that some of the variability in the abundance of native vegetation in 

crested wheatgrass stands is likely caused by environmental differences. Big sagebrush wasthe 

most common native species in stands of crested wheatgrass.
177

Table 1. Mean, median and standard error of basal cover of Sandberg bluegrass (Poa secunda), 

crested wheatgrass (Agropyron cristatum), large native perennial bunchgrass (LNPB), annual 

grass, perennial forbs, and annual forbs; foliar cover of shrubs and ground cover of moss and 

biological soil crusts across 121 crested wheatgrass stands sampled in southeastern Oregon.

Mean

Sandberg

bluegrass

Crested

wheatgrass LNPB

Annual

grass

Perennial

forbs

Annual

forbs Shrubs Moss

Cryptobiotic crust

3.3 5.9 0.4 0.3 0.2 0.1 6.0 3.7 1.1

Median 2.4 5.2 0.0 0.1 0.0 0.1 3.4 3.3 0.5

Std err 0.3 0.4 0.1 0.1 0.0 0.0 0.6 0.3 0.2

Table 2. Mean, median and standard error of density (plants. m-2) of functional groups across 121 

crested wheatgrass stands sampled in southeastern Oregon.

Mean

Sandberg 

bluegrass

Crested 

wheatgrass LNPB

Annual 

grass

Perennial

forbs

Annual 

forbs Shrubs

25.9 7.6 1.0 72.2 4.4 77.8 0.5

Median 19.8 7.7 0.1 13.3 0.7 29.3 0.4

Std err 2.4 0.4 0.2 12.9 0.6 11.5 < 0.1

Environmental factors explained as little as 24% of the variability in annual forb density and as 

much as 51% of the variability of annual grass density. Perennial forb basal cover, annual grass 

and Sandberg bluegrass basal cover and density variability were also well explained (43-46%) by 

environmental factors.

Though a variety of environmental factors were included in regression models, soil texture 

appeared to be quite important. Soil texture was included in the best regression models for cover 

and density of all plant functional groups (Tables 3 and 4).

Large native perennial bunchgrasses (LNPB) was the least likely functional group to occur in 

crested wheatgrass communities we sampled across southeastern Oregon. Although 10% of our 

sampled sites had a higher density of native perennial bunchgrasses than crested wheatgrass, we 

found, overall, few LNPB in most stands of crested wheatgrass. This likely confounded our 

ability to find meaningful associations between LNPB and environmental factors. LNPG basal 

cover and density were negatively correlated with sandy soil (Tables 3 and 4).

Perennial forb density and basal cover were, similarly to large native perennial bunchgrasses, 

lower on sandy soil.

Annual forbs, annual grasses, and Sandberg bluegrass were the three most prevalent herbaceous 

functional groups in stands of crested wheatgrass, and all tend to avoid resource limitations by 

growing early in the season when resources are more plentiful (Volire et al. 2009; Ludlow et al. 

1989).
178

Wyoming big sagebrush (Artemisia tridentata Nutt. Ssp. wyomingensis) constituted the majority 

of the shrub cover and density measurements. Similarly to Davies et al. (2007), we did not find a 

strong correlation between Wyoming big sagebrush density, cover, or any of the other measured 

environmental variables. Our data does suggest that, in agreement with Davies et al. (2006, 

2007) and in contrast to Williams (2009), Wyoming big sagebrush was negatively correlated 

with more sandy soils. Plant communities with higher numbers of shrubs, most of which were 

Wyoming big sagebrush, tended to be associated with soils higher in silt and lower in 

precipitation. However, associations were relatively weak as only 25% of the variation in 

Wyoming big sagebrush density was explained by environmental variables. Therefore, soil 

texture may not have much effect on the likelihood of sagebrush reestablishment into crested 

wheatgrass seedings. Although the relationship was not strong, there was a negative correlation

between crested wheatgrass basal cover and Wyoming big sagebrush foliar cover (adj. R2 =0.26, 

p < 0.001). However, there did not appear to be a relationship between Wyoming big sagebrush 

and crested wheatgrass density (R2 < 0.10, p >0.05).

Though we could only explain a small portion of the variability in Wyoming big sagebrush

density and cover, it occurred on 63% of the seedings. Wyoming big sagebrush recovery after 

disturbance can be slow, often taking several decades to a century (Watts and Wambolt 1996; 

Baker 2006; Boyd and Svejcar 2011). Thus, having 15% of the seedings with 10% or higher 

sagebrush cover suggests that sagebrush recovery can occur and provide habitat for sagebrush 

associated wildlife species. At 10% sagebrush cover these seedings meet the minimal sagebrush 

cover requirements suggested by Connelly et al. (2000) for Greater sage-grouse (Centrocercus 

urophasianus) winter habitat. Similarly, McAdoo et al. (1989) reported that sagebrush associated 

species reoccupied crested wheatgrass seedings after sagebrush reestablished.

Most communities seeded to crested wheatgrass were degraded, burned in a wildfire and/or 

treated to remove shrub species prior to seeding (Heady 1988), thus the limited native vegetation 

at some sites may have been an artifact of plant community composition at the time it was 

seeded. The type and severity of disturbance as well as plant composition prior to seeding is 

likely to have affected current plant composition on the site (Burke and Grime 1996).

