Conservation and Restoration of Sagebrush 

Ecosystems and Sage-Grouse:

An Assessment of USDA Forest Service Science

Deborah M. Finch, Douglas A. Boyce, Jr., Jeanne C. Chambers, Chris J. Colt, 

R. Kasten Dumroese, Stanley G. Kitchen, Clinton McCarthy, Susan E. Meyer, 

Bryce A. Richardson, Mary M. Rowland, Mark A. Rumble, Michael K. Schwartz, 

Monica S. Tomosy, and Michael J. Wisdom

United States Department of Agriculture

Forest Rocky Mountain General Technical Report

Service Research Station RMRS-GTR-348 March 2016
Finch, Deborah M.; Boyce, Douglas A., Jr.; Chambers, Jeanne C.; Colt, Chris J.; Dumroese, 

R. Kasten; Kitchen, Stanley G.; McCarthy, Clinton; Meyer, Susan E.; Richardson, Bryce A.; 

Rowland, Mary M.; Rumble, Mark A.; Schwartz, Michael K.; Tomosy, Monica S.; Wisdom, 

Michael J. 2016. Conservation and restoration of sagebrush ecosystems and sage-grouse: 

An assessment of USDA Forest Service Science. Gen. Tech. Rep. RMRS-GTR-348. Fort 

Collins, CO; U.S. Department of Agriculture, Forest Service, Rocky Mountain Research 

Station, 54 p.

Abstract

Sagebrush ecosystems are among the largest and most threatened ecosystems in North 

America. Greater sage-grouse has served as the bellwether for species conservation in 

these ecosystems and has been considered for listing under the Endangered Species Act 

eight times. In September 2015, the decision was made not to list greater sage-grouse, but 

to reevaluate its status in 5 years. Concerns over sage-grouse and associated habitats have 

set in motion sweeping Federal and State land management plan changes and proactive science-based conservation actions to address threats within the realm of management control.

For nearly a century, the Forest Service (FS), U.S. Department of Agriculture (USDA), has 

studied sagebrush ecosystems and for decades has focused on sage-grouse biology and 

habitat requirements. Our team of FS scientists and managers prepared this assessment 

to summarize FS strengths, capabilities, partners, past and current research, and potential 

future high-priority research areas for conservation and restoration of sagebrush ecosystems 

and sage-grouse. We identified research and science-based management needs of the 

National Forest System where lands are important for breeding and brood-rearing habitats for 

sage-grouse. We recommend expanded research and science delivery by FS scientists. This 

work will help meet continuing widespread concerns and calls for science-based conservation 

to mitigate threats to sagebrush ecosystems, conserve populations of sage-grouse and other 

sagebrush-obligate species, and restore sagebrush ecosystems throughout the western 

United States.

Keywords: sagebrush, sage-grouse, disturbance, fire, invasive species, restoration

Cover photos

Top left: A high elevation sagebrush hillside on the Uinta-Wasatch-Cache National Forest, 

Utah, after snowmelt in mid to late July. Photo by Teresa Prendusi, USDA Forest Service.

Top right: Female greater sage-grouse observed at a high-elevation mountain big sagebrush 

site, Inyo National Forest, California. Photo by Chris Balzotti, Stanford University, used 

with permission.

Bottom left: RMRS technicians monitoring a post-fire native grass, forb and sagebrush 

seeding in northern Utah. Photo by Matt Fisk, USDA Forest Service.

Bottom right: Wyoming big sagebrush, Owyhee County (southwest) Idaho. Photo by Matt 

Fisk, USDA Forest Service.
i

Author Information (all are current or former USDA Forest 

Service employees)

Deborah M. Finch is a Supervisory Biologist and Program Manager with the Rocky Mountain 

Research Station (RMRS) in Albuquerque, New Mexico.

Douglas A. Boyce, Jr., is a National Wildlife Ecologist with the National Forest System in 

Washington, DC.

Jeanne C. Chambers is a Research Ecologist with RMRS in Reno, Nevada.

Chris J. Colt is a Wildlife Biologist with the Caribou-Targhee National Forest in Pocatello, 

Idaho.

R. Kasten Dumroese is a Research Plant Physiologist with RMRS in Moscow, Idaho.

Stanley G. Kitchen is a Research Botanist with RMRS in Provo, Utah.

Clinton McCarthy is a retired Regional Wildlife Ecologist with the Intermountain Region in 

Ogden, Utah.

Susan E. Meyer is a Research Ecologist with RMRS in Provo, Utah.

Bryce A. Richardson is a Research Geneticist with RMRS in Provo, Utah.

Mary M. Rowland is a Research Wildlife Biologist with the Pacific Northwest Research 

Station (PNW) in La Grande, Oregon.

Mark A. Rumble is a Research Wildlife Biologist, Retired, with RMRS in Rapid City, South 

Dakota.

Michael K. Schwartz is a Research Wildlife Biologist with RMRS in Missoula, Montana.

Monica S. Tomosy is a National Wildlife Program Leader with Research and Development in 

Washington, DC.

Michael J. Wisdom is a Research Wildlife Biologist with PNW in La Grande, OR.

iii

Executive Summary

Sagebrush ecosystems are among the largest and most threatened ecosystems in North 

America. Greater sage-grouse (GRSG) has served as the bellwether for species conservation in these ecosystems and has been considered for listing under the Endangered Species 

Act (ESA) eight times. Most recently, in 2010, GRSG was warranted for listing under the 

ESA, but precluded by higher priority listing actions. In September 2015, the decision was 

made not to list GRSG, but to reevaluate the listing decision again in 5 years. Concerns 

over sage-grouse and associated habitats have set in motion sweeping Federal and State 

land management plan changes and proactive science-based conservation actions to address 

threats within the realm of management control. Nevertheless, persistent ecosystem threats, 

such as wildfire and invasive annual grasses, will continue to affect sagebrush ecosystems 

in the western portion of the GRSG range, and land development and land use activities will 

continue to affect sagebrush ecosystems in the eastern portion of the GRSG range. Sciencedriven management activities played a large role in preventing designation of GRSG as an 

endangered species in 2015. Ongoing management will be required to continue to mitigate 

threats to sagebrush ecosystems and to restore habitat for GRSG and other sagebrushobligate species. 

USDA Forest Service (FS) scientists have studied sagebrush ecosystems for nearly a 

century, and have focused on sage-grouse biology and habitat requirements for decades. This 

science has been included in many conservation and management plans produced by State 

and Federal agencies. Given FS science capacity, germane publications, research facilities 

and strategic locations, history of sagebrush science, and the importance of National Forest 

System (NFS) lands for breeding and brood-rearing habitats for sage-grouse, we recommend expanded research and science delivery by FS scientists. This work will help address 

continuing widespread concerns and calls for science-based conservation to mitigate threats 

to sagebrush ecosystems, conserve populations of sage-grouse and other sagebrush-obligate 

species, and restore sagebrush ecosystems throughout the western United States. 

Our team of FS scientists and managers prepared this assessment to summarize the agency’s strengths, capabilities, partners, past and current research, and potential future priority 

research areas for conservation and restoration of sagebrush ecosystems and sage-grouse. 

We identified four areas of strength, leadership, and knowledge development:

•  Evaluating links among sage-grouse population ecology, monitoring, and habitat 

•  Understanding disturbances and stressors in sagebrush ecosystems 

•  Analyzing and designing landscapes to improve habitat connectivity

•  Developing methods, models, and plant materials to restore sagebrush habitats. 

Our assessment identifies high-priority research and science-based management needs 

of the NFS and contributes to preparation of the USDA Forest Service Sage-Grouse 

Conservation Science Strategy (Finch et al. 2015).

v

Contents

Executive Summary ........................................... iii

1.0—Overview.................................................1

2.0—Forest Service Research and Development Strengths 

and Leadership: A Review of Knowledge ....................5

2.1—Sage-Grouse Population Ecology, Monitoring, and Habitat Links .5

2.1.1—Modeling greater sage-grouse and its habitat ............5

2.1.2—Assessment and monitoring of sage-grouse habitat .......6

2.1.3—Quantification of core and edge habitats across 

sage-grouse range....................................6

2.1.4—Linking sage-grouse populations and demography 

to habitat change .....................................7

2.2—Disturbances and Stressors in Sagebrush Ecosystems.........8

2.2.1—Invasion of annual grasses into sagebrush ecosystems ...10

2.2.2—Expansion of pinyon and juniper into sagebrush 

ecosystems ........................................10

2.2.3—Altered fire regimes and post-fire recovery .............10

2.2.4—Sage-grouse response to disturbance impacts on habitat. .12

2.2.5—Assessments of sage-grouse vulnerability to stressors ....13

2.3—Landscape Analysis and Design to Improve Habitat 

Connectivity .........................................14

2.3.1—Landscape connectivity and corridors .................14

2.3.2—Assessment of sage-grouse gene flow and connectivity ...15

2.4—Methods, Models, and Plant Materials to Restore Sagebrush 

Habitats ............................................16

2.4.1—Sagebrush genetic and taxonomic complexity...........16

2.4.2—Adaptive genetic variation and seed transfer guidelines ...17

2.4.3—Native plant development—seed and seedling 

biology and plant propagation ..........................19

2.4.4—Restoration methods—practices and equipment for 

weed control and seeding .............................21

2.4.5—Responding to threats: Loss of habitat through 

invasive plants, fire, and other stressors and disturbances ....21

2.4.6—Monitoring effectiveness of restoration treatments .......23

2.4.7—Conceptual models—linking habitat restoration to 

sage-grouse recovery ................................24

3.0—Research Needs..........................................26

4.0—Science and Communication to Aid Management 

Decisions on National Forest System Lands ................29

4.1—Questionnaire to Determine Knowledge Needs and Gaps ......30

4.2—Results of sage-grouse/sagebrush questionnaire.............30
vi

4.3—Identifying new areas of Forest Service research and 

science delivery to meet management needs ...............31

4.4—Role of syntheses, decision support tools, assessments, and 

databases...........................................31

4.5—Enhance science delivery and communication—what more

can be done .........................................32

5.0—Facilities, Centers, and Projects Having Key Locations, 

Contacts, and Expertise .................................33

6.0—Summary................................................34

7.0—Acknowledgments ........................................35

8.0—References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35

Appendix: Results of Sage-grouse/Sagebrush Questionnaire ........53

Respondents .............................................53

Selected Key Findings by Subject Area.........................53
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 1

1.0—Overview

Greater sage-grouse (Centrocercus urophasianus (GRSG) (fig. 1) is a high-profile species facing many anthropogenic and persistent ecosystem threats; it has been considered for 

Federal regulatory protections under the Endangered Species Act (ESA 1973) eight times 

(U.S. Fish and Wildlife Service 2015). Sage-grouse and more than 350 other species rely on 

sagebrush (Artemisia spp.) ecosystems (Schroeder et al. 2004), which are among the largest 

and most imperiled systems in North America (Noss et al. 1995). State and Federal agencies, 

research institutions, and stakeholders across 11 States in the western United States have 

undertaken tremendous conservation and research efforts to reduce threats to sagebrush 

ecosystems and greater sage-grouse. In 2010, the U.S. Fish and Wildlife Service (FWS), 

Department of the Interior (DOI), determined that greater sage-grouse warranted protection 

under the ESA, but that action was precluded by higher priority listing actions. The FWS 

found that the bi-state population of GRSG bordering California and Nevada, considered a 

distinct population segment, did not warrant ESA protection as a threatened species (U.S. 

Fish and Wildlife Service 2015a). As part of a 2011 court settlement, FWS agreed to make 

an initial determination on whether to propose GRSG for listing by September 30, 2015. In 

the interim years, unprecedented conservation partnerships across the western United States 

have developed with the aim to significantly reduce threats to GRSG across 90 percent of its 

breeding habitat. Subsequently, after a thorough status review, FWS concluded that GRSG 

remains relatively abundant and well-distributed across its 70-million ha range and in the 

foreseeable future does not face the risk of extinction. Thus protection under the ESA was 

not warranted as of 2015, but the listing decision would be reevaluated in 5 years (U.S. Fish 

and Wildlife Service 2015b).

Questions remain, however, on whether these conservation partnerships and their GRSG 

conservation plans and requirements will be consistently upheld by agencies, remain intact 

Figure 1—A displaying male greater sage-grouse on a lek in Butte County, South Dakota (photo by 

Steve Fairbairn, U.S. Fish and Wildlife Service).
2 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

if challenged by environmental groups, and mitigate threats to the bird and its habitat 

(Snodgrass and Jensen 2015). Two primary factors shaping the discussion of whether 

protection of GRSG was warranted under the ESA were threats to habitats important to 

sage-grouse populations and the lack of regulatory mechanisms, primarily in land use plans, 

to ensure protection for GRSG and its habitats. In response, the DOI’s Bureau of Land 

Management (BLM) and the Department of Agriculture’s Forest Service (FS), revised land 

use plans to incorporate measures to conserve, enhance, restore, or develop GRSG habitat 

and sagebrush ecosystems by reducing, eliminating, or minimizing threats, including climate 

change, that may force development of suitable GRSG habitat outside of its current range.

Sagebrush ecosystems remain threatened by a variety of stressors, and managers have 

called for science solutions to develop best practices for restoring damaged ecosystems. In 

January 2015, DOI released a Secretarial Order (SO 3666) for a comprehensive sciencebased policy to prevent and suppress rangeland fire and to restore sagebrush landscapes 

impacted by fire. Actions to prevent fire and restore habitats were deemed essential for conserving sagebrush ecosystems and GRSG habitat, particularly in the Great Basin (the Great 

Basin includes most of Nevada, half of Utah, and sections of Idaho, Wyoming, Oregon, and 

California). Continued collaboration and activities among State and Federal partners to proactively identify and implement actions to manage stressors and restore damaged sagebrush 

ecosystems remain necessary.

Scientists are recommending a holistic, multiscale approach for managing sagebrush 

habitats and sagebrush-obligate species (fig. 2) (Chambers et al. 2014d; Herrick et al. 2912; 

Knick and Connelly 2011; Rowland et al. 2006). At landscape scales, sage-grouse may serve 

as an “umbrella species” (species for which conservation indirectly protects other species) 

for other sagebrush-obligate species because of its requirement for large, contiguous areas 

of sagebrush (Rowland et al. 2006). For example, moderate to strong positive associations 

occur between GRSG and three sagebrush-obligate passerines (Brewer’s sparrow [Spizella 

breweri], sage sparrow [Amphispiza belli], and sage thrasher [Oreoscoptes montanus]) 

whose populations have declined (Hanser and Knick 2011). This relationship indicates that 

Figure 2—Estimated proportion of habitat for species of conservation concern in the Great Basin that would 

benefit from management of greater sage-grouse habitat within the historical range of sage-grouse in the Great 

Basin. Sagebrush-obligate species are indicated by white bars (from Rowland et al. 2006).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 3

a holistic management approach that addresses large-scale persistent threats to sage-grouse 

habitat may be expected to benefit sagebrush ecosystems and most sagebrush-obligate 

species (e.g., Chambers et al. 2014c). Davies et al. (2011) conclude that “the sagebrush 

(Artemisia) ecosystem is a prime example of an area where many conservation objectives 

could be simultaneously achieved by developing a comprehensive ecosystem conservation 

plan.” Multiple conservation objectives can be simultaneously met in sagebrush ecosystems 

by integrating scientific knowledge on factors such as ecosystem resilience and resistance 

and species habitat requirements into comprehensive ecosystem conservation plans 

(Chambers et al. 2014c,d). However, specific conservation actions for individual species 

including sage-grouse must necessarily recognize landscape-scale heterogeneity and differences in individual species’ habitat requirements (Hanser and Knick 2011; Rowland et al. 

2006).

The Forest Service, with a rich history of research and collective expertise in studying 

sagebrush ecosystems, has contributed substantially to sagebrush conservation, threat mitigation, and restoration efforts. FS scientists’ capacity to explore novel areas of sage-grouse 

and sagebrush biome science is facilitated by research station facilities in strategic locations. 

In particular, FS scientists continue to contribute unique science, services, and management 

tools to partners needing information in the following research areas pertinent to sagebrush 

ecosystem restoration and sage-grouse conservation:

•  Application of landscape analyses to planning

•  Genomics for conserving plant and animal populations

•  Effects, prevention, and control of fire and invasive species

•  Restoration science and applications

•  Seed and plant materials development sciences

•  Application of resilience and resistance concepts

•  Climate change adaptation science and models

•  Effectiveness monitoring and adaptive management.

Since the mid-1980s, the FS Research & Development (R&D) Mission Area has addressed key science and management questions about sagebrush ecosystems, specifically the 

role of sagebrush genetics in restoration science, the impacts and control of pinyon–juniper 

(Pinus-Juniperus spp.) expansion, and effects of annual grass invasions, especially in the 

Great Basin. As threats to GRSG became a dominant consideration for managing sagebrush 

habitats, FS R&D began integrating findings from earlier research into assessments, models, 

and new research for understanding and managing sage-grouse populations and habitats. 