Future Plans

We are currently analyzing data to determine the relationship between management (i.e. time 

since seeding/fire and grazing history) and the abundance of native vegetation in crested 

wheatgrass stands and whether environmental characteristics of a site interact with management 

history.

Management Applications and/or Seed Production Guidelines

Our data suggests that there are some environmental factors, especially related to soil texture, 

that influence plant community composition of crested wheatgrass seedings regardless of 

management and initial floristics. Our results also suggest that the effects of seeding crested 

wheatgrass likely vary considerably by site characteristics and that native vegetation can cooccupy some rangelands seeded with crested wheatgrass. This information may be useful for 

prioritizing where restoration of sagebrush communities seeded with crested wheatgrassshould 

be applied and assessing the potential effects of seeding crested wheatgrass.
179

Table 3. Multiple linear regression models for functional group cover in crested wheatgrass 

stands for directional chart shown in Table 4. All models shown have adjusted R2 >0.20.

Functional

Group Adj. R2 Best fit regression equation S. E.

Sandberg 

bluegrass 0.46 219.13 + 0.10*Silt – 1.89*Longitude

Crested 

wheatgrass 0.29 -11.89 + 32.84*Nitrogen + 0.77*C:N ratio - 0.096*Silt + 1.76*pH –0.14*Gravel

Sqrt(LNPB) 0.24 2.51 – 0.0066*Sand – 0.23*pH + 0.019*Gravel

Sqrt(Annual

grass) 0.46 1.08 – 0.0071*Sand – 0.013*Silt – 0.00056*Elevation + 0.025*Litter

Sqrt(Perennial 

forb) 0.43 2.95 + 2.16*Nitrogen – 0.0050*Sand – 0.36*pH – 0.0052*Litter

Sqrt(Shrubs) 0.26 11.59 – 0.021*Sand + 0.0022*Elevation – 1.01*pH – 0.022*Litter –

0.015*Precipitation

Sqrt(Wyoming

big sagebrush) 0.25 15.18 + 0.79*Carbon – 1.46*pH – 0.036*Litter – 0.013*Precipitation

Table 4. Multiple linear regression models for functional group density in crested wheatgrass 

stands for directional chart shown in table 3. All models shown have adjusted R2 >0.20.

Functional

Group Adj. R2 Best fit regression equation

S.

E.

Sandberg 

bluegrass 0.49 1291.38 + 0.19*Silt + 11.81*Longitude

Sqrt(LNPB) 0.31 3.50 -0.0085*Sand – 0.41*pH + 0.046*Gravel cover

Sqrt(Annual

grass) 0.51 8.08 - 0.074*Silt – 0.010*Elevation + 0.40*Litter cover –0.13*Gravel cover

SqrtPerennial

forbs) 0.43 12.02 – 0.040*Sand – 1.31*pH

Sqrt(Annual

forbs) 0.24 299.81 + 0.15*Clay + 2.56*Longitude + 0.16*Bare ground + 0.29*Rock cover

Sqrt(Shrubs) 0.24 3.19 – 0.0044*Sand - 0.00055*Elevation - 0.30*pH – 0.0084*Litter –

0.0032*Precipitation

Sqrt(Wyoming

big sagebrush) 0.25 2.79 – 0.0034*Sand - 0.28*pH - 0.012*Litter
180

Publications

Nafus, A. M.; Svejcar, T. J.; Ganskopp, D. C.; Davies, K. W. [In press]. Abundances of coplanted native bunchgrasses and crested wheatgrass after 13 years. Rangeland Ecology and 

Management.
181

Project Title Seeding Native Species Following Wildfire inWyoming

Big Sagebrush (Artemisia tridentata ssp.wyomingensis): 

Effects of a New Fire Incident

Project Agreement No. 11-IA-11221632-077

Principal Investigators and Contact Information

Jeff Ott, Post-doctoral Research Geneticist 

USDA Forest Service

Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4360, FAX 208.373.4360

jeott@fs.fed.us

Nancy Shaw, Research Botanist (Emeritus) 

USDA Forest Service

Rocky Mountain Research Station 

322 E. Front Street, Suite 401

Boise, ID 83702

208.373.4360, FAX 208.373.4360

nancylshaw@fs.fed.us

Additional Collaborators

Mike Pellant Robert Cox

USDI Bureau of Land Management Department of Natural Resources Management 

Idaho State Office Texas Tech University

1387 Vinnell Way Box 42125

Boise, ID 83702 Lubbock, TX 79409

208.373.3823, FAX 208.373.3805 806.742.2280, FAX 806.742.2841

mike_pellant@blm.gov robert.cox@ttu.edu

Dennis Eggett Bruce Roundy

Department of Statistics Department of Plant and Wildlife Sciences

233A TMCB 463 WIDB

Brigham Young University Brigham Young University

Provo, UT 84602 Provo, UT 84602

801.422.4199, FAX 801.422.0635 801.422.8137

theegg@byu.edu Bruce_Roundy@byu.edu

Partial funding provided by the Joint Fire Sciences Program and the National Fire Plan.
182

Project Description

Objectives

A post-fire seeding experiment was implemented across four sites in the northern GreatBasin 

beginning in 2007. The experiment was designed to compare the effectiveness of different 

seeding techniques for establishing native plants and suppressing exotic annual weeds in burned 

Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) communities. Previousreports 

have presented results based on the first 2 years of vegetation monitoring following treatment. 

Additional data have been collected from some sites during the third or fifth year. In this report, 

we highlight third-year monitoring results for a site that partially re-burned between the second 

and third year.

Methods

Study Site and Experimental Treatments

The Saylor Creek study site is located at 42o39'43" N, 115o28'18" W, ca. 1200 m elevation in 

Elmore County, Idaho. Mean annual precipitation is 223 mm/year (Centre for Forest 

Conservation Genetics 2014) and surface soil texture ranges from loam to silt loam to very fine 

sandy loam (Soil Survey Staff 2015). The Black Butte wildfire burned through mature Wyoming 

big sagebrush at this site on 29 June 2010. Experimental treatments were applied the following 

fall (27-28 Oct. 2010) or winter (15 Feb. 2011) in five blocks that were fenced to exclude 

livestock. Within blocks, each treatment was randomly assigned to a 30 m × 70 m rectangular 

plot.

Treatments differed by seeding method (rangeland drill vs. aerial broadcast), drill type 

(conventional vs. minimum-till), aerial broadcast timing (fall vs. winter) and sagebrush seeding 

rate (50 vs. 250 vs. 500 pure live seed/m2

). Rangeland drills were configured to place large seeds 

(drill mix) and small seeds (broadcast mix) in alternate rows (Table 1). The drill mix passed 

through the drill assembly and was placed in furrows, while the broadcast mix was allowed to 

fall to the soil surface of the intervening rows. Aluminum pipes 7.6 cm in diameter were added 

to the conventional drill to direct the broadcast mix to the soil surface where seeds were covered 

by drag chains. The minimum-till drill was equipped with imprinter units to press the broadcast 

mix into the soil. Aerial seeding was simulated by hand-seeding the broadcast mix on plots that 

had been drill-seeded in alternate rows with the broadcast seed boxes empty. Drill treatments 

applied with empty seed boxes for both mixes served as non-seeded controls in addition to a

fully untreated (non-drilled/non-seeded) control.

Detailed comparisons of treatment effects on plant establishment and survival were presented in 

previous years’ reports. In the current report we simplify our analysis by merging treatmentsinto 

two categories, seeded and non-seeded. Previous reports focused on data from the first two years 

following treatment, but here we add data from a third year so that the effects of a second 

wildfire can be evaluated. The Kinyon Road fire burned through the Saylor Creek site in early 

July 2012, shortly after second-year data collection. It burned irregularly leaving a mosaic of

burned and unburned sections.
183

Table 1. Seed mixes for post-fire seeding at Saylor Creek site.

Common Name Scientific Name Source/Cultivar/Germplasm PLS*(%) PLS*/m2

Drill Mix

Bluebunch wheatgrass Pseudoroegneria spicata Anatone Germplasm 80.0 60

Bottlebrush squirreltail Elymus elymoides Emigrant Germplasm 97.0 35

Indian ricegrass Achnatherum hymenoides 'Rimrock' 96.6 50

Thurber needlegrass Achnatherum thuberianum Snake River Plain - pooled 55.4 30

Needle-and-thread Hesperostipa comata Millard Co., UT 77.6 20

Munro’s globemallow Sphaeralcea munroana Uintah Co., UT (1550 m) 61.0 40

Basalt milkvetch Astragalus filipes Deschutes Co., OR (1330 m) 90.0 14

Broadcast Mix

Sandberg bluegrass Poa secunda Mountain Home Germplasm 82.0 100

Wyoming big sagebrush A. tridentata spp. wyomingensis Power Co., ID (1390 m) 28.4 50-500**

Rubber rabbitbrush Ericameria nauseosa Utah Co., UT (1650 m) 40.5 85

Western yarrow Achillea millefolium Eagle Germplasm 94.3 100

Royal penstemon Penstemon speciosus Northern Great Basin - pooled 57.8 15

*Pure live seed

**Big sagebrush seeding rate varied by treatment (50, 250, or 500PLS/m2

)

Data Collection and Analysis

Vegetation data were collected using protocols modified from Herrick et al. (2005) and Wirth 

and Pyke (2007). Five 20m transects were established 10m apart, perpendicular to the long axis 

of each plot. Canopy cover by species was recorded by line point intercept at 20 points along 

each transect in late spring of the first three years following treatment (2011-2013). Density of 

seeded forbs and shrubs was recorded in belts 2m wide centered on transects during the first and 

third year. To the extent that newly-burned patches could be discerned in the third year, each 

cover point and density count was categorized as burned or non-burned (or

borderline/uncertain). The percentage of belt areas within these categories was estimated so that 

densities by burn category could be calculated.

Differences between treatments (simplified to seeded and non-seeded) and years were analyzed 

using general linear mixed effects models in SAS 9.3 (SAS Institute Inc. 2011) with treatment 

and year as fixed effects and block as a random effect. Mean comparisons were applied using 

Tukey’s HSD at alpha = 0.05.

Using third-year data from the seeded treatment, deviations from expected cover or density in 

newly-burned vs. non-burned categories were analyzed using statistical tests in R (R Core

Team 2012). We used binomial proportion tests to compare observed versus expected cover 

proportions (canopy hits: total points) and chi-square contingency table tests to analyze density 

(density count and belt area on one table axis; burned and non-burned categories on the other).