This new research focused on controlling cheatgrass (Bromus tectorum) invasions in sagebrush ecosystems, treating and managing fire and postfire landscapes, restoring ecosystems 

affected by multiple disturbances, and linking sage-grouse biology and habitat requirements 

to management of plant communities and landscapes. Research expanded after 2000 to 

evaluate and control the spread of other invasive species; examine stressors such as energy 

development, drought, and climate change; assess effectiveness of sagebrush restoration 

actions; assess habitat and bird vulnerabilities; and evaluate sage-grouse and sagebrush 

genetics for improving restoration strategies and landscape design.
4 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Yet concerns about the conservation of the sagebrush ecosystem and sage-grouse continue to mount, creating expectations for natural resource agencies to demonstrate an ability 

to effectively manage sagebrush habitat and conserve GRSG across 11 western States (up to 

35 national forests or grasslands in the National Forest System), as well as to conserve the 

threatened Gunnison sage-grouse (Centrocercus minimus). The latter species is a Federally 

listed species (U.S. Fish and Wildlife Service 2014) whose small population occurs in Utah 

and Colorado, including the GrandMesa/Uncompahgre/Gunnison National Forest. Although 

the focus of this assessment is on GRSG, we recognize that key points are most likely applicable to Gunnison sage-grouse.

To be successful, land managers largely depend on integrating relevant and applicable 

scientific information. FS R&D has demonstrated strong leadership in identifying threats 

to natural resources, developing methods and models to address those threats, and providing managers science-based tools and options. FS R&D is committed to ensuring that 

its relevant and applicable science findings are available and delivered into the hands of 

practitioners through partnerships, publications, conferences, webinars, and consultations. 

Where FS R&D has unique research capabilities, it is committed to effectively responding to requests for additional information and to the need for more research and testing of 

hypotheses.

We have four objectives with this assessment. First, we intend to promote understanding 

and recognition of FS R&D strengths, science leadership, and collaborative opportunities 

in key areas of sagebrush and sage-grouse science, identify research emphasis areas within 

FS R&D based on a review of existing knowledge and future needs, stimulate increased 

development and delivery of knowledge about sagebrush ecosystems and sage-grouse conservation, and serve as a communication tool. This assessment aims to improve awareness 

of the FS’s collective expertise, partnerships, and existing facility resources by summarizing 

the sagebrush ecosystem and sage-grouse conservation science we conduct. It is intended to 

identify, for the benefit of other agencies and stakeholders, the FS’s current research niches 

of expertise that can be expanded in the future.

Second, we intend for this assessment to serve as a planning guide for managers, staff, 

and scientists working on, or concerned about, sagebrush ecosystems and sage-grouse 

related issues. To this end, information from this assessment was integrated into the USDA 

Forest Service Sage-Grouse Conservation Science Strategy (Finch et al. 2015).

Third, we hope that this assessment, coordinated through scientists using a team approach, serves to foster synergistic collaborations among FS R&D scientists, their partners, 

and the management community. We invited sagebrush and sage-grouse scientists to share 

what they do across locations and disciplines and to succinctly synthesize their efforts into 

this single document.

Fourth, our intention is to support our colleagues in the National Forest System in achieving sagebrush ecosystem and sage-grouse conservation objectives. Much of our science is 

developed to aid management decisions on NFS and other Federal lands. For GRSG, NFS 

manages 8 percent of the GRSG range; most of the remainder is on BLM-managed lands. 

BLM and FS are the leads for developing Federal management action plans for GRSG. 

Importantly, the FS-managed GRSG habitat occurs mostly at higher elevation and may be 

disproportionately important to the future of GRSG because of the greater resilience of 

higher elevation habitat to invasive species and climate change.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 5

Our assessment presents areas of research emphasis where FS scientists have demonstrated significant leadership and strengths, as reflected by a review of FS R&D publications, 

science delivery, applications by stakeholders, and future research needs. Four research 

emphasis areas of FS R&D, identified based on existing knowledge and future needs and 

priorities, and outlined in the Sage-Grouse Conservation Science Strategy (Finch et al. 2015) 

are:

•  Evaluating links between sage-grouse population ecology, monitoring, and habitat

•  Understanding disturbances and stressors in sagebrush ecosystems

•  Analyzing and designing landscapes to improve habitat connectivity

•  Developing methods, models, and plant materials to restore sagebrush habitats.

This assessment is divided into five sections. The first section is an overview. The second section assesses existing knowledge developed by FS R&D scientists who exemplify 

strengths and science leadership, clarifies and emphasizes how these scientists are experts in 

these research emphasis areas, and frames the science within the context of each of the four 

research emphasis areas. The third section discusses research needs identified through the 

assessment process. The fourth section identifies NFS needs for future science and identifies 

science delivery methods to meet those needs. Finally, the fifth section reviews key facilities 

and locations having relevant science expertise and capacity.

2.0—Forest Service Research and Development 

Strengths and Leadership: A Review of Knowledge

2.1—Sage-grouse Population Ecology, Monitoring, and Habitat Links

Several comprehensive publications describe the basic ecology and habitat requirements 

of greater sage-grouse (Connelly et al. 2000; Schroeder et al. 1999). A landmark contribution 

was the book Greater Sage-grouse: Ecology and Conservation of a Landscape Species and 

its Habitat (Knick and Connelly 2011). Forest Service (FS) research contributed to specific 

chapters in this book (e.g., Wisdom et al. 2011) that deal with management and conservation 

status, basic sage-grouse ecology, sagebrush ecology, and factors associated with extirpation 

of GRSG. FS scientists have developed a plethora of sage-grouse models, many of which 

were subsequently used in broad-scale assessments of the species and its habitat. Other 

research focuses on links between sage-grouse demographics and habitat status or human 

disturbance. Below we describe primary lines of FS research on these topics.

2.1.1—Modeling greater sage-grouse and its habitat

Mechanistic models that clearly link animal distributions and performance to habitat 

variables are useful tools for species conservation and management. Forest Service scientists 

have developed a variety of models that address greater sage-grouse (GRSG) ecology and 

habitat requirements. As part of the Interior Columbia Basin Ecosystem Management Project 

(ICBEMP) in the late 1990s, the science strategy team developed more than 30 Bayesian 

belief network models for key species of conservation concern, including GRSG. These 

models combined empirical and hypothesized relations in probability-based projections 

of conditions. The models were subsequently used to estimate abundance and distribution 

of habitat and population outcomes under a suite of management alternatives across the 
6 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

58 million ha of the Great Basin (Marcot et al. 2001; Raphael et al. 2001). Wisdom et al. 

(2002a) validated two restoration models by using areas of extirpated versus occupied range. 

One restoration scenario assumed a 50-percent reduction in detrimental grazing effects and 

a sixfold increase in actively restored areas compared with future management proposed 

by the FS and the Bureau of Land Management (BLM). The second scenario assumed a 

100-percent reduction in detrimental grazing effects and the same increase in active restoration as the first scenario. In an ecoregional assessment in the Wyoming Basin, Hanser et al. 

(2011a) developed a statistical model relating GRSG occurrence and abundance to environmental covariates.

2.1.2—Assessment and monitoring of sage-grouse habitat

Connelly et al. (2000) offered guidelines to manage sage-grouse populations and habitats 

and later provided guidance on habitat monitoring (Connelly et al. 2003). More recently, the 

BLM and FS collaborated to develop a comprehensive monitoring framework with methods 

to monitor habitat and evaluate effectiveness of FS and BLM land management plans to 

conserve GRSG and its habitat (DOI, BLM and USDA Forest Service 2014). The technical guide of Rowland and Vojta (2013) describes standardized methods of wildlife habitat 

monitoring on NFS and other lands that are now widely adopted. The methods are included 

in State wildlife action plans and land management units that are revising forest plans under 

the 2012 FS planning rule (USDA Forest Service 2012). One chapter in this technical guide 

features example habitat monitoring programs for three species, including GRSG (Goldstein 

et al. 2013). It illustrates conceptual models at landscape and site scales for processes acting 

as stressors on GRSG and projected impacts on its habitat, and discusses specific monitoring 

objectives and indicators at each scale, including fragmentation and connectivity.

FS scientists have spearheaded large-scale GRSG habitat assessments. Through the 

ICBEMP, FS scientists evaluated GRSG habitat conditions across the interior Columbia 

basin under various management scenarios (Hemstrom et al. 2002; Raphael et al. 2001; 

Wisdom et al. 2000, 2002). In recognition of that work, the BLM requested that FS researchers lead a broad-scale habitat assessment for GRSG and other sagebrush-associated 

species in the Great Basin ecoregion; information was shared through a book (Wisdom et al. 

2005b) and other publications (Rowland and Wisdom 2009; Rowland et al. 2006, 2010b). 

Using methods developed in the Great Basin, Wisdom and Rowland subsequently led an 

ecoregional assessment of GRSG and sagebrush communities in the 350,000 km2 Wyoming 

Basins and contributed to several chapters in the resulting book (see Hanser et al. 2011b). 

These chapters cover selection of sagebrush-associated species of concern (Rowland et al. 

2011), a model of GRSG occupancy and abundance (Hanser et al. 2011a), and management 

considerations (Knick et al. 2011).

2.1.3—Quantification of core and edge habitats across sage-grouse range

Patterns of species occurrence can help in evaluating factors associated with population 

losses. Wisdom et al. (2011) evaluated environmental conditions in areas of former (extirpated) range and range still occupied by GRSG and Gunnison sage-grouse. They identified 

the environmental conditions most strongly associated with landscape-scale extirpations 

based on the strongest environmental differences between areas of extirpation and current 

occupation. A variety of landscape metrics related to concepts of “core” versus “edge” areas 

were considered in the evaluation. Mean size of core areas (defined as the mean size of 
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 7

contiguous sagebrush patches) were >12 times larger in occupied range (3,964 ha) than in 

extirpated range (332 ha). Mean size of core areas were included in the second-best model, 

which provided >80 percent accuracy in discriminating between occupied and extirpated 

range. The single-best factor in correctly classifying occupied versus extirpated range was 

the percent area of sagebrush. Occupied range had nearly twice as much sagebrush area 

(46 percent) as extirpated range (24 percent); these results served as an additional indicator 

suggesting that the size of core areas was substantially larger in occupied range, which further corroborated results by Aldridge et al. (2008). Finally, Wisdom et al. (2011) identified 

“strongholds” of occupied range―these had large core areas and minimal edge, and were far 

from areas of extirpated range. In contrast, areas associated with a high risk of extirpation 

had small core areas that were often isolated and disjunct. These results provide a strong 

management framework for landscape designs to maintain and restore desirable characteristics of suitable habitat across the historical range of both sage-grouse species.

2.1.4—Linking sage-grouse populations and demography to habitat change

Understanding the demographics of life-history stages and linking these to environmental 

factors can improve management to positively influence sage-grouse populations and conservation planning. FS R&D has extensive experience evaluating demographic parameters 

and environmental covariates at multiple scales throughout the range of GRSG.

Habitat alteration may affect GRSG populations’ site occupancy, size of leks (male 

courtship assemblage areas) (e.g., Hess and Beck 2012; Smith et al. 2005; Wisdom et al. 

2011), or survival during individual life-history stages, such as juvenile or brood-rearing. 

FS scientist Rumble collaborated with many partners, including State wildlife agencies (in 

North Dakota, South Dakota, and Wyoming) and two universities to link sage-grouse habitat 

and demographic parameters in the eastern portion of the sage-grouse range (e.g., fig. 3) 

(Fremgen et al. 2015; Herman-Brunson et al. 2009; Kaczor et al. 2011a,b; Schreiber 2014; 

Schreiber et al. 2015a,b; Swanson et al. 2013). Rumble and his colleagues have studied 

Figure 3—Vegetation sampling site in greater sage-grouse nesting habitat, Carbon County, Wyoming 

(photo by Brian Dickerson, USDA Forest Service).
8 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

movement patterns and habitat selection of telemetered GRSG (fig. 4) in relation to natural 

disturbances and plan to evaluate effects of wind energy development on GRGS populations 

(e.g., Hanser et al. 2014; Schreiber et al. 2015a,b).

By using the Bayesian belief network models developed for GRSG as part of the 

ICBEMP (see 2.1.1—Modeling greater sage-grouse and its habitat), population outcomes 

were projected across the range of the species within the Interior Columbia Basin; potential 

outcomes ranged across a gradient from well-distributed and abundant to scarce, patchy, and 

at risk of extirpation (Raphael et al. 2001). Managers then applied model results to guide 

selection of alternatives in the environmental impact statement for this basin.

2.2—Disturbances and Stressors in Sagebrush Ecosystems

Sage-grouse are “landscape” species (species that use and may influence large landscapes) and declines in populations can often be attributed to the loss, fragmentation, and 

degradation of sagebrush ecosystems (Connelly et al. 2011). Primary drivers of change in 

sagebrush ecosystems are the direct effects of human land use and development and their 

interactions with more complex factors, such as altered fire regimes, invasive species, expansion of trees to larger areas, drought stress, and livestock grazing (Connelly et al. 2011). 

Effects of these disturbances and stressors are manifest at temporal scales ranging from 

years to centuries and spatial scales ranging from sites to large landscapes (Arkle et al. 2014; 

Doherty et al. 2010). Superimposing rapid climate changes over disturbances makes assessment and management of shrubland ecosystems and their services increasingly difficult 

(Finch 2012). The influences of a changing climate potentially have profound implications 

Figure 4—Greater sage-grouse with solar-powered PTT-100 global positioning system transmitter in a 

study of movement patterns in Wyoming (photo by Brian Dickerson, USDA Forest Service).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 9

for the distribution of sagebrush ecosystems. Kitchen et al. (in preparation) show the 

historical environmental settings for the development and establishment of sagebrush communities, their trajectories since the Pleistocene, and the implications for current and future 

distribution of sagebrush communities. Still and Richardson (2015) and Balzotti et al. (in review) modeled climate envelopes that are useful in identifying areas where sagebrush habitat 

is most likely to change in a warming climate. Balzotti et al. (in review) also integrated the 

confounding effects of invasive species. This information is valuable for understanding 

implications of changes to sage-grouse habitat and for identifying where management effort 

can best conserve habitat connectively.

A primary strength of FS research is in developing the required understanding of 

disturbance effects on sagebrush ecosystems across scales necessary to develop effective 

management strategies. Key areas of FS science leadership in sagebrush ecosystems pertain 

to disturbances and stressors associated with invasive species, conifer expansion, and fire 

(fig. 5A,B,C), as described next.

A

B

C

Figure 5—(A) Landscape 

converted to B tectorum

in north-central Nevada 

(photo by Nolan Preece, independent artist, used with 

permission), (B) sagebrush 

fire in south-central Idaho 

(photo by Doug Shinneman, 

U.S. Geological Survey), 

and (C) pinyon and juniper expansion in central 

Nevada (photo by Jeanne 

Chambers, USDA Forest 

Service).
10 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

2.2.1—Invasion of annual grasses into sagebrush ecosystems

Invasion and expansion of annual grasses is resulting in an annual grass-driven fire 

cycle and progressive conversion of lower elevation, warmer, and drier sagebrush ecosystems to ecosystems dominated by annual grasses, especially cheatgrass (Kitchen 2014). 

Successful management strategies require a detailed understanding of attributes of sagebrush ecosystems that determine resistance to invasion. To gain that understanding, FS 

scientists examined effects of climatic factors (temperature and precipitation) and common 

disturbances (fire, grazing) on soil water and nutrients (Whittaker et al. 2008), and on 

establishment, reproduction, and persistence of the most widespread annual invader―

cheatgrass (Beckstead and Augsburger 2004; Beckstead et al. 2010, 2011; Blank et al. 2007; 

Chambers et al. 2009, 2014d; Jones et al. 2015a, 2015b; Roundy et al. 2007). To understand 

implications for its adaptation, spread, and control, FS scientists examined germination 

characteristics and genetic structure among cheatgrass populations (Merrill et al. 2012) and 

interactions of cheatgrass with native plant species (Goergen and Chambers 2009) and sagebrush plant communities (Jones 2014). This research was the basis for creating conceptual 

and statistical models of factors that influence resistance to annual grass invasion across 

the environmental gradients that characterize sagebrush ecosystems (e.g., fig. 6B) (Brooks 

and Chambers 2011; Chambers et al. 2014b). These models are used widely to develop 

management strategies (Miller et al. 2013, 2014b, 2015). More recent collaborative research 

has examined how climate change is likely to affect both expansion and contraction of the 

invasive annual brome grasses, cheatgrass and red brome (Bromus rubens L.) (Bradley et al. 

2015).