Results and Discussion

Treatment Comparisons (Seeded vs. Non-seeded) Across Three Years

Seeded species generally had low cover in 2011, the first year following fire and seeding 

treatment, then increased dramatically in cover by 2012 (Figure 1A-B). By 2013, following the 

second (partial) burn, cover of most seeded species decreased to levels intermediate between
184

2011 and 2012 (Figure 1B). Each year, cover of seeded species was higher in seeded than nonseeded treatments, with the exception of Sandberg bluegrass (Poa secunda), which was abundant 

in non-seeded treatments as fire-surviving residual plants (Fig 1B). Seeding clearly had a 

significant effect on plant establishment which persisted even in the aftermath of the second fire.

Like seeded species, cover of non-seeded species categories (native perennials, native annuals, 

cheatgrass and exotic annual forbs) generally increased between 2011 and 2012 then decreased 

in 2013 (Figure 1A). However, cheatgrass cover decreased between 2012 and 2013 only in the 

seeded treatments, and was significantly higher in non-seeded than seeded treatments in both 

2012 and 2013 (Figure 1A). Exotic annual forbs (primarily tumblemustard, Sisymbrium 

altissimum) had minimal cover except in the non-seeded treatment in 2012. These results provide 

evidence that seeded plants competitively suppressed cheatgrass and exotic annual forbs in the 

seeded treatments.

Of the seeded forbs and shrubs counted in belt transects, the species with the highest first-year 

(2011) establishment in seeded treatments (Table 2) were western yarrow (Achillea millefolium; 

3940 plants/acre), royal penstemon (Penstemon speciosus; 2,860 plants/acre) and Wyoming big 

sagebrush (2,333 plants/acre). By the third year (2013), density had dropped by 38% for western 

yarrow, 51% for royal penstemon and 82% for Wyoming big sagebrush. Munro’s globemallow 

(Sphaeralcea munroana) and rubber rabbitbrush (Ericameria nauseosa) had comparatively 

lower first-year densities of 140-282 plants/acre, but their densities had increased to 219-624 

plants/acre by the third year (Table 2). Basalt milkvetch (Astragalus filipes) began at 225 

plants/acre then declined to 143 plants/acre during this period (Table 2).

Effects of 2012 Fire on 2013 Seeded Treatments

Declines in density and cover from 2011-2012 to 2013 (Figure 1, Table 2) might have been 

caused by the intervening fire, but other factors such as precipitation patterns could have also 

have played a role. Statistical tests comparing newly-burned versus non-burned sections of the 

2013 seeded treatments allowed us to assess the influence of the fire relative to other factors. 

Species with higher than expected density/cover in non-burned sections can be inferred to have 

been negatively impacted by the fire, while those that were higher than expected in burned 

sections can be inferred to have been positively impacted by the fire even if they had decreased 

relative to previous years.

Wyoming big sagebrush, rubber rabbitbrush, western yarrow and Sandberg bluegrass were 

higher than expected in non-burned sections, suggesting that the fire had a net negative impact 

on these species (Tables 3, 4). The contrast between observed and expected was most

pronounced for Wyoming big sagebrush (Table 3), which is known for its limited post-fire

resprouting capacity. Royal penstemon and Indian ricegrass (Achnatherum hymenoides) showed 

the opposite pattern of higher than expected density or cover in burned sections (Tables 3, 4), 

suggesting a stimulatory effect of the fire on regrowth and/or seed germination. Bluebunch

wheatgrass (Pseudoroegneria spicata), bottlebrush squirreltail (Elymus elymoides), Munro’s

globemallow and basalt milkvetch occurred in both burned and non-burned sections without 

higher than expected abundance in one or the other (Tables 3, 4). Thurber’s needlegrass

(Achnatherum thuberianum) and needle-and-thread (Hesperostipa comata) were recorded only 

in burned sections, but were too uncommon for reliable statistical inference.
185

Figure 1. Canopy cover by treatment (seeded vs. not), year (2011-2013) and species/category at 

the Saylor Creek site. The entire site burned in 2010 and partially reburned in 2012; reburned 

sections are not differentiated here. Within cells, means with the same letter were not 

significantly different (p < 0.05). Year differences are shown separate from treatment differences 

in cases where the interaction was non-significant.
186

Table 2. Densities of seeded forbs and shrubs in seeded treatments at 

the Saylor Creek site in 2011 (first year following seeding) and 2013 

(following new fire; reburned and non-burned sections not 

differentiated).

2011 2013 Higher Year**

Basalt milkvetch 225 ± 40* 143 ± 26 2011

Munro's globemallow 282 ± 46 624 ± 100 2013

Wyoming big sagebrush 2333 ± 741 404 ± 129 2011

Rubber rabbitbrush 140 ± 16 219 ± 25 2013

Western yarrow 3940 ± 683 2433 ± 422 2011

Royal penstemon 2860 ± 497 1378 ± 240 2011

*values shown are plants/acre ± standard error

**year with significantly higher density (p < 0.05)

Table 3. Results of chi-square tests comparing densities of seeded forbs and shrubs in 

burned and non-burned sections of seeded treatments at the Saylor Creek site. Observed 

values are counts within a total area of 10,000 m

2

.

Burned Non-burned Higher than

Observed(Expected) Observed(Expected) p value Expected: 

Basalt milkvetch 341(330) 64(75) 0.1626

Munro's globemallow 1305(1290) 279(294) 0.3255

Wyoming big sagebrush 141(770) 885(256) <0.0001 Non-burned

Rubber rabbitbrush 424(476) 165(113) <0.0001 Non-burned

Western yarrow 4273(4752) 1937(1458) <0.0001 Non-burned

Royal penstemon 3195(2963) 348(580) <0.0001 Burned

Among non-seeded species, perennial western wheatgrass (Pascopyrum smithii) had higher than 

expected cover in burned sections, while the annuals western tansymustard (Descurania pinnata) 

and clasping pepperweed (Lepidium perfoliatum) had higher than expected cover in non-burned 

sections (Table 4). Although live cheatgrass cover did not differ from expected between burned 

and non-burned sections, litter cover (dominated by dead cheatgrass from previous years) was 

proportionally lower in the burned sections (Table 4).

Future Plans

Data will be re-collected from the Saylor Creek site in 2015, adding to fifth-year data that has 

already been collected from other sites of the broader study. These data will be used to evaluate 

post-fire seeding effectiveness beyond the typical 1-3 year monitoring time frame.

Management Applications

This study demonstrates that post-fire seeding with native species can be successful onWyoming 

big sagebrush sites with characteristics similar to the Saylor Creek site. Seeded native perennials

can compete with exotic annuals leading to lower abundance of exotics than in the absence of 
187

seeding. Once established, native seedings have the capacity to rebound following fire even 

though they may not immediately return to pre-burn levels of cover and density. Species 

demonstrating high resilience to fire should be sought for seeding in fire-prone areas. Species 

that are more sensitive to fire, such as Wyoming big sagebrush, may need to be reseeded 

following each fire episode.

Table 4. Results of binomial proportion tests comparing cover of prominent 

species and cover categories in burned and non-burned sections of seeded 

treatments at the Saylor Creek site. Proportions shown are hits/total points within 

burned and non-burned.

Publications

Daniels, Amy; Shaw, Nancy; Peterson, Dave; Nislow, Keith; Tomosy, Monica; Rowland, Mary. 

2014. Facing climate change in forests and fields: U. S. Forest Service taps into sciencemanagement partnerships. The Wildlife Professional. (Spring 2014): 31-35.

Kiehl, Kathrin; Kirmer, Anita; Shaw, Nancy; Tischew, Sabine. Guidelines for native seed 

production and grassland restoration. Cambridge Scholars Publishing. Newcastle upon Tyne, 

UK: Cambridge Scholars Publishing. 315 p.

Kiehl, Kathrin; Kirmer, Anita; Tischew, Sabine; Shaw, Nancy. 2014. Chapter 1: Native seed 

production and grassland restoration. Introduction. In: Kiehl, Kathrin; Kirmer, Anita; Shaw, 

Nancy; Tischew, Sabine. Guidelines for native seed production and grassland restoration.

Cambridge Scholars Publishing. Newcastle upon Tyne, UK: Cambridge Scholars Publishing. p. 

2-13.

Burned Non-burned p value

Higher than

Expected: 

Native seeded perennials 0.221 0.229 0.6536

Bluebunch wheatgrass 0.101 0.099 0.8640

Bottlebrush squirreltail 0.011 0.003 0.0666

Indian ricegrass 0.027 0.002 <0.0001 Burned

Sandberg bluegrass 0.061 0.095 0.0003 Non-burned

Western yarrow 0.014 0.040 <0.0001 Non-burned

Native non-seeded perennials 0.089 0.055 0.0012 Burned

Western wheatgrass 0.082 0.047 0.0006 Burned

Native annuals 0.005 0.014 0.0110 Non-burned

Western tansymustard 0.005 0.014 0.0110 Non-burned

Exotic annual grass (cheatgrass) 0.330 0.336 0.7714

Exotic annual forbs 0.002 0.023 <0.0001 Non-burned

Clasping pepperweed <0.001 0.018 <0.0001 Non-burned

Tumblemustard <0.001 0.001 0.7944

Litter 0.267 0.563 <0.0001 Non-burned
188

Shaw, Nancy; Jensen, Scott. 2014. Chapter 4. 2: Restoration of Great Basin ecosystems using 

native seed. In: Kiehl, Kathrin; Kirmer, Anita; Shaw, Nancy; Tischew, Sabine. Guidelines for 

native seed production and grassland restoration. Cambridge Scholars Publishing. Newcastle 

upon Tyne, UK: Cambridge Scholars Publishing. p. 141-159.

Shock, Clinton C.; Feibert, Erik B. G.; Shaw, Nancy L.; Shock, Myrtle P.; Saunders, Lamont D. 

in press. Irrigation to enhance native seed production for Great Basin restoration. Natural Areas 

Journal.

Tischew, Sabine; Kirmer, Anita; Kiehl, Kathrin; Shaw, Nancy. 2014. Chapter 5. 1. Planning and 

implementation of restoration projects using native seed and plant material. In: Kiehl, Kathrin; 

Kirmer, Anita; Shaw, Nancy; Tischew, Sabine. Guidelines for native seed production and 

grassland restoration. Cambridge Scholars Publishing. Newcastle upon Tyne, UK: Cambridge 

Scholars Publishing. p. 286-300.