2.2.2—Expansion of pinyon and juniper into sagebrush ecosystems

Conifer expansion, particularly by species of pinyon (and juniper), is a primary threat to 

sagebrush communities on national forest lands because it typically results in a decrease in 

understory sagebrush, grasses, and forbs (fine fuels), an increase in woody fuels, and less 

frequent but more intense fires (Miller et al. 2013). FS researchers and collaborators examined pinyon juniper ecology (Chambers 2001; Chambers et al. 1999), and the changes that 

occur in soil resources (Rau et al. 2007, 2008) and plant community composition (Chambers 

2000, 2005), reductions in sagebrush cover in relation to pinyon-juniper expansion 

(Rowland et al. 2010a), and the consequences for fuel loads with progressive tree expansion 

and infilling (Miller et al. 2008, 2014b; Roundy et al. 2014a,b). They also examined resilience of these areas to wildfire and management treatments and resistance to invasive annual 

grasses across the environmental gradients that characterize these ecosystems (fig. 6A,B) 

(Chambers et al. 2014b,c). This research was the basis for management guides for prioritizing areas for treatment and selecting the best treatment options (Chambers et al. 2014d; 

Miller et al. 2014a, 2015; Pyke et al. 2014, 2015).

2.2.3—Altered fire regimes and post-fire recovery

Fire is a dominant disturbance process regulating composition and structure of sagebrush 

ecosystems. Understanding variability in natural and altered disturbance regimes and their 

effects on vegetation is critical for developing effective management and restoration strategies. FS scientists and collaborators characterized historical (multi-century) fire regimes for 

big sagebrush (Artemisia tridentata) ecosystems, and evaluated the changes that are occurring in these fire regimes (Heyerdahl et al. 2006; Kitchen and McArthur 2007; Miller and 
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 11

Heyerdahl 2008; Miller et al. 2008). Working across a diverse range of big sagebrush sites, 

they assessed the drivers of, and variability in, sagebrush population recovery (fig. 7) and 

tree occupation (Goodrich et al. 2008; Nelson et al. 2014), and quantified the effects of postfire sagebrush seed production on recovery rate (Landeen 2015). On landscapes prone to tree 

invasion, sagebrush dominance is facilitated by fire-free intervals long enough for sagebrush 

recovery but short enough to prevent tree invasion and dominance (Miller and Rose 1999; 

Miller et al. 2013).

Figure 6—(A) Resilience to disturbance and (B) resistance to cheatgrass (B. tectorum) over a 

typical soil temperature and moisture gradient in sagebrush ecosystems of the western United 

States (adapted from Chambers et al. 2014b).
12 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Landscape-scale vegetation dynamics across a range of fire frequencies and climate 

scenarios are being modeled in order to assess the resilience of big sagebrush ecosystems 

at various spatial and temporal scales. Fire frequency is expected to increase with increasing air temperature and drought. If fire frequency is too great, it may stimulate a phase 

change from sagebrush-dominated vegetation to steppe and will likely become unsuitable 

for sage-grouse. Even without fire in the system, climate change is likely to reduce current 

distribution of sagebrush (Friggens et al. 2012; Richardson et al. 2012a). Habitat for sagegrouse may shift to higher latitudes and elevations under warming climates (Finch et al. 

2012) and may be needed at higher elevations (e.g., FS lands), where habitat will be less at 

risk of fire and invasion by annual invasive grasses.

2.2.4—Sage-grouse response to disturbance impacts on habitat

Forest Service researchers have extensive experience in evaluating effects of landscape 

change and land management actions on wildlife species, including sage-grouse, through 

direct evaluations of movement patterns, and demographics and resource selection models 

applied to evaluate various types of disturbance. This expertise is particularly relevant to the 

evaluation of land disturbance from invasive species, fires, or anthropogenic development 

(e.g., energy sources, roads, transmission lines), and stems from a long history of research 

on relationships between wildlife populations and vegetation patterns. A key example is 

the FS-led evaluation of factors associated with extirpation of sage-grouse by Wisdom et 

al. (2011), which identified a wide variety of anthropogenic and vegetation factors strongly 

associated with extirpation (see 2.1.3—Quantification of core and edge habitats across 

Figure 7—Vegetative recovery 5 years after a fire in a mountain big sagebrush community. Scattered 

sagebrush plants grew from seeds that survived the fire and are now large enough to begin producing 

the seeds that will give rise to a second postfire generation. Plant density is sufficient for full sagebrush 

recovery in 25–35 years after the fire (photo by Stanley G. Kitchen, USDA Forest Service).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 13

sage-grouse range). Results are being used for conservation planning across the range of 

the species on public and private lands. Another example of FS expertise deals with wind 

energy, projected to compose about 20 percent of the U.S. energy demand by 2030 (U.S. 

Department of Energy 2008). High-quality wind resources overlap the range of sage-grouse, 

particularly in the eastern portion of the range (Becker et al. 2009). FS scientists have expertise to evaluate effects of wind energy development on populations and habitat, and research 

on these effects on GRSG habitat use and movements is underway in Wyoming (Mark 

Rumble, personal communication, USDA Forest Service, Rapid City, SD, 2015; Schreiber 

et al. 2015a,b). In addition, Rowland (2004) identified and synthesized effects of a large 

suite of management activities on GRSG habitats and populations across the species range, 

including prescribed fire, livestock grazing, and land use change.

Degradation of sagebrush habitat during a drought year was evaluated by using 

MODIS (moderate resolution imaging spectroradiometer) imagery “ForWarn”: 

http://forwarn.forestthreats.org/. Field observations noted a significant loss of leaves and 

branches from decadent old sagebrush at this south-central Oregon site, which was experiencing a single year of severe drought (25 percent of average precipitation). A dynamic 

vegetation model (Drapek et al. 2015) will be used to show expected end-of-century sagebrush habitat distribution and sagebrush productivity levels in the region (Grulke et al., in 

preparation).

FS researchers also have a long history of evaluating gene flow of sensitive species and 

how gene flow is altered by changes in the landscape (see 2.3.2—Assessment of sage-grouse 

gene flow and connectivity). Of particular value is the use of spatially explicit genetic data 

to test hypotheses of how genes move across landscapes and how this movement will be 

affected by disturbances (Cushman et al. 2006; Manel et al. 2003; Schwartz et al. 2009; 

Shirk et al. 2015). Currently, FS researchers are analyzing thousands of feather samples from 

sage-grouse leks to evaluate gene flow and genetic continuity across the northern portion of 

the species’ range (see 2.3.2—Assessment of sage-grouse gene flow and connectivity). This 

research will also be used to assess how stressors such as land conversion, energy development, and fire may influence movement (e.g., Shirk et al. 2015).

2.2.5—Assessments of sage-grouse vulnerability to stressors

Recent FS efforts have focused on assessing the vulnerability of sage-grouse and their 

habitat to the effects of climate change and other natural and anthropogenic stressors in 

the Intermountain Region (Balzotti et al., in review). Contemporary and projected future 

conditions were evaluated based on past distribution patterns reconstructed from paleoclimatological and historical data. Specific objectives included: (1) describing pre-Euroamerican 

settlement sagebrush distribution and changes during the last several millennia; (2) developing models of climate and sagebrush ecological niches and assessing their implications for 

sage-grouse habitat occupancy and use; (3) identifying natural and anthropogenic causes for 

sagebrush degradation and loss, with emphasis on those associated with wildfire, invasive 

species, and conifer expansion on NFS lands; (4) identifying habitats at greatest risk for 

loss and disruption of sage-grouse population connectivity; and (5) providing management 

options, including identifying and prioritizing those areas where sagebrush restoration efforts could best conserve habitats and connectivity of sage-grouse habitats. Objectives 4 

and 5 were also the focus of an FS-led assessment of habitat threats in the Great Basin that 
14 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

provided broad-scale evaluation of the risks of sagebrush loss from pinyon-juniper expansion and cheatgrass invasion (Rowland et al. 2010b; Wisdom et al. 2005b,c).

2.3—Landscape Analysis and Design to Improve Habitat Connectivity

FS R&D has been instrumental in studying changes in sage-grouse and the landscapes 

they inhabit. Fragmentation of the species’ preferred habitat and effects of this fragmentation 

on sage-grouse population cohesion and dispersal have made these studies of paramount 

importance. This research spans multiple scales and uses multiple approaches, from experiments to species distribution modeling to landscape-scale genetic approaches. Recent 

analyses also are addressing the influence of climate change on these systems and the implications for connectivity of sage-grouse habitats.

2.3.1—Landscape connectivity and corridors

A variety of natural and anthropogenic disturbances have fragmented sage-grouse 

habitats. These disturbances include removal of sagebrush through prescribed fire and 

mechanical means, natural fire, energy development and mining, exurban development, 

agricultural development, and road and transmission line corridors. These disturbances must 

also be viewed in the context of a changing climate.

FS scientists have evaluated broad-scale fragmentation in sagebrush ecosystems through 

several projects. As part of the ICBEMP (see 2.1.1—Modeling greater sage-grouse and 

its habitat), Hann et al. (2003) developed a spatially explicit disturbance departure and 

fragmentation index across the basin and found high levels of departure and fragmentation 

in low-elevation rangelands. Other ICBEMP analyses led by FS scientists projected future 

trajectories and effects of land management on the persistence of sage-grouse habitats and 

populations (Hemstrom et al. 2002; Wisdom et al. 2002a,b). Landscape configurations related to landscape connectivity, as measured by habitat size, arrangement, and fragmentation, 

were considered. In the Great Basin ecoregion, Wisdom et al. (2005) evaluated risk of cheatgrass encroachment for habitats of several species groups, including sagebrush-associated 

species such as GRSG. Their composite risk map can be used to depict habitat connectivity 

by highlighting watersheds in the Great Basin that have low cheatgrass invasion risk and 

relatively high habitat abundance. In an additional analysis, Rowland and Wisdom (2009) 

identified habitat networks in the Great Basin for sagebrush-dependent vertebrate species, 

including GRSG, based on sagebrush habitat area and configurations.

As discussed earlier (2.1.1—Modeling greater sage-grouse and its habitat), Wisdom 

et al. (2011) evaluated environmental conditions in areas of former (extirpated) range and 

range still occupied by sage-grouse. They considered a wide variety of landscape metrics, 

including multiple measures of habitat fragmentation, core size, patch size, edge area, roads, 

and transmission lines, all of which are part of, or directly influence, measures of landscape 

connectivity. Moreover, they identified habitat “strongholds” for sage-grouse persistence 

based on characteristics of landscape connectivity and isolation, which depicted de facto 

corridors. These FS-led evaluations have been widely used in conservation planning by 

public and private land managers across the range of sage-grouse.

Modeling potential occupied area of a species and the factors that affect its distribution 

has become a useful tool in ecological research and management. In particular, bioclimate 

models are becoming even more important, but, like any model, they require assumptions 
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 15

and appropriate interpretation (Pearson and Dawson 2003; Soberon and Nakamura 2009). 

Two primary issues with bioclimatic models are that they (1) rarely take into account 

intraspecific genetic variation (note that common garden genecological and reciprocal transplant studies, used in seed zone construction for example, can be used to understand how 

intraspecific genetic variation interacts with climate; see Kilkenny 2015) and (2) rarely take 

into account how biotic interactions might affect species distributions and possibly could 

be mistaken for climatic interactions. FS R&D has developed many of the analytical tools 

(Iverson et al. 2008; Rehfeldt et al. 2006) and data (FS, Forest Inventory and Analysis) used 

to create these models. For sagebrush, three such models have been developed that can be 

used in connectivity planning. Two models used land cover imagery to assess cheatgrass and 

disturbance threats (Bradley 2010) and evaluate the effects of ecohydrology on sagebrush 

occurrence (Schlaepfer et al. 2012). Recently, Still and Richardson (2015) used presence and 

absence data of Wyoming big sagebrush (A. tridentata subsp. wyomingensis) to develop a 

subspecies model.

Currently, FS-managed GRSG habitat tends to occur at higher elevations on more mesic 

sites than GRSG habitat on non-FS lands, as mentioned earlier. Under a changing climate, 

the total area of sagebrush communities and high-quality sage-grouse habitat may potentially shrink and move upslope (Friggens et al. 2012), increasing the importance of these 

FS-managed habitats. To plan for this possibility, Balzotti et al. (in review) modeled and 

analyzed climate patterns at regional and sub-regional scales and evaluated potential changes in sagebrush and pinyon-juniper communities on national forests in Utah and Nevada. 

When overlaid with the distribution of sage-grouse habitats, the model allows managers to 

evaluate areas where conservation may have the greatest benefit for population retention and 

recovery.

2.3.2—Assessment of sage-grouse gene flow and connectivity

As noted earlier (2.2.4—Sage-grouse response to disturbance impacts on habitat), gene 

flow of sensitive animal species, including gene flow altered by changes in the landscape, 

has long been a focus of Forest Service research. Using spatially explicit genetic data to test 

various hypotheses is particularly valuable, especially when considering disturbances to 

the ecosystem (Manel et al. 2003; Schwartz et al. 2009). This discipline is called landscape 

genetics and was developed and advanced by FS researchers (Cushman et al. 2006; Manel 

et al. 2003; Schwartz 2001). It is used in ongoing FS research that is exploring how disturbances, including exurban expansion, agricultural conversion, sagebrush fragmentation, 

energy development, and invasive species, affect gene flow across multiple scales (Braun 

1998; Copeland et al. 2009; Knick et al. 2003; Naugle 2011; Naugle et al. 2004, 2006, 2011; 

Shirk et al. 2015).

Using genetic data collected from feathers and blood, and observations of bird presence 

in about 70 different breeding areas in eastern Washington, FS scientists in collaboration 

with other research institutions, developed a model for predicting how GRSG move across 

the landscape under different future disturbance scenarios (Shirk et al. 2015). Their findings 

showed that transmission lines are barriers that limit GRSG movement between patches of 

habitat, which could interrupt gene flow among subpopulations and reduce genetic diversity. 

At the smaller scale, research is exploring how potential disturbances influence movement 

among Core Breeding Areas in Montana (Cross et al., in review). A larger collaboration of 

scientists with the U.S. Geological Survey, Department of the Interior and the FS is asking a 
16 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

similar question across the entire distribution of GRSG (Cross et al., in review; Hanks et al., 

in review; Oyler-McCance et al., in preparation).

Cross et al. (in review) analyzed 16-locus microsatellite genotypes from 2,108 GRSG 

associated with 508 spatially distinct locations across Montana, North Dakota, and South 

Dakota. Subpopulation structure aligned closely with ecologically distinctive areas of the 

landscape created by historical biogeographic processes. This finding suggests that contemporary GRSG fine-scale genetic population structure is shaped by the underlying landscape 

physiography. The ability to match genetically distinct populations with suitable landscapes 

may be useful when designing corridors for connecting populations, or for reintroducing 

populations to vacated areas. Finally, new approaches applying graph theory are being used 

in conjunction with the FS’s multiscale sage-grouse genetics work to explore the impacts of 

degradation or loss of individual leks to the overall connectivity of the system.

2.4—Methods, Models, and Plant Materials to Restore Sagebrush 

Habitats

FS researchers and collaborators are instrumental in developing effective restoration 

approaches and treatments for sagebrush ecosystems. FS researchers are developing climatically adapted plant materials with high genetic diversity that confer resistance and resilience 

to a community, as well as seed transfer guidelines for planting the right seed in the right 

place. The establishment of partnerships such as the Great Basin Native Plant Project (see 

2.4.3—Native plant development—seed and seedling biology and plant propagation) has 

been instrumental in advancing seed science and plant material development for application 

in restoration of the sagebrush ecosystem. Additional strategies focus on active restoration 

methods, such as vegetation management to reduce fuels and remove trees, control of exotic 

annual grasses, and reestablishing native, perennial vegetation.

Timely restoration of degraded sagebrush plant communities at landscape scales is 

essential for maintaining sage-grouse habitats. An understanding of factors that regulate 

plant community resilience to disturbance and resistance to invasion by annual grasses is 

necessary for prioritizing restoration activities, selecting the most effective treatments, and 

choosing the most appropriate plant materials. Treatments that reduce woody fuel loads, 

increase herbaceous perennials, and decrease annual invasive abundance promote increases 

in resilience and resistance while restoring structure and function needed for sagebrushdependent wildlife (Brooks and Chambers 2011; Chambers et al. 2014b,c,d; Korfmacher 

et al. 2003; Tausch et al. 2009). Restoration using seed mixes that contain seed adapted to 

appropriate climate zones and conditions will promote resilience and resistance of sagebrush 

communities as well (Richardson et al. 2015). Although passive restoration measures, such 

as changes in land use, can often be effective in improving ecological condition, these measures alone are insufficient to meet the burgeoning restoration needs in the face of multiple 

threats. Among these threats are altered fire regimes including large high-severity fires, 

current and projected rates of spread and dominance by trees and exotic annuals, existing 

poor condition of herbaceous vegetation in many areas, and projected increased stress due to 

climate change.

2.4.1—Sagebrush genetic and taxonomic complexity

Restoration of sagebrush ecosystems is complicated by the taxonomic and genetic variability of sagebrush. FS common garden research on big sagebrush has shown that three 
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 17

widespread subspecies (tridentata, wyomingensis, and vaseyana) (fig. 8) are genetically distinct (Richardson et al. 2012b) and adapted to different habitats (McArthur and Welch 1982; 

McArthur et al. 1988; Wang et al. 1997). Correct subspecies identification and use of source 

plant materials (seed or outplanted seedlings) are critical for successful sagebrush restoration. Past efforts to identify and match seed sources to planting sites have been hampered by 

a lack of reliable tests of seed-lot identity. Of particular importance to sage-grouse habitat 

restoration is proper identification of Wyoming big sagebrush (subspecies wyomingensis), 

which is the predominant subspecies seeded following large wildland fires. A recent study 

demonstrated that seeds of Wyoming big sagebrush can be distinguished from those of basin 

big sagebrush (subspecies tridentata), based on weight; and a retrospective analysis of past 

BLM seed purchases (years 2012 and 2013) showed that Wyoming big sagebrush seed lots 

were largely composed of basin big sagebrush (Richardson et al. 2015). This research can 

be used to develop a certification step to distinguish seeds of these two subspecies of big 

sagebrush. It also suggests that restoration failure of burned sagebrush sites may be linked to 

use of seed mixes having seed of the wrong sagebrush subspecies.

2.4.2—Adaptive genetic variation and seed transfer guidelines

Climate is a principal factor affecting adaptation within a plant species. Knowledge of 

how plants are climatically adapted is fundamentally important to the process of seed collection and proper transfer to restoration sites. Seed collection should be focused on regions 

that are most likely to be threatened by disturbance so that restoration can proceed with seed 

that is appropriately adapted to the threatened region. The research approach to elucidate 

climatic adaptation is long established (Turesson 1925), and the large body of subsequent 

research primarily focused on tree species. During the last decade, FS has begun to apply 

this approach to native grasses, forbs, and shrubs (Bower et al. 2014; Johnson et al. 2013; 

Richardson et al. 2014; St. Clair et al. 2013), a number of which are important species in 

sage-grouse related plant communities. Genecological research has been used to determine 

areas where plant species are under threat from climate change and to identify promising 

seed sources for use in changed climates (Kilkenny 2015). Data from common garden 

Figure 8—Indication of the different growth rates among big sagebrush subspecies: tridentata (far right), 

wyomingensis (middle), and vaseyana (far left) (photo by Bryce Richardson, USDA Forest Service).
18 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

genecological and reciprocal transplant studies, used in seed zone construction, can be used 

to estimate the effects of intraspecific genetic variation on species distribution under climate 

change. Currently, FS researchers at the Rocky Mountain Research Station and the Pacific 

Northwest Research Station are performing a large-scale reciprocal transplant study (using 

16 common garden sites) on bluebunch wheatgrass (Pseudoroegneria spicata), a keystone 

structural species in many sagebrush habitats, to determine in unprecedented detail the climatic adaptation of 78 wild seed sources (Kilkenny et al. 2013).

The FS-managed Western Wildland Environmental Threat Assessment Center has a 

comprehensive seed zone website where seed zones are displayed (http://www.wwetac.

net/index_files/seedzone.shtml). Bower et al. (2014) developed generalized provisional 

seed zones that can be applied to any plant species in the United States to help guide seed 

movement in the face of climate change. Preliminary work for understanding the adaptive 

genetic variation in big sagebrush ecosystems is available (fig. 9; Richardson et al. 2013). 

Figure 9—Predicted landscape survivorship of Wyoming big sagebrush based on common garden 

mortality patterns (Richardson et al. 2013). These patterns are largely explained by climate variables 

describing cold winter and spring temperatures. This model is projected based on climate surface data 

and partitioned into groups from low to high survival (red = low survival, green = high survival). Survival 

is one of several traits that can be combined to develop seed transfer zones.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 19

In addition, FS research has addressed other genetic characteristics of sagebrush species 

(Meyer 1994; Miglia et al. 2007; Smith et al. 2002; Welch and McArthur 1981).

2.4.3—Native plant development—seed and seedling biology and plant 

propagation

Active restoration of sagebrush communities generally includes planting seeds and seedlings of native plants. Chambers and MacMahon (1994) developed a conceptual model of 

seed movement and fate that is helpful in understanding how likely a seed will successfully 

establish. Availability of adequately adapted seed sources of forbs, shrubs, and many grasses 

is limited (fig. 10), creating a bottleneck to large-scale restoration (Roundy et al. 1997; Shaw 

and Jensen 2014). A multi-state partnership initiated by FS and BLM in 2001, known formerly as the Great Basin Native Plant Increase and Selection Project (Shaw et al. 2012) and 

now as the Great Basin Native Plant Project (GBNPP), is helping to address this problem. 

More than 20 Federal, State, and private cooperators are involved in the GBNPP. Current 

research conducted by GBNPP and others focuses on developing seed zones where plant 

materials can be transferred with little risk of being poorly adapted to their new location, 

increasing seed availability, understanding native plant response to climate change, and 

evaluating native plant interactions with invasive species. The GBNPP hosts an active website (http://www.greatbasinnpp.org/) where its publications, presentations, and annual reports 

are posted.

In 2001, FS also created the Reforestation, Nurseries and Genetic Resources Program 

(RNGR) (http://rngr.net/), with the mission to supply people who grow seedlings with the 

latest technical information, and to provide links to other organizations and individuals with 

similar interests (Haase et al. 2011). Scientists and other professionals on the RNGR team 

Figure 10—Although many species of forbs are already in production, reliable methods are needed to 

economically produce a broader suite of forb species with appropriate genetics to meet restoration objectives (photo by Matt Fisk, USDA Forest Service).
20 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

contribute regularly to the literature on native plant propagation and nursery science (e.g., 

Dumroese et al. 2012). These contributions include papers on the influence of container size 

and storage on big sagebrush seedling growth, survival, morphology, and cold hardiness 

(Herriman et al. 2009; Overton et al. 2013), as well as broader reviews about forbs, pollinators, and GRSG (Dumroese et al. 2015, in press), assisted plant migration (Williams and 

Dumroese 2013), and tribal nursery plants (Dumroese et al. 2009). All of these publications 

have value for understanding ideas and approaches for restoring sagebrush ecosystems.

Knowledge of life-history traits is necessary to add new species to the restoration menu 

(Kitchen 1994; Shaw and Jensen 2014). FS scientists and collaborators have developed 

conceptual approaches for seed movement and fate in any ecosystem (Chambers and 

MacMahon 1994) and have investigated seed dormancy and germination regulation (Bonner 

and Karrfalt 2008; Meyer 1994; Meyer and Kitchen 1994a,b; Meyer and Monsen 1991; 

Meyer et al. 1995; Scholten et al. 2009), seedling emergence and growth (Chambers 2002; 

Chambers and Linnerooth 2001; Kitchen 1995; Monsen and Kitchen 1994; Shock et al. 

2012, 2015), pollination strategies (Cane 2008), and fecundity and plant longevity (Kitchen 

1995; Shock et al. 2015; St. Clair et al. 2013) for numerous species native to sagebrush ecosystems. Specialized equipment and cultural practices are being developed by FS scientists 

and collaborators to improve seed yields for species grown in agronomic settings (Shock et 

al. 2014, 2015) and managed wildland stands (Armstrong 2007; Ott et al., in preparation) 

(fig. 11). FS scientists have developed standardized seed testing protocols and storage practices needed to stabilize markets and ensure seed quality for many native species (Karrfalt 

and Shaw 2013; Kitchen 2001).

Figure 11—A rangeland drill in use at the burned site of the Scooby Fire, in northern Utah, in 2008. This 

drill was modified for seeding diverse mixtures of native plant species. It is an example of the specialized seeding techniques under development for restoring sagebrush ecosystems following wildfire in the 

Great Basin (photo by Beth Newingham, USDA Agricultural Research Service).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 21

Novel equipment for collecting seeds has been described (Jensen 2004) and guidelines 

are available for proper collection and certification of seed from natural stands (Adair et 

al. 2006; Currans et al. 1997; Young et al. 1995). FS researchers and their collaborators 

continue to develop cultural practices needed to increase the availability and quality of seed 

needed for sagebrush ecosystem restoration (Shaw et al. 2012).

2.4.4—Restoration methods—practices and equipment for weed control and 

seeding

Active restoration of arid and semiarid sagebrush ecosystems that lack the necessary 

perennial plants for unassisted recovery is particularly challenging because of low, and 

highly variable, precipitation, episodic plant establishment, and widespread invasion by 

exotic weeds. FS researchers and their collaborators have a productive history of developing 

the necessary plant establishment methods and seeding equipment for restoring sagebrush 

and its communities. They have produced hundreds of publications on the subject (Kiehl 

et al. 2014; Kitchen and Monsen 1995; Meyer and Warren 2015; Monsen and Meyer 

1990; Monsen et al. 1992, 2004a,b). Research by the FS as part of the Shrubland Research 

Consortium has kept researchers and practitioners up-to-date on current restoration topics 

and approaches through biennial symposia and publication of 16 symposium proceedings 

(1983–2011). The GBNPP (see 2.4.3—Native plant development—seed and seedling biology and plant propagation) generates research products, annual conferences, and webinars 

focused on improving the availability of native plant materials and on providing the knowledge and technology required for their use in restoring diverse native plant communities 

across the Great Basin. Research on restoration methods is broad and includes topics such 

as testing the use of drilling to establish native species after fire (Thompson et al. 2006), 

determining the effectiveness of different types of drills for establishing native mixes 

containing seeds of different sizes (Ott et al., in preparation; Taylor et al. 2014), evaluating 

species compatibility and invasion resistance within restored communities (Allen and Meyer 

2014; Parkinson et al. 2013), developing methods and testing seed mixes for restoring key 

plants (e.g., forbs) required by GRSG (Dumroese et al., in press), and testing use of carbon 

addition and repeated burning to decrease soil nitrogen availability and invasive annual grass 

(cheatgrass) establishment and reproduction (Jones et al. 2015b; Mazzola et al. 2008, 2011).

Outplanting of seedlings to reestablish difficult-to-seed species such as sagebrush is increasing (McAdoo et al. 2013) and is supported by FS research that examines (1) native plant 

nursery cultural practices including seedling storage, (2) the consequences of those practices 

on seedling physiology and outplanting success, and (3) the biophysical limitations of outplanting sites (Davis et al., in press; Overton et al. 2013; Pinto et al. 2012, 2015) (fig. 12).

2.4.5—Responding to threats: Loss of habitat through invasive plants, fire, 

and other stressors and disturbances

FS researchers and collaborators developed and tested many of the vegetation management treatments currently used to reduce woody fuels and increase native perennial 

vegetation in sagebrush ecosystems ( Monsen et al. 2004b). The FS’s research uses syntheses and regional studies to develop basic information and conceptual models on resilience 

and resistance of sagebrush systems to better target management actions (Board et al. 2011; 

Brooks and Chambers 2011; Chambers et al. 2007, 2014b,c,d; D’Antonio and Chambers 

2006; McIver et al. 2010; Miller et al. 2013; Monsen and Kitchen 1994). The Sagebrush 
22 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Treatment Evaluation Project (SageSTEP; http://www.sagestep.org/) provides regional 

information on effects of prescribed fire, mowing, and herbicide treatments in sagebrush 

ecosystems exhibiting cheatgrass invasion (Pyke et al. 2014; Rau et al. 2014) and prescribed 

fire, cut-and-leave, and mastication (shredding) treatments on those exhibiting pinyon or juniper expansion (Miller et al. 2014a; Roundy et al. 2014a,b). An interdisciplinary framework 

is used to couple environmental indicators of resilience and resistance with sage-grouse 

habitat requirements to prioritize management actions on the landscape (Chambers et al. 

2014c). Field guides have been developed that use resilience and resistance concepts to 

select appropriate sites for treatment (fig. 13) and to determine the most effective treatments 

(fig. 14) (Miller et al. 2014a, 2015; Pyke et al. 2015).

Recent work by Kitchen et al. (in preparation) portrays the influences of climate change 

in the distribution and abundance of sagebrush communities during the last several millennia, and the implications of climate change on projected distributions. This is intended to 

provide a valuable backdrop to the environmental envelope of sagebrush communities, and 

offers insight to areas where restoration may be most beneficial. Richardson et al. (2013) 

and Balzotti et al. (in review) developed climate envelopes for sagebrush, pinyon-juniper 

and invasive species, and modeled potential changes in the communities as a result of 

climate change. When coupled with anthropogenic landscape changes, this work suggests 

current and future conservation and restoration opportunities relative to these stressors.

For more than two decades, Meyer and her colleagues have evaluated biocontrol 

approaches for controlling cheatgrass infestations, including examinations of the head 

smut Ustilago bullata (e.g., Meyer et al. 2000, 2001, 2010a), the fungal seed pathogen 

Figure 12—Sagebrush being grown as bareroot stock at the USFS Lucky Peak Nursery near Boise, 

Idaho. Shrub seedlings like these could be outplanted as part of islands to act as microsite engineers 

to foster establishment of forbs required to support a variety of wildlife species (photo by R. Kasten 

Dumroese, USDA Forest Service).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 23

Pyrenophora semeniperda (e.g., Beckstead et al. 2010, 2011; Masi et al. 2014; Meyer et 

al. 2008, 2010b, 2013, 2015; Soliai et al. 2014), and underlying reasons for cheatgrass dieoffs (Baughman et al. 2016; Meyer et al. 2014). Currently the most promising biocontrol 

organism Meyer’s team has evaluated is a fungal seed pathogen that can kill dormant 

cheatgrass seeds and sometimes a high proportion of germinable seeds as well. This pathogen (Pyrenophora semeniperda) has been dubbed “black fingers of death” because of the 

fingerlike, black fruiting bodies that protrude from killed seeds (fig. 15). Their research has 

included extensive work on the population and evolutionary genetics of both cheatgrass and 

its pathogens, as well as biocontrol technology research aimed directly at developing effective field application methods. In some field inoculation treatments with the black fingers 

of death pathogen, complete control of the dormant carryover seed bank has been achieved. 

The goal of a practical, safe, and cost-effective commercial product for cheatgrass biocontrol 

on rangelands is now within reach. This biocontrol product addresses the problem of ungerminated seeds that carry over across years and hamper establishment success in restoration 

seedings even after successful control of germinated cheatgrass seeds or established stands.

2.4.6—Monitoring effectiveness of restoration treatments

Understanding the effectiveness of restoration treatments is the basis for adjusting methods to increase restoration success. Along with collaborators, FS researchers have evaluated 

restoration treatment effectiveness across the environmental gradients and ecological types 

Figure 14—In 2015, the Forest Service published 

A field guide for rapid assessment of post-wildfire 

recovery potential in sagebrush and piñon-juniper 

ecosystems in the Great Basin: Evaluating resilience to disturbance and resistance to invasive 

annual grasses and predicting vegetation response.

Gen. Tech. Rep. RMRS-GTR-338. Fort Collins, CO: 

USDA Forest Service, Rocky Mountain Research 

Station.

Figure 13—In 2014, the Forest Service published 

A field guide to selecting the most appropriate treatments in sagebrush and pinyon-juniper ecosystems 

in the Great Basin: Evaluating resilience to disturbance and resistance to invasive annual grasses 

and predicting vegetation response. Gen. Tech. Rep. 

RMRS-GTR-322. Fort Collins, CO: USDA Forest 

Service, Rocky Mountain Research Station.
24 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

that characterize sagebrush ecosystems. Evaluations include (1) effectiveness of broadcast 

versus drill seeding or other seeding techniques, (2) success in establishing big sagebrush, 

(3) tradeoffs between seeding aggressive introduced species and native species recovery, and 

(4) effects on invasive plant species (Knutson et al. 2014; Ott et al. 2003, in preparation). FS 

researchers also have participated in modeling the predicted probability of sage-grouse occupancy on treated sites based on sagebrush species and establishment, perennial grass cover, 

and invasive annual cover (Arkle et al. 2014).

2.4.7—Conceptual models—linking habitat restoration to sage-grouse 

recovery

FS scientists led the development and implementation of a variety of groundbreaking 

conceptual models for sage-grouse habitat restoration and recovery as part of ICBEMP, and 

later in ecoregional assessments in the Great Basin and across the entire range of GRSG 

(see 2.1.1—Modeling greater sage-grouse and its habitat). Conceptual models developed by 

FS scientists for sage-grouse restoration and recovery for application across the 58-million 

ha ICBEMP area include: (1) novel state-and-transitions models for sagebrush to evaluate 

future sagebrush habitat trajectories (Hemstrom et al. 2002); (2) innovative Bayesian belief 

network models developed and applied by Wisdom et al. (2002a) for evaluating current and 

future sage-grouse habitats; (3) development of a new habitat network approach for sagebrush-dependent species, including sage-grouse, and demonstration of this conceptual model 

Figure 15—Fruiting bodies of the seed pathogen Pyrenophora semeniperda, which illustrate why the pathogen was given its nickname “black fingers of death” (photo by Susan 

Meyer, USDA Forest Service).
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 25

in restoration planning (Wisdom et al. 2002b); and (4) additional integration and synthesis of 

sage-grouse models for restoration planning (Wisdom et al. 2005a).