Taylor, Megan M.; Hild, Ann L.; Shaw, Nancy L.; Norton, Urszula; Collier, Timothy R. 2014. 

Plant recruitment and soil microbial characteristics of rehabilitation seedings following wildfire 

in northern Utah. Restoration Ecology. 5 p. doi: 10. 1111/rec. 12112

Reports

Kilkenny, Francis; Shaw, Nancy; Gucker, Corey. 2014. Great Basin Native Plant Project: 2013 

progress report. Boise, ID: U. S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station. 211 p.

Individual reports:

Richardson, Bryce; Shaw, Nancy; Germino, Matthew J. Ecological genetics of big sagebrush 

(Artemisia tridentata): genetic structure and climate based seed zone mapping, p. 17-23.

Shock, Clinton C.; Fiebert, Erik B.; Saunders, Lamont D.; Shaw, Nancy; Sampangi, Ram K. 

Seed production of Great Basin native forbs, p. 108-154.

Ott, Jeff; Shaw, Nancy. Seeding Native Species following Wildfire in Wyoming Big 

Sagebrush (Artemisia tridentata ssp. wyomingensis).

Shock, Clinton C.; Feibert, Erik B. G.; Saunders, Lamont D.; Shaw, Nancy. 2014. Directsurface 

seeding systems for successful establishment of native wildflowers. In: Shock, Clinton C. 

Oregon State University, Malheur Experiment Station Annual Report 2013, Department of Crop 

and Soil Science Ext/CrS 149. p. 159-165.

Shock, Clinton C.; Feibert, Erik B. G.; Saunders, Lamont D.; Shaw, Nancy; Sampangi, Ram. 

2014. Irrigation requirements for native perennial wildflower seed production in a semi-arid

environment. In: Shock, Clinton C. Oregon State University, Malheur Experiment Station 

Annual Report 2013, Department of Crop and Soil Science Ext/CrS 149. p. 166-183.
189

Shock, Clinton C.; Feibert, Erik B. G.; Saunders, Lamont D.; Shaw, Nancy. 2014. Irrigation 

requirements for native wildflower seed production for perennial and annual species planted in 

the fall of 2009. In: Shock, Clinton C. Oregon State University, Malheur Experiment Station 

Annual Report 2013, Department of Crop and Soil Science Ext/CrS 149. p. 184-192.

Shock, Clinton C.; Feibert, Erik B. G.; Saunders, Lamont D.; Shaw, Nancy. 2014. Irrigation 

requirements of annual native wildflower species for seed production, fall 2012 planting. In: 

Shock, Clinton C. Oregon State University, Malheur Experiment Station Annual Report 2013, 

Department of Crop and Soil Science Ext/CrS 149. p. 193– 197.

Shock, Clinton C.; Feibert, Erik B. G.; Saunders, Lamont D.; Shaw, Nancy. 2014. Tolerance of 

native wildflower seedlings to pre-emergence and post-emergence herbicides. In: Shock, Clinton

C. Oregon State University, Malheur Experiment Station Annual Report 2013, Department of 

Crop and Soil Science Ext/CrS 149. p. 198-205.

Presentations

Ott, J. E.; Shaw, N. L.; Cox, R.; Pellant, M.; Roundy, B. A.; Eggett, D. 2013. Native plant 

seeding techniques for burned Wyoming big sagebrush communities: comparisons of drilling 

and broadcasting methods across multiple sites. Poster presented at Society for Ecological 

Restoration 5th World Conference; 2013 October 4-12; Madison, WI. Abstract.

Ott, J.; Shaw, N. 2014. Seeding native species following fire in Wyoming big sagebrush

(Artemisia tridentata subsp. wyomingensis). Great Basin Native Plant Annual Meeting; 2014 

March 17-18; Boise, ID.

Ott, J. E.; Shaw, N. L.; Cox, R. D.; Pellant, M. 2014. Effects of drilling and native plantseeding 

on vegetation in the northern Great Basin. Society for Ecological Restoration Northwest and 

Great Basin Chapter Conference; 2014 October 6-10; Redmond, OR.

Ott, J.; N. Shaw; R. Cox; M. Pellant; B. Roundy; D. Eggett. 2014. Comparison of native plant 

seeding techniques on burned ARTRW sites. 67th Annual Meeting of the Society for Range 

Management; 2014 February 9-14; Orlando, FL.

Hild, A.; Taylor, M.; Shaw, N. 2014. Exotic plant species and soil characteristics of 

rehabilitation seedings following wildfire in northern Utah. Society for Ecological Restoration 

Northwest and Great Basin Chapters Conference; 2014 October 6-10; Redmond, OR. (presented 

by Shaw).

Newingham, Beth A.; Ott, Jeffrey E.; Ganguli, Amy C.; Shaw, Nancy L. 2013. Collaborative 

efforts comparing the effects of two post-fire seed drills on ecosystem responses in the Great 

Basin. Poster presented at Great Basin Consortium Conference; 2013 December 9-10; Reno, NV. 

Abstract.
190

Newingham, B. A.; A. C. Ganguli; J. E. Ott; and N. L. Shaw. 2014. Linking soil erosion and 

plant recovery under different post-fire seeding techniques in Wyoming big sagebrush 

communities. Society for Ecological Restoration Northwest and Great Basin Chapter 

Conference; 2014 October 6-10; Redmond, OR.

Shaw, N. 2014. Sagebrush restoration and native plant for the Great Basin. USDI BLM Natural 

Resources Staff, September 2014, Washington, DC.

Additional Products

Steering Committee Member, National Seed Strategy (N. Shaw).

Conference Co-chair, Society for Ecological Restoration Northwest and Great Basin Chapters 

Joint Annual Conference (N. Shaw).

References

Centre for Forest Conservation Genetics. 2014. Climate of Western North America. [Online] 

Available: http://cfcg. forestry. ubc. ca/projects/climate-data/climatebcwna/#ClimateWNA 

[Accessed 4 June 2014].

Herrick, J. E. , J. W. Van Zee, K. M Havstad, L. M. Burkett, and W. G. Whitford. 2005. 

Monitoring manual for grassland, shrubland and savanna ecosystems, Volume I. Las Cruces, 

NM, USA: US Department of Agriculture, Agricultural Research Service, Jornada Experimental 

Range. 36 p.

R Core Team. 2012. R: A language and environment for statistical computing. R Foundation for 

Statistical Computing, Vienna, Austria. [Online] Available: http://www. R-project. org.

Soil Survey Staff, Natural Resources Conservation Service, United States Department of 

Agriculture. 2015. Web Soil Survey. [Online] Available: http://websoilsurvey. nrcs. usda. gov. 

[Accessed 15 January 2015].

Wirth, T. A. and D. A. Pyke. 2007. Monitoring post-fire vegetation rehabilitation projects – a 

common approach for non-forested ecosystems. U. S. Geological Survey Scientific Investigation 

Report 2006-5048, 36p.
191

Project Title A New Biological Control Strategy for Cheatgrass and

Ventenata dubia Based on Fungal Endophytes

Project Agreement No. 10-CR-11221632-182

Principal Investigators and Contact Information

George Newcombe, Professor 

University of Idaho

Department of Forest, Rangeland, and Fire Sciences 

PO Box 441133

Moscow, ID 83844-1133

208.885.5289

georgen@uidaho.edu

David L. Griffith, Ph. D. Candidate 

University of Idaho

Environmental Science & Water Resources Program 

875 Perimeter Dr.

Moscow, ID 83844-1142

208.310.0180

griffith@uidaho.edu

Sara Ashiglar, M.S. Candidate 

University of Idaho

Department of Forest, Rangeland, and Fire Sciences 

PO Box 441133

Moscow, ID 83844-1133

208.413.0056

sashiglar@gmail.com

Maryam Alomran, Ph.D. Candidate 

University of Idaho

Department of Plant, Soil and Entomological Sciences 

PO Box 441133

Moscow, ID 83844-1133

509.263.7173

ms.a1@hotmail.com

Project Description

Fusarium Greenhouse Bioassays

Endophytic fungi identified were isolated from surface sterilized ventenata and cheatgrass plants 

collected from Latah County, ID. Twenty Fusarium spp. isolates (10 isolated from ventenata and 

10 from cheatgrass) were chosen for a series of greenhouse experiments. Fusarium spp. were of 

particular interest because members of this genus are often pathogenic on local grasses and 

cereal grain crops. Blocked, randomized seed trays were planted with three grasses: bluebunch

wheatgrass, cheatgrass and ventenata. Each seed was inoculated with spores and mycelia from 
192

the cheatgrass- or ventenata-derived isolates. The ten Fusarium spp. inoculum derived from 

ventenata and the ten Fusarium spp. inoculum derived from cheatgrass were first bulked

together as treatments on the three grass species seeds. A second set of experiments inoculated 

the three grass species with spores/mycelium from the individual Fusarium isolates. All 20

Fusarium isolates were sequenced in the ITS region and identified through comparison in the

GenBank database.

Results from the bulked inoculum treatments showed that both ventenata-derived and 

cheatgrass-derived Fusaria treatments significantly decreased germination and aboveground 

biomass of bluebunch wheatgrass compared to the control. Both ventenata-derived and 

cheatgrass-derived Fusaria treatments had no impact on cheatgrass germination, but did 

significantly decrease its aboveground biomass. The ventenata-derived Fusaria had a 

significantly negative effect on ventenata germination (unlike the cheatgrass-derived Fusaria 

which was not significantly different), while both ventenata-derived and cheatgrass-derived 

Fusaria had positive effects on aboveground biomass. Results from the single-isolate 

inoculations varied somewhat, though generally followed the same patterns. These experiments 

additionally showed a significant decrease in cheatgrass germination caused by several 

ventenata- and cheatgrass- derived Fusaria treatments, as well as a significant increase in 

ventenata germination by a few of the ventenata-derived Fusaria treatments and decrease in 

germination by a few cheatgrass-derived Fusaria treatments.

Lab experiments using assays to determine whether various ventenata and cheatgrass derived 

Fusaria would infest and kill seedlings was conducted. A highly pathogenic strain of Fusarium 

culmorum was obtained from WSU Plant Pathology to further test ventenata’s susceptibility to 

these relatively common pathogens. This is partly inspired by the ability of some nonpathogenic, endophytic Fusarium spp. to protect against disease caused by Fusarium pathogens 

in other systems. Although F. culmorum did not infest ventenata seeds or seedlings on water 

agar plates in the laboratory, it did cause disease (and death) of nearly all ventenata plants in a 

greenhouse experiment where ventenata and winter wheat were planted in the same pots and 

both inoculated with F. culmorum spore suspension. Data indicating whether the ventenataderived Fusarium species protect ventenata from infection or disease onset by F. culmorum is 

still being analyzed.