In the Great Basin, FS researchers evaluated the use of (1) umbrella species concepts 

for managing sagebrush-dependent species (Rowland et al. 2005, 2006), and (2) restoration modeling using ecosystem resilience concepts for sagebrush habitats occupied by 

sage-grouse (Wisdom and Chambers 2009). Rowland and Wisdom (2009) also developed 

and applied new habitat network approaches for sagebrush-dependent vertebrate species 

in the Great Basin, including sage-grouse. FS-led conceptual models for sagebrush habitat 

restoration and sage-grouse recovery across the historical range of the species include those 

by Wisdom et al. (2005a), which introduced novel modeling paradigms and management 

approaches for prioritizing landscape conservation and restoration.

Most recently, an interdisciplinary working group of the Western Association of Fish and 

Wildlife Agencies (WAFWA) developed a strategic, multidisciplinary approach to reduce 

impacts of invasive annual grasses and altered fire regimes on sagebrush ecosystems and 

sage-grouse (Chambers et al. 2014d). The approach uses information about factors that 

influence sagebrush ecosystem resilience to disturbance and invasive annual grasses (Brooks 

and Chambers 2011; Chambers et al. 2014b,c; Wisdom and Chambers 2009) and distribution, relative abundance, and persistence of sage-grouse populations (Aldridge et al. 2007, 

2008; Doherty et al. 2010; Knick et al. 2013; Wisdom et al. 2011) to develop management 

strategies at landscape and site scales (fig. 16; Chambers et al. 2014d). A sage-grouse habitat 

matrix links relative resilience and resistance of sagebrush ecosystems with sage-grouse 

habitat requirements for landscape cover of sagebrush to help decisionmakers assess risks 

and determine appropriate management strategies at landscape scales. Focal areas for 

management are assessed by overlaying matrix components with sage-grouse Priority Areas 

for Conservation (PACs), breeding bird densities, and specific habitat threats. Decision 

tools are discussed for determining the suitability of focal areas for treatment and the most 

Figure 16—Landscape indicators of resilience and resistance, species habitat, and persistent habitat threats 

for sagebrush ecosystems and sage-grouse in the western portion of the species’ range. Soil temperature 

and moisture regimes are used to indicate landscape resistance and resilience, and sagebrush landscape 

cover is used to indicate sage-grouse habitat abundance. Wildfire is a persistent threat. See Chambers et al. 

(2014d) for a detailed explanation.

Resilience and Resistance Species Habitat Persistent Threat
26 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

appropriate management treatments. Management actions considered include fire operation 

(preparedness, prevention, and suppression), fuels management, postfire rehabilitation, and 

restoration/recovery.

Understanding the role of climate change is an important factor in prioritizing restoration 

opportunities (Richardson 2015; Still and Richardson 2015). Balzotti and colleagues (in 

review) have evaluated and modeled the implications of climate change on regional and subregional scales and the potential shifts in sagebrush and pinyon-juniper communities. They 

provide recommendations for identifying areas where conservation is most needed to retain 

connectivity of sage-grouse habitats.

These many and varied conceptual models developed and applied by FS scientists have 

been widely used in restoration planning for sage-grouse habitat at both landscape and site 

scales across the species range, and demonstrate a historical area of strength in FS research 

on sage-grouse and sagebrush ecosystems. This holistic approach has broad applicability for 

the conservation of threatened ecosystems and species across the western United States.

3.0—Research Needs

This section describes research needs associated with the four high-priority areas identified in the previous section. These research needs were identified based on stakeholder 

interests in relation to what areas of research Forest Service, Research and Development (FS 

R&D) scientists can begin to address given their current expertise, cooperators, and existing facilities and locations. Additional capacity such as increased numbers of scientists and 

resources (e.g., funds, technical support) is needed to address many of these needs.

1. Evaluate Links Between Sage-Grouse Population Ecology, Monitoring, and Habitat

•  Assess and monitor the contribution of NFS lands to sage-grouse population persistence, especially in the context of climate change

•  Evaluate the effects of livestock grazing on vital rates of sage-grouse populations

•  Determine the contributions of life-history stages (e.g., nesting versus brood-rearing) 

to sage-grouse population growth

•  Evaluate the impacts of disease and predation on sage-grouse survival, nesting success, 

habitat use, and population persistence

•  Find cost-effective surrogates to help monitor sage-grouse populations

•  Determine how establishment of core areas, as identified by Farm Bill (Conservation 

Reserve Program) policy and State efforts, and management of them over time, affects 

sage-grouse population growth rates and nesting success

•  Assess the impacts of restoration efforts (e.g., juniper and pinyon reduction or invasive 

species removal on sage-grouse growth rates and survival

•  Evaluate sage-grouse habitat use and population movement responses to different 

restoration treatments of burned or degraded sagebrush sites

•  Determine effects of transmission lines and other linear infrastructure, such as livestock 

fences, on habitat, dispersal (gene flow), and population dynamics of sage-grouse

•  Evaluate effects of agriculture and suburban developments, within and adjacent to current sage-grouse range, on population persistence

2. Understand Disturbances and Stressors in Sagebrush Ecosystems
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 27

•  Identify factors that influence resistance of sagebrush ecosystems to medusahead grass 

(Taeniatherum caput-medusae), North African grass (Ventenata dubia), and other 

recent annual invaders

•  Identify cover, species composition, and characteristics of perennial grasses and forbs 

required for sagebrush ecosystems to recover after disturbance and to resist invasive 

annual grasses

•  Determine potential effects of a warming, drying climate on spread and contraction 

of invasive annual grasses, pinyon and juniper species, sagebrush species, sagebrush 

communities, and sage-grouse

•  Assess effects of multiple fires on persistence and recovery of big sagebrush and its 

sage-grouse populations in the context of resilience and resistance

•  Assess impacts of past fire on the size, severity, and spatial patterns of future wildland 

fire in sagebrush ecosystems

•  Evaluate effects of livestock grazing on postfire sagebrush ecosystem recovery and 

rates of invasive annual grass and tree expansion

•  Evaluate effects of livestock grazing on habitat occupancy, use, and movement by 

sage-grouse

•  Assess effects of tree expansion on shrub species (e.g., bitterbrush, Purshia tridentata) 

known to co-dominate with sagebrush

•  Assess effects of energy development (e.g., wind, oil, gas) and associated infrastructure

(e.g., roads, pipelines) on sage-grouse habitats and populations and interpopulation 

movements

•  Study effects of communication towers and associated roads on sage-grouse land use

patterns

•  Project the effects of climate change on vegetation composition and fuels characteristics and how these changes will affect future fire regimes of sagebrush habitats.

3. Analyze and Design Landscapes to Improve Habitat Connectivity

•  Evaluate the use of foundational species and core landscape elements as habitat indicators for sage-grouse and sagebrush-obligate species

•  Evaluate additional biophysical indicators of ecosystem resilience and resistance that 

can be used to prioritize management activities and select appropriate management 

actions

•  Determine functional genetic variants that are identified to be under selection and assess how these vary in the different sage-grouse Distinct Population Segments (DPSs; 

i.e., smallest taxonomic unit protected under the ESA) and among different core areas

•  Develop seed zones and seed transfer guidelines for grass, shrub, and forb species to 

facilitate restoration of spatial links between sagebrush communities and sage-grouse 

populations

•  Evaluate and test restoration approaches and seeding and outplanting practices for improving habitat connectivity between sagebrush habitats and sage-grouse populations
28 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

•  Understand the relative contributions of environmental, demographic, and genetic 

effects on the decline of sage-grouse at the periphery of their range where populations 

are declining

•  Assess how fire management and other land management and restoration activities 

influence the movement of sage-grouse among core breeding areas

•  Evaluate potential changes in sage-grouse habitat connectivity and fragmentation under 

a variety of climate change scenarios

•  Conduct a spatial analysis of contributions by landowner to core sage-grouse habitats 

in order to better understand management limitations and opportunities

•  Conduct large-scale analyses of overlaps in sage-grouse populations with those of other 

sagebrush-dependent wildlife to better understand GRSG’s role as an umbrella species

•  Evaluate the individual and cumulative effects of all human activities and land uses on 

resistance and resilience of sagebrush habitats at ecoregional scales.

4. Develop Methods, Models, and Plant Materials to Restore Sagebrush Habitats

•  Develop suitable plant materials, with attention to native grasses, shrubs, and forbs, for 

restoring sagebrush communities and sage-grouse habitats in sites affected by fires, 

invasive species, and other stressors and disturbances

•  Develop seed zones and seed transfer guidelines for grass, shrub, and forb species that 

are important components of sagebrush ecosystems

•  Identify the climate envelopes for sagebrush, pinyon-juniper, and annual invasive communities and model their current and potential future distributions

•  Intersect sage-grouse habitats with those areas that will potentially be most affected by 

climate change to identify the areas where conservation and restoration will have the

greatest benefits

•  Develop protocols for incorporating climate change analyses into landscape 

assessments

•  Create high-resolution models of resilience to disturbance and resistance to invasive 

annual grasses that include climate, soils, and vegetation

•  Determine successful restoration approaches for recovering sage-grouse habitat and 

encouraging colonization by sage-grouse

•  Design and test restoration approaches and alternatives for warmer and drier sagebrush 

ecosystems with low resilience and resistance

•  Design and test restoration approaches and alternatives for pinyon and juniper communities that pose the greatest threats to sage-grouse habitats

•  Develop knowledge of the effects of livestock grazing (timing, amount, and duration) 

on restoration outcomes

•  Develop additional information on treatment effectiveness based on regional data and 

repeated monitoring over time

•  Develop information on treatment effectiveness based on new analytical approaches 

such as meta-data analyses

•  Understand climatic adaptation and develop seed transfer zones in big and dwarf sagebrushes and other key restoration species of their communities
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 29

•  Develop knowledge about and determine appropriateness of establishing novel 

sagebrush habitat in areas where newly emerging climate envelopes may allow it to 

develop.

4.0—Science and Communication to Aid Management 

Decisions on National Forest System Lands

Forests and grasslands in the National Forest System (NFS) across the western United 

States support key habitats for sage-grouse, especially higher elevation brood-rearing habitats in mountain big sagebrush communities (fig. 17). The Forest Service (FS) and Bureau 

of Land Management have worked to evaluate sage-grouse and sagebrush conservation 

measures in their land use plans. Due to recently adopted conservation measures, the U.S. 

Fish and Wildlife Service determined in 2015 that neither the greater sage-grouse nor the 

bi-state Distinct Population Segment (DPS) warranted listing under the Endangered Species 

Act, but Gunnison sage-grouse does have protection as a threatened species. Therefore, it 

is important to understand the kinds of knowledge and tools that FS land managers need to 

address sage-grouse and sagebrush conservation. This information is necessary to assess the 

effectiveness of conservation measures for sage-grouse, as well as to ensure that adaptive 

management and monitoring provide accurate information for future management. It is 

equally important to identify the knowledge gaps to improve or establish conservation measures that are informed by the best available science. In addition to sage-grouse, numerous 

species depend on sagebrush communities and are of conservation concern (fig. 2) (Rich et 

al. 2005; Rowland et al. 2006). Healthy sagebrush plant communities that support wildlife 

Figure 17—Female greater sage-grouse observed at a high-elevation mountain big sagebrush site 

(about 3050 m), Inyo National Forest, California (Photo by Chris Balzotti, Stanford University, used 

with permission).
30 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

such as the GRSG also support healthy pollinator communities, particularly because both 

GRSG and pollinators thrive on a high biodiversity of native forbs (see Dumroese et al., in 

press). GRSG populations require large landscapes and specific habitat conditions at broad 

scales to meet their seasonal life requirements. Rowland et al. (2006) and Hanser and Knick 

(2011) showed that GRSG serves as an umbrella species at landscape scales because of its 

widespread distribution and the broad range of sagebrush habitats it uses. Thus, a holistic 

management approach that addresses large-scale persistent threats to sage-grouse habitat 

may be expected to benefit sagebrush ecosystems and most sagebrush-obligate species 

(fig. 2). However, specific conservation actions for individual species or groups of species 

must recognize landscape-scale heterogeneity and differences.

4.1—Questionnaire to Determine Knowledge Needs and Gaps

In May 2013, a working group tasked with addressing FS issues on NFS-administered 

lands initiated an internal questionnaire related to sage-grouse and sagebrush management. 

The purpose was to identify and prioritize the kinds and scopes of science-based information 

and tools needed by FS land managers to address sage-grouse and sagebrush conservation. 

The questionnaire went to FS employees in the Washington Office, several regions (1, 2, 4, 5, 

and 6), three research stations (Pacific Northwest, Rocky Mountain, and Pacific Southwest), 

and all national forests that might potentially manage sage-grouse and their habitats. The 

working group targeted an array of resource management disciplines and employees working 

at all levels within the organization.

The questionnaire asked participants to respond to 37 questions in the following categories: (1) ecosystem dynamics of sagebrush communities on NFS lands, (2) contribution of 

sagebrush communities on NFS lands in supporting broader species/system conservation, 

(3) sage-grouse populations, (4) habitat management and restoration, and (5) conservation 

guidance for sage-grouse. Questionnaire results will inform those involved in developing an 

environmental impact statement (EIS), help in decisionmaking regarding compliance with the 

National Forest Management Act (NFMA), and support sage-grouse and sagebrush management. See Appendix for more details about the questionnaire.

4.2—Results of Sage-Grouse/Sagebrush Questionnaire

Respondents agreed that understanding sagebrush community dynamics and ecological 

relationships, especially for high-elevation mountain big sagebrush, was important, but 

disagreed about how much is known about effects of management on sagebrush restoration 

and whether sagebrush will be able to reestablish naturally as climate changes. They agreed 

that more attention should be given to managing sagebrush plant and animal communities on 

NFS lands and that knowledge was needed on effects of disturbance processes on sagebrush 

landscape patterns. Respondents indicated that more knowledge was needed about the

contributions of NFS lands and habitats to sage-grouse population health and persistence. 

Respondents strongly agreed that research and monitoring were needed to assess the effects 

of disturbances, including those from land management, and the effectiveness of habitat 

restoration on sage-grouse and sagebrush ecosystems. Syntheses of research findings that 

provided guidance to managers were viewed positively. Opinions differed about the sufficiency of information to address threats and risks to sage-grouse habitats and whether GRSG 

is an appropriate surrogate for monitoring other species.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 31

4.3—Identifying New Areas of Forest Service Research and Science 

Delivery to Meet Management Needs

Results of the questionnaire demonstrate that many new avenues of research can be 

initiated to meet the needs identified by NFS managers. Several of these needs corroborate 

those identified in section 3.0, “Research Needs.” Additional needs and gaps in information 

identified by questionnaire respondents can be addressed by FS R&D and partners through 

collaborative projects with NFS employees, and via consultations, syntheses, tool development, assessments, and science delivery and communication mechanisms.

4.4—Role of Syntheses, Decision Support Tools, Assessments, and 

Databases

FS R&D scientists are well-known for developing syntheses of existing literature in order 

to support management needs, and many examples are listed in the references section of this 

assessment. These often take the form of FS General Technical Reports and other agency 

serial publications, but they can also be published by journals and in monograph series 

of professional societies. Syntheses are usually collaborative efforts involving numerous 

scientists and experts within the FS as well as from other institutions, such as universities, 

other State and Federal natural resources agencies, and non-governmental organizations (e.g., 

Miller et al. 2013). A synthesis on a topic such as sage-grouse habitat restoration or landscape ecology is often requested by, or aimed at, land managers.

The Fire Effects Information System (FEIS; http://www.feis-crs.org/beta/) is an online 

collection of reviews of scientific literature on fire ecology and fire regimes. FEIS provides 

scientific information for resource management, restoration, rehabilitation, and fire management through three products: Species Reviews, Fire Studies, and Fire Regime Syntheses. 

Species Reviews are syntheses of literature that describe relationships of plants and animals 

with fire. Fire Studies provide detailed results from research and management projects on 

vegetation, fire characteristics, and fire effects. Fire Regime Syntheses describe historical 

and contemporary changes in fire regimes in particular ecosystems. Species Reviews are 

currently available for GRSG, Gunnison sage-grouse, sagebrush taxa, cheatgrass, and many 

other species important to sagebrush habitat (e.g., McWilliams 2002). FEIS is continually 

being updated to include new information and to respond to requests from land managers 

(see 4.5, Enhance Science Delivery and Communication—What More Can be Done).

Decision support tools and assessments can take various forms to help meet needs for 

management of sage-grouse habitats and landscapes. Tools can include field guides and 

publications that identify models, methods, geographic information systems (GIS) maps, 

and approaches to address questions about sage-grouse biology, disturbances, landscapes, 

and restoration (e.g., Miller et al. 2014a, 2015; Tausch et al. 2009). Tools can also be 

instruments, devices, models, methods, maps, databases, and approaches that are available 

directly to practitioners who access them online, by video, or through consultations, lectures, 

presentations, and training sessions. Tools are typically developed to meet a management or 

science application need and are designed so that new techniques do not have to be developed 

for routine management projects. Routine tools are also often used to test hypotheses where 

the question is not about the tool type but rather about how new findings can be revealed 

by using the tool. Typical tools are those that solve problems, provide outcomes, generate 

maps, enable consistency in inventory and monitoring, count populations, and model future 
32 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

projected changes in land cover, climate, and disturbances, such as likelihood of fire or invasions of non-native species. The results of assessments are often considered to be tools.