This work is currently being prepared as part of a Ph.D. dissertation and for submission to 

peer-reviewed journals by David Griffith.

Ventenata Fungal Communities via Metagenomics

While limited isolations of fungi as described in the experiments above have been done, virtually 

nothing is known about the fungal symbionts and pathogens of ventenata. In the Palouse region 

of north-central Idaho and east-central Washington, ventenata appears to be a state of enemy 

release from pathogens according to field surveys done in the spring and summer of 2014, even 

when adjacent to other grasses showing active signs and symptoms of disease. Preliminary work 

has shown that ventenata germination is negatively affected in laboratory conditions by wheat 

pathogen Fusarium culmorum, as well as other Fusaria (see above). It is not known whether 

these pathogens are commonly present and affect ventenata in the field. Nor is there any other

known information on other fungal symbionts and how they may be driving (or limiting) the 

invasion of ventenata.
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Samples of ventenata plants were collected from 22 sites across Latah, Nez Perce, Lewis and 

Clearwater counties in Idaho, Gallatin County in Montana, and Grant County in Oregon in the 

spring and summer of 2014. Surrounding ecotypes and crops were noted along with latitude and 

longitude. Elevation ranged approximately 1,000 to 4,900 feet. The samples were surface 

sterilized and freeze-dried for storage. DNA extracted from plants, and associated endophytic 

fungal material, will be sequenced using a MiSeq Illumina processor. This method will allow 

broad scale environmental sampling of the fungal and potentially bacterial metagenomics using 

millions of short reads that can be compared to known species in the GenBank database. It will 

offer a snapshot of ventenata fungal communities across a geographic and elevational range.

DNA are currently be extracted and PCR conditions optimized. Analysis of sequence data is 

projected for spring 2015. This work is supporting an M.S. project by Sara Ashiglar.

Management Applications

The work on fungal endophytes will provide baseline knowledge on how cheatgrass and 

ventenata are successfully invading native perennial grasslands and out-competing native 

species. The metagenomics work describing fungal communities of ventenata will be compared 

with future work looking for potential biocontrol agents on ventenata fungal communities from 

its native range.

Three scientists (Newcombe, Ashiglar, and Griffith) attended the MtnSEON ventenata-focused 

working group (“The Processes, Patterns and Mechanisms of Ventenata dubia Invasion in 

Complex Western Landscapes”) meeting in Moscow, ID. At this meeting, attention was brought 

to key researchers of the potential problems of ventenata infestations in areas where it is not yet 

well established. Collaborations between scientists across the West ensued to help understand the 

gaps in knowledge and inconsistencies in monitoring protocols about this species. The social and 

economic implications of the spread of ventenata across the west is more clearly understood and 

will be brought up in future meetings with relevant stakeholders. Known management protocols 

for control, and areas where such protocols need to be developed, were discussed and 

disseminated.

One scientist (Griffith) attended the MtnSEON Blue Mountain Working Group meeting held in 

LaGrande, OR on January 27, 2015. This meeting focused on stakeholder and management input 

into ongoing and future research goals for the Starkey Experimental Forest and Range. Detection 

and control of ventenata, and the impact of cheatgrass, medusahead, and other annual grass 

invaders in our region was discussed. Griffith will be attending another meeting in Portland, OR, 

at the invitation of USFS, of the Blue Mountain Working Group focused on designing research 

goals, where he will present knowledge about cheatgrass and ventenata microbial ecology 

derived from the funded project.
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Presentations

Griffith, David L.; Newcombe, George; Ashiglar, Sara. 2014. Endophytic fungi linked to 

invasion of Ventenata dubia. Great Basin Native Plant Project annual meeting; 2014 March 17-

18; Boise, ID. Poster.

Griffith, David L.; Newcombe, George; Ashiglar, Sara. 2015. The Mountain Social Ecological 

Observatory Network (MtnSEON), Blue Mountain Working Group meeting; 2015 January 30-

31; Moscow, ID. Workshop participation and research discussion.

Ashiglar, Sara. 2015. Ventenata and Fusarium. 2015. The Mountain Social Ecological 

Observatory Network (MtnSEON), Blue Mountain Working Group meeting; 2015 January 30-

31; Moscow, ID. Presentation.

Publications and Additional Products

A manuscript, titled “Revisiting the life cycle of Sordaria fimicola” was submitted to PLOS ONE

and has currently been resubmitted after first reviews were received and revisions made.

A manuscript is being prepared for submission to The Journal of Ecology on field experiments 

demonstrating the effects on biomass and fecundity of inoculating cheatgrass with Sordaria 

fimicola.

A manuscript is being prepared for submission on laboratory and greenhouse experiments with 

ventenata and cheatgrass-derived Fusaria, and their effects on the growth and competitive 

abilities of cheatgrass, ventenata, and bluebunch wheatgrass.

Research conducted under this agreement will inform a third chapter of Griffith’s dissertation, on 

Social Ecological Systems frameworks for research and management focused on plant invasion 

in the American West. This chapter will be revised for submission in summer of 2015.