An assessment is an evaluation or estimation of the nature, quality, quantity, or response 

of organisms, habitats, landscapes, communities, stressors, and disturbances at any given 

time or place or during a period of time or across space. Assessments associated with 

sage-grouse may focus on populations in an area as small as a forest stand or as large as its 

entire range, the amount of occupied habitat, the availability of habitat with potential for occupancy, or the effects of changes on habitat or sage-grouse populations from stressors such 

as fire, invasive species, grazing, or climate change.

4.5—Enhance science Delivery and Communication—What More Can 

be Done

Modern communication tools have enhanced the delivery of science enormously during 

the last 20 years. In addition to traditional methods for delivering science findings, such as 

meeting presentations, posters, and lectures, more managers and scientists are using vehicles 

such as webinars, videoconferences, websites, social media, and interactive online databases 

to share information. For example, FEIS enables more than a half million people (the 

public and land managers) each year to search for Species Reviews, Fire Studies, and Fire 

Regime Syntheses (see 4.4—Role of Syntheses, Decision Support Tools, Assessments, and 

Databases) by species’ name, geographic location, Federal agency, plant community, life 

form, invasiveness, and nativity. Online newsletters have replaced many printed versions, 

journal publications often appear “online early” before they are printed with page numbers, 

and many journals are “online only.” FS partners with numerous State and Federal agencies, 

universities, and non-governmental organizations to design collaborative research and management projects and publish and deliver science findings about sagebrush and sage-grouse 

at regional, national, and international levels. FS was instrumental in establishing the Great 

Basin Research and Management Partnership, a group of 20 organizations that focus on 

developing science and technology transfer products applicable for managers. FS scientists 

and managers participate in and contribute to many other key partnerships that generate publications, syntheses, workshops and conferences, webinars, websites, and online newsletters 

such as:

•  SageSTEP, The Sagebrush Treatment Evaluation Project

•  Great Basin Consortium

•  Great Basin Fire Science Exchange

•  Range-wide Interagency Sage-Grouse Conservation Team, led by the Western 

Association of Fish and Wildlife Agencies

•  Landscape Conservation Cooperatives managed by agencies in the Department of the 

Interior

•  U.S. Geological Survey Climate Science Centers

•  Climate Hubs managed by USDA agencies

•  SO 3336 task and action groups responding to the SO3336 final report, An Integrated 

Rangeland Fire Management Strategy

•  Many other partnerships throughout the western United States.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 33

FS scientist Jeanne Chambers began the “Great Basin Fact Sheet” series about fire, 

invasive species, restoration, and other issues in the Great Basin. This series provides 

managers with brief, accessible summaries of current science concepts related to 

conservation and restoration of sagebrush, as well as associated management strategies. Several FS scientists have contributed to the series. Many fact sheets have been 

produced and are posted on websites and newsletters managed by several interagency 

partnerships, including the Great Basin Landscape Conservation Cooperative’s site 

http://www.greatbasinlcc.org/update/new-great-basin-fact-sheet-series, the Great Basin 

Fire Sciences Exchange site (http://www.gbfiresci.org/), and the Sage Grouse Initiative’s 

site http://www.sagegrouseinitiative.com/category/great-basin-factsheet-series/. 

Treesearch (http://www.treesearch.fs.fed.us/) is an online database system hosted by FS 

R&D that shares free, full-text publications written or cowritten by FS scientists and published by the FS or in other outlets, such as journals, conference proceedings, or books. The 

Treesearch website has been delivering publications since January 2004; more than 40,000 

publications are included with thousands added annually. Treesearch is searchable by using 

keywords, author name, title, and year. A mid-October 2015 search using the single keyword 

“sagebrush” revealed 258 full-text publications in the database.

Conferences, workshops, special sessions at professional society meetings, and training 

sessions are excellent interactive modes for communicating new science findings, establishing researcher and manager connections, discovering new techniques, and gaining new 

knowledge. Libraries now make articles available online and articles are accessed easily 

with online search tools. Online chat sessions are available on many subjects, and literature 

resource services can be easily found on the Web.

A productive way for managers to engage scientists is when they are in the early stages 

of planning management projects, such as a habitat restoration project or during forest plan 

revisions. Another approach to ensure communication and use of science results is for scientists to involve managers during the proposal and implementation phases of science studies 

and assessments through collaboration on shared goals and activities.

5.0—Facilities, Centers, and Projects Having Key 

Locations, Contacts, and Expertise

The Forest Service, Research and Development (FS R&D) has several facilities located 

in sagebrush ecosystems near sage-grouse populations. These are staffed by scientists 

and other professionals with the experience and expertise to conduct continuing and new 

research and science delivery for the benefit of sagebrush and sage-grouse conservation. 

Scientific capacity and resources are expected to decline in the next few years as wellknown scientists in key facilities retire and budget trends continue. Some relevant facilities 

may close owing to declining availability of resources. Workforce planning efforts, capacitybuilding, and additional funding targeted at key locations will be needed to renew capacity 

and resources.

Facilities with past or current studies and expertise on sagebrush and sage-grouse and 

contacts are:

•  Grassland, Shrubland and Desert Ecosystems Science Program, RMRS, Albuquerque, 

New Mexico; Boise, Idaho; Bozeman, Montana; Moscow, Idaho; Provo, Utah; Rapid 

City, South Dakota; Reno, Nevada. Dr. Deborah Finch.
34 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

•  Great Basin Ecology Laboratory, RMRS, Reno, Nevado. Dr. Jeanne Chambers.

•  Great Basin Native Plant Project, RMRS, Boise, Idaho. Dr. Francis Kilkenny and Dr. 

Nancy Shaw, Emeritus.

•  LaGrande Forestry and Range Sciences Laboratory, PNW, LaGrande, Oregon. Mary 

Rowland and Dr. Michael Wisdom.

•  Missoula Fire Sciences Laboratory, RMRS, Missoula, Montana. Dr. Colin Hardy.

•  National Genomics Center for Wildlife and Fish Conservation, RMRS. Missoula, 

Montana. Dr. Michael Schwartz.

•  National Center for Reforestation, Nurseries, and Genetics Resources (RNGR). 

Moscow, Idaho. Dr. Kasten Dumroese and Dr. Jeremiah Pinto.

•  Rapid City Forest and Grassland Research Laboratory, Rapid City, South Dakota. Dr. 

Jack Butler and Dr. Mark Rumble, Emeritus.

•  RMRS Center for Landscape Science, Flagstaff, Arizona. Dr. Samuel Cushman.

•  Shrub Sciences Laboratory, RMRS, Provo, Utah. Dr. Stanley Kitchen, Dr. Susan 

Meyer, Dr. Bryce Richardson, and Dr. Steven Warren.

•  Western Wildland Environmental Threats Assessment Center, PNW, Prineville, 

Oregon. Dr. Nancy Grulke.

•  Wildlife and Terrestrial Ecosystems Program, RMRS, Flagstaff, Arizona; Fort Collins, 

Colorado; Missoula, Montana. Dr. William Block.

6.0—Summary

Forest Service scientists are contributing to the restoration of sagebrush ecosystems and 

conservation of sage-grouse by providing science-based information and specific recommendations for future conservation and management efforts. Immediate research efforts are

needed to establish a scientific foundation for effective management responses that minimize 

threats from fire and invasive species, for restoring functioning sagebrush ecosystems in 

the western portion of the range, for preparing for consequences of climate change, and for 

minimizing threats from energy development in the eastern portion of the range. For decades 

FS Research and Development (R&D) has been studying sagebrush ecosystem dynamics, 

investigating mechanisms to manage for resilient and resistant sagebrush ecosystems, and 

publishing syntheses on the plants and animals that compose sagebrush communities. With this 

extensive experience, FS R&D is prepared to play a relevant role in sagebrush and sage-grouse 

conservation. When this work is combined with FS research on sage-grouse genetics, biology, 

and ecology, along with that of other agencies and organizations, FS expertise will be valuable to efforts for making progress on sage-grouse habitat management. The Forest Service 

will continue to collaborate with others, with the goal of serving the needs of managers 

striving to retain or regain sustainable sagebrush ecosystems and sage-grouse or return them 

to sustainable conditions.

Here we reported on the unique strengths and areas of FS R&D science leadership for 

conserving sage-grouse and restoring sagebrush ecosystems, and we evaluated stakeholder 

interests to identify emphasis areas and priorities for future focus. To meet widespread 

concerns and increased calls for science-based conservation to prevent further declines of 

sage-grouse populations and loss of sagebrush ecosystems, we recommend that research 
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 35

capacity and science delivery be continued and expanded. Because of the importance of the 

National Forest System for sage-grouse breeding and brood-rearing habitats and the FS R&D 

research capacity, strategic locations of facilities, and germane research already completed, 

we believe the Forest Service can make important contributions for sage-grouse management on FS and other landscapes throughout the western United States.

7.0—Acknowledgments

Drs. Francis Kilkenny, Jeffrey Ott, and Nancy Shaw reviewed this manuscript at various 

stages of its development. In addition, BLM comments on the 2015 FS R&D sagebrush and 

sage-grouse conservation science strategy were incorporated into this report. Robin Innes 

provided input on the Fire Effects Information System. Editorial work by Cynthia Moser 

was beneficial in improving the manuscript prior to layout. We are grateful for insightful 

suggestions and support from Cynthia West. We thank Toral Patel-Weynand for vision and 

support for this project. Finally, we acknowledge all Forest Service scientists, and their collaborators, sponsors, postdoctoral researchers, and students who have developed knowledge 

on sagebrush and sage-grouse.

8.0—References

Adair, R.; Johnson, R.C.; Hellier, B.C.; Kaiser, W.J. 2006. Collecting taper-tip onion (Allium 

acuminatum Hook.) in the Great Basin using traditional and GIS methods. Native Plants 

Journal. 7: 141–148.

Aldridge, C.L.; Boyce, M.S. 2007. Linking occurrence and fitness to persistence: habitat 

based approach for endangered greater sage-grouse. Ecological Applications. 17: 

508–526.

Aldridge, C L.; Nielson, S.E.; Beyer, H.L.; Boyce, M.S.; Connelly, J.W.; Knick, S.T.; 

Schroeder, M.A. 2008. Range-wide patterns of greater sage-grouse persistence. Diversity 

and Distributions. 17: 983–994.

Allen, P.S.; Meyer, S.E. 2014. Community structure affects annual grass weed invasion 

during restoration of a shrub-steppe ecosystem. Invasive Plant Science and Management. 

7: 1–13.

Arkle, R.S.; Pilliod, D.S.; Hanser, S.E.; Brooks, M.L.; Chambers, J.C.; Grace, J.B.; Knutson, 

K.C.; Pyke, D.A.; Welty, J.L.; Wirth, T.A. 2014. Quantifying restoration effectiveness 

using multi-scale habitat models: implications for sage-grouse in the Great Basin. 

Ecosphere. 5: 1–32.

Armstrong, J.C. 2007. Improving sustainable seed yield in Wyoming big sagebrush . Provo, 

UT: Brigham Young University. 29 p. Thesis.

Balzotti, C.C.; Kitchen, S.A.; McCarthy, C.W. [In review]. Beyond the single species 

climate envelope: A multifaceted approach to mapping climate change vulnerability. Fort 

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

Station.

Baughman, O.W.; Meyer, S.E.; Aanderud, Z.T.; Leger, E.A. 2016. Cheatgrass die-offs as 

an opportunity for restoration in the Great Basin, USA: Will local or commercial native 

plants succeed where exotic invaders fail? Journal of Arid Environments. 124: 193–204.
36 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Becker, J.M.; Duberstein, C.A.; Tagestad, J.D.; Downs, J.L. 2009. Sage-grouse and wind 

energy: biology, habits, and potential effects from development. PNNL-18567. U.S. 

Department of Energy report under contract DE-ACO5-76RL01830. Springfield, VA:

National Technical Information Service. 35 p.

Beckstead, J.; Augspurger, C.K. 2004. An experimental test of resistance to cheatgrass 

invasion: limiting resources at different life stages. Biological Invasions. 6: 417–432.

Beckstead, J.; Meyer, S.E.; Street, L.E.; Allen, P.S. 2010. Effect of fire on a seed bank 

pathogen and on seeds of its host Bromus tectorum. In: Pendleton, R. Meyer, S.; Schultz, 

B., eds. Conference proceedings: Seed Ecology III - The Third International Society for 

Seed Science Meeting on Seeds and the Environment - “Seeds and Change”; 2010 June 

20–June 24; Salt Lake City, UT. Albuquerque, NM: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station: 10–11.

Beckstead, J.; Street, L.E.; Meyer, S.E.; Allen, P.S. 2011. Fire effects on the cheatgrass seed 

bank pathogen Pyrenophora semeniperda. Rangeland Ecology and Management. 64: 

148–157.

Blank, R.B.; Chambers, J.C.; Roundy, B.A.; Meyer, S.E.; Whittaker, A. 2007. Nutrient 

availability in rangeland soils: influence of prescribed burning, herbaceous vegetation 

removal, over-seeding with Bromus tectorum, season, and elevation. Rangeland Ecology 

and Management. 60: 644–655.

Board, D.I.; Chambers, J.C.; Wright, J.G. 2011. Effects of spring prescribed fire in expanding 

pinyon-juniper woodlands on seedling establishment of sagebrush species. Natural

Resources and Environmental Issues. 16: 1–10.

Bonner, F.T.; Karrfalt, R.P., eds. 2008. The woody plant seed manual. Agric. Handb. 727. 

Washington, DC: U.S. Department of Agriculture, Forest Service. 1223 p.

Bower, A.D.; St. Clair, J.B.; Erickson, V. 2014. Generalized provisional seed zones for 

native plants. Ecological Applications. 24: 913–-919.

Bradley, B.A. 2010. Assessing ecosystem threats from global and regional change: 

Hierarchical modeling of risk to sagebrush ecosystems from climate change, land use and 

invasive species in Nevada, USA. Ecography. 33: 198–208.

Bradley, B.A., C.A. Curtis; J.C. Chambers. 2016. Chapter 9. Bromus response to climate and 

projected change with climate change. In: Germino, M.J.; J.C. Chambers; C.S. Brown, 

editors. Exotic Brome-Grasses in Arid and Semiarid Ecosystems of the Western US: 

Causes, Consequences, and Management Implications. Springer Press, New York.

Braun, C.E. 1998. Sage-grouse declines in western North America: What are the problems? 

Proceedings of the Western Association of State Fish and Wildlife Agencies. 78: 139–156.

Brooks, M.L.; Chambers, J.C. 2011. Resistance to invasion and resilience to fire in desert

shrublands of North America. Rangeland Ecology and Management. 60: 431–438.

Cane, J.H. 2008. Breeding biologies, seed production and species-rich bee guilds of Cleome

lutea and C. serrulata (Cleomaceae). Plant Species Biology. 23: 152–158.

Chambers, J.C. 2000. Seed movements and seedling fates in disturbed sagebrush steppe

ecosystems: Implications for restoration. Ecological Applications. 10: 1400–1413.

Chambers, J.C. 2001. Pinus monophylla establishment in an expanding Pinus-Juniperus 

woodland: Environmental conditions, facilitation and interacting factors. Journal of 

Vegetation Science. 12: 27–40.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 37

Chambers, J.C. 2005. Fire related restoration issues in woodland and rangeland ecosystems. 

In: Taylor, L.; Zelnik, J.; Cadwallader, S.; Hughes, B., comps. Mixed fire regimes: 

Ecology and management. Symposium Proceedings; 2004 November 17–19. Association 

of Fire Ecology MIXC03. Spokane, WA: 149–160.

Chambers, J.C.; Eldredge, E.P.; Snyder, K.A.; Board, D.I.; Forbis de Queiroz, T.; Hubbard, 

V. 2014a. Restoring abandoned agricultural lands in cold desert shrublands: Tradeoffs 

between water availability and invasive species. Invasive Plant Science and Management. 

7: 176–189.

Chambers, J.C.; Leger, E.; Goergen, E. 2009. Cold desert fire and invasive species 

management: Resources, strategies, tactics and responses. Rangelands. 31: 14–20.

Chambers, J.C.; Linnerooth, A.R. 2001. Restoring sagebrush dominated riparian corridors 

using alternative state and threshold concepts: Environmental and seedling establishment

response. Applied Vegetation Science. 4: 157–166.

Chambers, J.C.; MacMahon, J.A. 1994. A day in the life of a seed: Movements and fates of 

seeds and their implications for natural and managed systems. Annual Review of Ecology 

and Systematics. 25: 263–292.Chambers, J.C.; Miller, R.F.; Board, D.I.; Grace, J.B.; 

Pyke, D.A.; Roundy, B.A.; Schupp, E.W.; Tausch, R.J. 2014b. Resilience and resistance 

of sagebrush ecosystems: Implications for state and transition models and management 

treatments. Rangeland Ecology and Management. 67: 440–454.

Chambers, J.C.; Miller, R.F.; Grace, J.B.; Pyke, D.A.; Bradley, B.; Hardegree, S.P.; 

D’Antonio, C. 2014c. Resilience to disturbance and resistance to invasive alien grasses in 

arid and semi-arid ecosystems: Lessons from the cold desert. Ecosystems. 17: 360–375.

Chambers, J.C.; Pyke, D.A.; Maestas, J.; Pellent, M.; Boyd, C.S.; Campbell, S.; Espinosa, 

S.; Havelina, D.; Mayer, K.; Wuenschel, A. 2014d. Using resistance and resilience 

concepts to reduce impacts of annual grasses and altered fire regimes on the sagebrush 

ecosystem and sage-grouse ― A strategic multi-scale approach. Gen. Tech. Rep. RMRSGTR-326. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 73 p.

Chambers, J.C.; Roundy, B.A.; Blank, R.R.; Meyer, S.E.; Whittaker, A. 2007. What 

makes Great Basin sagebrush ecosystems invasible by Bromus tectorum? Ecological 

Monographs. 77: 117–145.

Chambers, J.C.; Vander Wall, S.B.; Schupp, E.W. 1999. Seed and seedling ecology of 

piñon and juniper species in the pygmy woodlands of western North America. Botanical

Reviews. 65: 1–38.

Condon, L.; Weisberg, P.J.; Chambers, J.C. 2011. Abiotic and biotic influences on Bromus 

tectorum invasion and Artemisia tridentata recovery after fire. International Journal of 

Wildland Fire. 20: 597–604.

Connelly, J.W.; Knick, S.T.; Braun, C.E.; Baker, W.L.; Beever, E.A.; Christiansen, T.; 

Doherty, K.E.; Garton, E.O.; Hanser, S.E. ; Johnson, D.H. ; Leu, M. ; Miller, R.F.; 

Naugle, D.E.; Oyler-McCance, S.J.; Pyke, D.A.; Reese, K.P.; Schroeder, M.A.; Stiver, 

S.J.; Walker, B.L.; Wisdom, M.J. 2011. Conservation of greater sage-grouse: A synthesis 

of current trends and future management. In: Knick, S.T.; Connelly, J.W., eds. Greater 

sage-grouse: ecology and conservation of a landscape species and habitats. Berkeley, CA: 

University of California Press: 549–563.
38 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Connelly, J.W.; Reese, K.P.; Schroeder, M.A. 2003. Monitoring of greater sage-grouse 

habitats and populations. Station Bull. 80. Moscow, ID: University of Idaho, College of 

Natural Resources Experiment Station.

Connelly, J.W.; Schroeder, M.A.; Sands, A.R.; Braun, C.E. 2000. Guidelines to manage 

sage-grouse populations and their habitats. Wildlife Society Bulletin. 28: 967–985.

Copeland, H.E.; Doherty, K.E.; Naugle, D.E.; Pocewicz, A.; Kiesecker, J.M. 2009. Mapping 

oil and gas development potential in the US Intermountain West and estimating impacts 

to species. PloS ONE. 4(10): e7400.

Cross, T.B., D. Naugle, J.C. Carlson, and M.K. Schwartz. [In review]. Greater sagegrouse hierarchical population genetic structure: alignment with physiogeographic and 

contemporary landscape features, and insights into management. Conservation Genetics.

Currans, S.P.; Kitchen, S.G.; Lambert, S.M. 1997. Ensuring identity and quality of native 

seeds. In: Shaw, N.L.; Roundy, B.A., comps. Proceedings: Using seeds of native 

species on rangelands. Gen. Tech. Rep. INT-GTR-372. Ogden, UT: U.S. Department of 

Agriculture, Forest Service, Intermountain Research Station: 17–20.

Cushman, S.A.; McKelvey, K.S.; Hayden, J.; Schwartz, M.K. 2006. Gene flow in complex 

landscapes: Testing multiple hypotheses with causal modeling. The American Naturalist.

168: 486–499.

D’Antonio, C.; Chambers, J.C. 2006. Using ecological theory to manage or restore

ecosystems affected by invasive plant species. In: Falk, D.; Palmer, M.; Zedler, J., eds. 

Foundations of restoration ecology. Covelo, CA: Island Press: 260–279.

Davies, K.W.; Boyd, C.S.; Beck, J.L.; Bates, J.D.; Svejcar, T.J.; Gregg, M.A. 2011. Saving 

the sagebrush sea: An ecosystem conservation plan for big sagebrush plant communities. 

Biological Conservation. 144: 2573–2584.

Davis, A.S.; Herriman, K.R.; Apostol, K.G.; Kildisheva, O.A.; Ross-Davis, A.L.; Dumroese, 

R.K. [In press]. Do container volume, site preparation, and field fertilization affect 

restoration potential of Wyoming big sagebrush? Natural Areas Journal.

Doherty, K.E.; Naugle, D.E.; Walker, B.L. 2010. Greater sage-grouse nesting habitat: 

The importance of managing at multiple scales. Journal of Wildlife Management. 74: 

1544–1553.

Drapek, R.J.; Kim, J.B.; Neilson, R.P. 2015. The Dynamic General Vegetation Model MC1 

over the United States and Canada at a 5-arcminute resolution: model inputs and outputs. 

Gen. Tech. Rep. PNW-GTR-904. Portland, OR: U.S. Department of Agriculture, Forest 

Service, Pacific Northwest Research Station. 57 p.

Dumroese, R.K.; Landis, T.D.; Luna, T. 2012. Raising native plants in nurseries: basic 

concepts. Gen. Tech. Rep. RMRS-GTR-274. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 84 p.

Dumroese, R.K.; Luna, T.; Landis, T.D., eds. 2009. Nursery manual for native plants: 

A guide for tribal nurseries - Volume 1: Nursery management. Agric. Handb. 730. 

Washington, DC: U.S. Department of Agriculture, Forest Service. 302 p.

Dumroese, R.K.; Luna, T.; Pinto, J.R.; Landis, T.D. [In press]. Forbs: Foundation for 

restoration of pollinators, monarch butterflies, and Greater Sage-Grouse in the western 

United States. Natural Areas Journal.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 39

Dumroese, R.K.; Luna, T.; Richardson, B.A.; Kilkenny, F.F.; Runyon, J.B. 2015. Conserving 

and restoring habitat for Greater Sage-Grouse and other sagebrush-obligate wildlife: The 

crucial link of forbs and sagebrush diversity. Native Plants Journal. 16: 276–299.

Endangered Species Act of 1973 [ESA]; 16 U.S.C. 1531-1536, 1538-1540.

Finch, D.; Boyce, D.; Chambers, J.; Colt, C.; McCarthy, C.; Kitchen, S.; Richardson, B.; 

Rowland, M.; Rumble, M.; Schwartz, M.; Tomosy, M.; Wisdom, M. 2015. USDA 

Forest Service Sage-Grouse Conservation Science Strategy. Washington, DC: U.S. 

Department of Agriculture, Forest Service. http://www.fs.fed.us/research/docs/wildlifefish/sage-grouse-conservation-strategy.pdf. [Accessed January 31, 2016].

Finch, D.M., ed. 2012. Climate change in grasslands, shrublands, and deserts of the interior 

American West: A review and needs assessment. Gen. Tech. Rep. RMRS-GTR-285. Fort 

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

Station. 139 p.

Finch, D.M.; Smith, D.M.; LeDee, O.; Cartron, J.E.; Rumble, M.A. 2012. Chapter 5: 

Climate change, animal species, and habitats: Adaptation and issues. In: Finch, D.M., ed. 

Climate change in grasslands, shrublands, and deserts of the interior American West: A 

review and needs assessment. Gen. Tech. Rep. RMRS-GTR-285. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station: 60–79.

Fremgen, A.L.; Hansen, C.P.; Rumble, M.A.; Gamo, R.S.; Millspaugh, J.J. 2016. Male 

greater sage-grouse detectability on leks. Journal of Wildlife Management. 80:266–274.

Friggens, M.M.; Warwell, M.V.; Chambers, J.C.; Kitchen, S.G. 2012. Chapter 1: Modeling 

and predicting vegetation response of western USA grasslands, shrublands, and deserts 

to climate change. In: Finch, D.M., ed. Climate change in grasslands, shrublands, and 

deserts of the interior American West: A review and needs assessment. Gen. Tech. Rep. 

RMRS-GTR-285. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station: 1–20.

Goergen, E.; Chambers, J.C. 2009. Influence of a native legume on soil N and plant response

following prescribed fire in sagebrush steppe. International Journal of Wildland Fire

Science. 18: 1–11.

Goldstein, M.I.; Suring, L.H.; Vojta, C.D.; Rowland, M.M.; McCarthy, C. 2013. Chapter 10: 

Developing a habitat monitoring program: three examples from National Forest planning. 

In Rowland, M.M.; Vojta, C.D. Vojta, tech. eds. A technical guide for monitoring wildlife 

habitat. Gen. Tech. Rep. WO-GTR-89. Washington, DC: U.S. Department of Agriculture, 

Forest Service: 10-1 through 10-74.

Goodrich, S.; Huber, A.; Monroe, B. 2008. Trend in mountain big sagebrush crown cover 

and ground cover on burned sites, Uinta Mountains and West Tavaputs Plateau, Utah. 

In: Kitchen, S.G.; Pendleton, R.L.; Monaco, T.A.; Vernon, J., comps. Proceedings―

Shrublands under fire: Disturbance and recovery in a changing world. Proc. RMRS-P-52. 

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

Research Station: 151–160.

Haase, D.L.; Pinto, J.R.; Dumroese, K.R.; Hernandez, G.; Karrfalt, B.; Overton, R. 2011. 

RNGR: A national resource for reforestation, restoration, and nursery professionals. Tree 

Planters Notes. 54(1): 28–34.
40 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Hanks, E.M.; Hooten, M.B.; Knick, S.T.; Oyler-McCance, S.J.; Fike, J.A.; Cross, T.B.; 

Schwartz, M.K. [In review]. Latent spatial models and sampling design for landscape 

genetics. Annals of Applied Statistics.

Hann, W.J.; Wisdom, M.J.; Rowland, M.M. 2003. Disturbance departure and fragmentation 

of natural systems in the interior Columbia Basin. Res. Pap. PNW-RP-545. Portland, OR: 

U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 19 p.

Hansen, C.P.; Rumble, M.A.; Gamo, S.; Millspaugh, J.J. 2014. Auxiliary VHF transmitter 

to aid recovery of solar Argos/GPS PTT. Res. Note RMRS-RN-72. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station. 11 p.

Hanser, S.E.; Aldridge, C.L.; Leu, M.; Rowland, M.M.; Nielsen, S.E.; Knick, S.T. 2011a. 

Greater sage-grouse: General use and roost site occurrence with pellet counts as a measure 

of relative abundance. In: Hanser, S.E.; Leu, M.; Knick, S.T.; Aldridge, C.L. eds. 

Sagebrush ecosystem conservation and management: Ecoregional assessment tools and 

models for the Wyoming Basins. Lawrence, KS: Allen Press: 112–140.

Hanser, S.E.; Knick, S.T. 2011. Greater sage-grouse as an umbrella species for shrubland 

passerine birds: A multiscale assessment. Studies in Avian Biology. 38: 475–488.

Hanser, S.E.; Leu, M.; Knick, S.T.; Aldridge, C.L., eds. 2011b. Sagebrush ecosystem

conservation and management: Ecoregional assessment tools and models for the

Wyoming Basins. Lawrence, KS: Allen Press. 429 p.

Hemstrom, M.A., M.J. Wisdom, W.J. Hann, M.M. Rowland, and R.A. Gravenmier. 2002. 

Sagebrush-steppe vegetation dynamics and restoration potential in the Interior Columbia

Basin, USA. Conservation Biology. 16: 1243–1255.

Herman-Brunson, K M.; Jensen, K.C.; Kaczor, N.W.; Swanson, C.C.; Rumble, M.A.; Klaver, 

R.W. 2009. Nesting ecology of greater sage-grouse Centrocercus urophasianus at the

eastern edge of their historic distribution. Wildlife Biology. 15 :237–246.

Herrick, J.E.; Duniway, M.C.; Pyke, D.A.; Bestelmeyer, B.T.; Wills, S.A.; Brown, J.R.; Karl, 

J.W.; Havstad, K.M. 2012. A holistic strategy for adaptive land management. Journal of 

Soil and Water Conservation. 67: 105A–113A.

Herriman, K.R.; Davis, A.S.; Dumroese, R.K. 2009. Influence of container size on Wyoming 

big sagebrush seedling morphology and cold hardiness. In: Dumroese, R.K.; Riley, L.E., 

tech. coords. National proceedings: Forest and Conservation Nursery Associations-2008. 

Proceedings RMRS-P-58. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station: 44–47.

Hess, J.; Beck, J.L.  2012.  Disturbance factors influencing greater sage-grouse lek 

abandonment in north-central Wyoming. Journal of Wildlife Management. 76: 1625–1634.

Heyerdahl, E.K.; Miller, R.F.; Parsons, R.A. 2006. History of fire and Douglas-fir 

establishment in a savanna and sagebrush–grassland mosaic, southwestern Montana, 

USA. Forest Ecology and Management. 230: 107–118.

Iverson, L.R.; Prasad, A.M.; Matthews, S.N.; Peters, M. 2008. Estimating potential 

habitat for 134 eastern US tree species under six climate scenarios. Forest Ecology and 

Management. 254: 390–406.

Jensen, S. 2004. Racquets, hoppers, and felt boards―Low-tech devices for processing seeds. 

Native Plants Journal. 5: 50–51.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 41

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.

Jones, R.; Chambers, J.C.; Board, D.I.; Johnson, D.W.; Blank, R.R. 2015a. The role of 

resource limitation in restoration of sagebrush ecosystems dominated by cheatgrass. 

Ecosphere. 6: Article 107.

Jones, R.; Chambers, J.C.; Johnson, D.W.; Blank, R.R.; Board, D.I. 2015b. Effect of repeated 

burning on plant and soil carbon and nitrogen in cheatgrass (Bromus tectorum) dominated 

ecosystems. Plant and Soil. 386: 47–64.

Kaczor, N.W.; Herman-Brunson, K.M.; Jensen, K.C.; Rumble, M.A.; Klaver, R.W.; 

Swanson, C.C. 2011a. Resource selection during brood-rearing by greater sage-grouse. 

Studies in Avian Biology. 39: 169–177.

Kaczor, N.W.; Jensen, K.C.; Klaver, R.W.; Rumble, M.A.; Herman-Brunson, K.M. Swanson, 

C.C. 2011b. Nesting success and resource selection of greater sage-grouse. Studies in 

Avian Biology. 39: 107–118.

Karrfalt, R.P.; Shaw, N. 2013. Banking Wyoming big sagebrush seeds. Native Plants Journal. 

14: 60–69.

Kiehl, K.; Kirmer, A.; Shaw, N.; Tischew, S., eds. 2014. Guidelines for native seed 

production and grassland restoration. Newcastle upon Tyne, UK: Cambridge Scholars. 

315 p.

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

In: Haase, D.L.; Pinto, J.R.; Wilkinson, K.M., tech. coord. National proceedings: Forest 

and Conservation Nursery Associations - 2012. Proc. RMRS-P-69. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station: 87–89.

Kilkenny, F.F. 2015. Genecological approaches to the effects of climate change on plant

populations. Natural Areas Journal. 35: 152–164.

Kitchen, S.G. 1994. Perennial forb life-history strategies on semiarid rangelands: 

Implications for revegetation. Monsen, S.B.; Kitchen, S.G., comps. Proceedings: Ecology 

and management of annual rangelands. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. 

Department of Agriculture, Forest Service, Intermountain Research Station: 342–346.

Kitchen, S.G. 1995. Return of the native―a look at select accessions of North American 

Lewis flax (Linum lewisii). In Roundy, B.A.; McArthur, E.D.; Haley, J.S.; Mann, D.K., 

comps. Proceedings: Wildland shrub and arid land restoration symposium. Gen. Tech. 

Rep. INT-GTR-315. Ogden, UT: U.S. Department of Agriculture, Forest Service, 

Intermountain Research Station: 321–326.

Kitchen, S.G. 2001. Intra-specific variability in germination behavior and seed testing 

protocols: The challenge of Intermountain species. Seed Technology. 23: 68–77.

Kitchen, S.G. 2014. Learning to live with cheatgrass: Giving up or a necessary paradigm 

shift? Rangelands. 36: 32–36.

Kitchen, S.G.; McArthur, E.D. 2007. Big and black sagebrush landscapes. In: Hood, 

S.M.; Miller, M., tech. eds. Fire ecology and management of the major ecosystems of 

southern Utah. Gen. Tech. Rep. RMRS-GTR-212. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station: 73–95.
42 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Kitchen S.G.; Monsen, S.B. 1994. Germination rate and emergence success in bluebunch 

wheatgrass. Journal of Range Management. 47: 145–150.

Kitchen, S.G.; Monsen, S.B. 1996. Arrowleaf balsamroot (Balsamorhiza sagittata) seed 

germination and establishment success (Utah). Restoration and Management Notes. 14: 

180–181.

Knick, S.T.; Connelly, J.W., eds. 2011. Greater sage-grouse: Ecology and conservation of a 

landscape species and its habitats. Studies in Avian Biology 38. Berkeley, CA: University 

of California Press. 644 p.

Knick, S.T.; Dobkin, D.S.; Rotenberry, J.T.; Schroeder, M.A.; Vander Haegen, W.M.; Van 

Riper, C., III. 2003. Teetering on the edge or too late? Conservation and research issues 

for avifauna of sagebrush habitats. The Condor. 105: 611–634.

Knick, Steven T.; Hanser, Steven E.; Leu, Matthias; Aldridge, Cameron L.; Neilsen, 

Scott E.; Rowland, Mary M.; Finn, Sean P.; Wisdom, Michael J. 2011. Chapter 11. 

Management considerations. In: Hanser, S.E.; Leu, M.; Knick, S.T.; Aldridge, C.L., eds. 

Sagebrush ecosystem conservation and management: ecoregional assessment tools and 

models for the Wyoming Basins. Lawrence, KS: Allen Press: 387–409.

Knick, S.T.; Hanser, S.E.; Preston, K.L. 2013. Modeling ecological minimum requirements 

for distribution of greater sage-grouse leks: Implications for population connectivity 

across their western range, U.S.A. Ecology and Evolution. 3: 1539–1551.

Knutson, K.C.; Pyke, D.A.; Wirth, T.A.; Arkle, R.S.; Pilliod, D.S.; Brooks, M.L.; Chambers, 

J.C.; Grace, J.B. 2014. Long-term effects of seeding after wildfire on vegetation in Great 

Basin shrubland ecosystems. Journal of Applied Ecology. 51: 1414–1424.

Korfmacher, J.L.; Chambers, J.C.; Tausch, R.J.; Roundy, B.A.; Meyer, S.E.; Kitchen, S. 

2003. Technical note: A technique for conducting small-plot burn treatments. Journal of 

Range Management. 56: 251–254.

Landeen, M.L. 2015. Mountain big sagebrush (Artemisia tridentata ssp. vaseyana) seed 

production. Provo, UT: Brigham Young University. 71 p. Master’s thesis.

Manel, S.; Schwartz, M.K.; Luikart, G.; Taberlet, P. 2003. Landscape genetics: The

combination of landscape ecology and population genetics. Trends in Ecology and 

Evolution. 18: 1807–1816.

Marcot, B.G.; Holthausen, R.S.; Raphael, M.G.; Rowland, M.M.; Wisdom, M.J. 2001. Using 

Bayesian belief networks to evaluate wildlife population viability under land management 

alternatives from an environmental impact statement. Forest Ecology and Management.

153: 29–42.

Masi, M.; Evidente, A.; Meyer, S.; Nicholson, J.; Munoz, A. 2014. Effect of strain and 

cultural conditions on the production of cytochalasin B by the potential mycoherbicide 

Pyrenophora semeniperda (Pleosporaceae, Pleosporales). Biocontrol Science and 

Technology. 24(1): 53–64.

Mazzola, M.B.; Allcock, K.G.; Chambers, J.C.; Blank, R.R.; Nowak, R.S. 2008. Effects of 

nitrogen availability and cheatgrass competition on the establishment of Vavilov Siberian 

wheatgrass. Rangeland Ecology and Management. 61: 475–484.

Mazzola, M.B.; Chambers, J.C.; Pyke, D.; Schupp, E.W.; Blank, R.R.; Allcock, K.G.; 

Nowak, R.S. 2011. Effects of resource availability and propagule supply on native species 

recruitment in sagebrush ecosystems invaded by Bromus tectorum. Biological Invasions. 

13: 513–526.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 43

McAdoo, J.K.; Boyd, C.S.; Sheley, R.L. 2013. Site, competition, and plant stock influence

transplant success of Wyoming Big Sagebrush. Rangeland Ecology and Management. 66: 

305–312.

McArthur, E.D.; Welch, B.L. 1982. Growth rate differences among big sagebrush (Artemisia 

tridentata) accessions and subspecies. Journal of Range Management. 35: 396–401.

McArthur, E.D.; Welch, B.L.; Sanderson, S.C. 1988. Natural and artificial hybridization 

between big sagebrush (Artemisia tridentata) subspecies. Journal of Heredity. 79: 

268–276.

McIver, J.; Brunson, M.; Bunting, S.; Chambers, J.; Devoe, N.; Doescher, P.; Grace, J.; 

Johnson, D.; Knick, S.; Miller, R.; Pellant, M.; Pierson, F.; Pyke, D.; Rollins, K.; Roundy, 

B.; Schupp, E.; Tausch, R.; Turner, D. 2010. The Sagebrush Steppe Treatment Evaluation 

Project (SageSTEP): A test of state-and-transition theory. Gen. Tech. Rep. RMRSGTR-237. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station. 16 p.

McWilliams, J. 2002. Centrocercus minimus, C. urophasianus. In: Fire Effects Information 

System. Missoula, MT: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station, Fire Sciences Laboratory. http://www.feis-crs.org/feis/. [Accessed 

August 17, 2015].

Merrill, K.R.; Meyer, S.E.; Coleman, C.E. 2012. Population genetic analysis of Bromus 

tectorum (Poaceae) indicates recent range expansion may be facilitated by specialist

genotypes. American Journal of Botany. 99: 529–537.

Meyer, S.E. 1994. Germination and establishment ecology of big sagebrush: implications 

for community restoration. In Monsen, S.B.; Kitchen, S.G., comps. Proceedings: Ecology 

and management of annual rangelands. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. 

Department of Agriculture, Forest Service, Intermountain Research Station: 244–251.

Meyer, S.E.; Beckstead, J.; Allen, P.S.; Smith, D.C. 2008. A seed bank pathogen causes 

seedborne disease: Pyrenophora semeniperda on undispersed grass seeds in western 

North America. Canadian Journal of Plant Pathology. 30(4): 525–533.

Meyer, S.E.; Clement, S.; Beckstead, J. 2013. Annual brome control using a native fungal 

seed pathogen. United States Patent Application Publication No. US 2013/0035231 A1. 

Alexandria, VA: U.S. Patent and Trademark Office. 23 p.

Meyer, S.E.; Franke, J.-L.; Baughman, O.W.; Beckstead, J.; Geary, B. 2014. Does Fusariumcaused seed mortality contribute to Bromus tectorum stand failure in the Great Basin? 

Weed Research. 54: 511–519.

Meyer, S.E.; Garvin, S.C.; Beckstead, J. 2001. Factors mediating cheatgrass invasion of 

intact salt desert shrubland. In McArthur, D.E.; Fairbanks, D.J., comps. Proceedings—

Shrubland ecosystem genetics and biodiversity. Proc. RMRS-P-21. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station: 224–232.

Meyer, S.E.; Kitchen, S.G. 1994a. Habitat-correlated variation in seed germination response

to chilling in Penstemon Section Glabri (Scrophulariaceae). American Midland Naturalist.

132: 349–365.

Meyer, S.E.; Kitchen, S.G. 1994b. Life history variation in blue flax (Linum perenne:

Linaceae): seed germination phenology. American Journal of Botany. 81: 528–535.
44 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Meyer S.E.; Kitchen, S.G.; Carlson, S.L. 1995. Seed germination patterns in Intermountain 

penstemons (Scrophulareaceae). American Journal of Botany. 82: 377–389.

Meyer, S.E.; Masi, M.; Clement, S.; Davis, T.L.; Beckstead, J. 2015. Mycelial growth rate 

and toxin production in the seed pathogen Pyrenophora semeniperda: Resource trade-offs 

and temporally varying selection. Plant Pathology. 60:1450–1460.

Meyer, S.E.; Monsen, S.B. 1991. Habitat-correlated variation in mountain big sagebrush 

(Artemisia tridentata ssp. vaseyana) seed germination patterns. Ecology. 72: 739–742.

Meyer, S.E.; Nelson, D.L.; Clement, S. 2001. Evidence for resistance polymorphism in the 

Bromus tectorum - Ustilago bullata pathosystem: Implications for biocontrol. Canadian 

Journal of Plant Pathology. 23: 19–27.

Meyer, S.E.; Nelson, D.L.; Clement, S.; Paulsen, A. 2000. Exploring the potential for 

biocontrol of cheatgrass with the head smut pathogen. In: Entwistle, P.G.; DeBolt, A.M.; 

Kaltenecker, J.H. Steenholf, K., comps. Proceedings: Sagebrush steppe ecosystem 

symposium; 1999 June 23–25; Boise, ID. BLM/ID/PT-001001+1150. Boise, ID: U. S. 

Department of the Interior, Bureau of Land Management, Idaho State Office: 70–72.

Meyer, S.E.; Nelson, D.L.; Clement, S.; Ramakrishnan, A. 2010a. Ecological genetics of 

the Bromus tectorum (Poaceae) - Ustilago Bullata (Ustilaginaceae): A role for frequency 

dependent selection? American Journal of Botany. 97(8): 1304–1312.

Meyer, S.E.; Stewart, T.E.; Clement, S. 2010b. The quick and the deadly: Growth versus 

virulence in a seed bank pathogen. New Phytologist. 187: 207–216.

Meyer, S.E.; Warren, T.W 2015. Seeding big sagebrush successfully on intermountain 

rangelands. Great Basin Fact Sheet Series. 10: 1–5.

Miglia, K.J.; McArthur, E.D.; Redman, R.S.; Rodriguez, R.J.; Zak, J.C.; Freeman, D.C. 

2007. Genotype, soil type, and locale effects on reciprocal transplant vigor, endophyte 

growth, and microbial functional diversity of a narrow sagebrush hybrid zone in Salt 

Creek Canyon, Utah. American Journal of Botany. 94: 425–436.

Miller R.F.; Chambers, J.C.; Pellant, M. 2014a. A field guide to selecting the most

appropriate treatments in sagebrush and pinyon-juniper ecosystems in the Great Basin:

Evaluating resilience to disturbance and resistance to invasive annual grasses and 

predicting vegetation response. Gen. Tech. Rep. RMRS-GTR-322. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station. 66 p.

Miller R.F.; Chambers, J.C.; Pellant, M. 2015. A field guide for rapid assessment of postwildfire recovery potential in sagebrush and piñon-juniper ecosystems in the Great 

Basin: Evaluating resilience to disturbance and resistance to invasive annual grasses and 

predicting vegetation response. Gen. Tech. Rep. RMRS-GTR-338. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station. 70 p.

Miller, R.F.; Chambers, J.C.; Pyke, D.A.; Pierson, F.B.; Williams, C.J. 2013. A review of fire 

effects on vegetation and soils in the Great Basin Region: Response and ecological site 

characteristics. Gen. Tech. Rep. RMRS-GTR-308. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 136 p.

Miller, R.F.; Heyerdahl, E.K. 2008. Fine-scale variation of historical fire regimes in 

sagebrush-steppe and juniper woodland: An example from California, USA. International 

Journal of Wildland Fire. 17: 245–254.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 45

Miller, R.F.; Ratchford, J.; Roundy, B.A.; Tausch, R.J.; Pereia, C.; Hulet, A.; Chambers, J.C. 

2014b. Response of conifer encroached shrublands in the Great Basin to prescribed fire

and mechanical treatments. Rangeland Ecology and Management. 67: 468–481.

Miller, R.F.; Rose, J.A. 1999. Fire history and western juniper encroachment in sagebrush 

steppe. Journal of Range Management. 52: 550–559.

Miller, R.F.; Tausch, R.J.; McArthur, E.D.; Johnson, D.D.; Sanderson, S.C. 2008. Age 

structure and expansion of piñon-juniper woodlands: A regional perspective in the 

Intermountain West. Res. Pap. RMRS-RP-69. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 15 p.

Monsen, S.; Stevens, R.; Shaw, N. 2004a. Restoring western ranges and wildlands. Vols. 

I, II, III. Gen. Tech. Rep. RMRS-GTR-136. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station.

 Monsen, S.B.; Kitchen, S.G., comps. 1994. Proceedings: Ecology and management of 

annual rangelands. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. Department of 

Agriculture, Forest Service, Intermountain Research Station. 416 p.

Monsen, S.B.; Kitchen, S.G.; Memmott, K.; Shaw, N.; Pellant, M.; Young, S.; Ogle, D.; 

St. John, L. 2004b. Notice of Release Anatone Bluebunch Wheatgrass (Selected Class 

Natural Population). Provo, UT: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station; Boise, ID: Department of the Interior, Bureau of Land 

Management; Ephraim, UT: Utah Department of Natural Resources, Division of Wildlife 

Resources; Logan, UT: Utah State University, Agricultural Experiment Station; Boise, 

ID: U.S. Department of Agriculture Natural Resources Conservation Service; Moscow, 

ID: University of Idaho, Agricultural Experiment Station. 9 p.

Monsen, S.B.; Meyer, S.E. 1990. Seeding equipment effects on establishment of big 

sagebrush on mine disturbances. In: Munkshower, F. ed. Fifth Billings Symposium on 

Disturbed Land Rehabilitation.Volume 1, Hardrock waste, analytical, and revegetation. 

1990 March 25–30; Billings, MT. Pub. 9003. Bozeman, MT: Montana State University, 

Reclamation Research Unit: 192–199.

Monsen, S.B.; Meyer, S.E.; Carlson, S.L. 1992. Sage-brush establishment enhanced by 

snowfencing. In: Rangeland Technology and Equipment Council, USDA Forest Service 

Technology and Development Program 2200-Range. 1992 Annual Report: 6–8.

Overton, E.C.; Pinto, J.R.; Davis, A.S. 2013. Insights into big sagebrush seedling storage 

practices. Native Plants Journal. 14: 225-230.

Naugle, D.E., ed. 2011. Energy development and wildlife conservation in western North 

America. Washington, DC: Island Press. 344 p.

Naugle, D.E.; Aldridge, C.L.; Walker, B.L.; Cornish, T.E.; Moynahan, B.J.; Holloran, M.J.; 

Brown, K.; Johnson, G.D.; Schmidtmann, E.T.; Mayer, R.T. 2004. West Nile virus: 

Pending crisis for greater sage-grouse. Ecology Letters. 7: 704-713.

Naugle, D.E.; Doherty, K.E.; Walker, B.L.; Copel, H.E.; Holloran, M.J.; Tack, J.D. 2011. 

Sage-grouse and cumulative impacts of energy development. In Naugle, D.E., ed. Energy 

development and wildlife conservation in western North America. Washington, DC: 

Island Press: 55–70.
46 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Naugle, D.E.; Walker, B.L.; Doherty, K.E. 2006. Sage-grouse population response to coalbed natural gas development in the Powder River Basin: Interim progress report on 

region-wide lek-count analyses. Missoula, MT: University of Montana, Wildlife Biology 

Program, College of Forestry and Conservation.

Nelson, Z.J.; Weisberg, P.J.; Kitchen, S.G. 2014. Influence of climate and environment on 

post-fire recovery of mountain big sagebrush. International Journal of Wildland Fire. 23: 

131–142.

Noss, R.F.; LaRoe, E.T., III; Scott, J.M. 1995. Endangered ecosystems of the United States: 

A preliminary assessment of loss and degradation. U.S. Department of the Interior, 

National Biological Service. National Biological Service Report 28. Washington DC.

Ott, J.E.; Cox, R.D.; Shaw, N.L.; Pellant, M.; Roundy, B.A. [In preparation]. Post-fire 

seeding techniques for Great Basin native plants: Comparisons of conventional and 

minimum-till drills.

Ott, J.E.; McArthur, E.D.; Roundy, B.A. 2003. Vegetation of chained and non-chained 

seedlings after wildfire in Utah. Journal of Range Management. 56: 81–91.

Overton, E.C.; Pinto, J.R.; Davis, A.S. 2013. Insights into big sagebrush seedling storage 

practices. Native Plants Journal. 14: 225–230.

Parkinson, H.; Zabinski, C.; Shaw, N. 2013. Impact of native grasses and cheatgrass 

(Bromus tectorum) on Great Basin forb seedling growth. Rangeland Ecology and 

Management. 66: 174–180.

Pearson, R.G.; Dawson, T.P. 2003. Predicting the impacts of climate change on the

distribution of species: Are bioclimate envelope models useful? Global Ecology and 

Biogeography. 12: 361–371.

Pinto, J.R.; Marshall, J.D.; Dumroese, R.K.; Davis, A.S.; Cobos, D.R. 2012. Photosynthetic 

response, carbon isotopic composition, survival, and growth of three stocktypes under 

water stress enhanced by vegetative competition. Canadian Journal of Forest Research. 

42: 333–344.

Pinto, J.R.; Marshall, J.D.; Dumroese, R.K.; Davis, A.S.; Cobos, D.R. 2015. Seedling 

establishment and physiological responses to temporal and spatial soil moisture changes. 

New Forests. 42. doi: 10.1007/s11056-015-9511-7.

Pyke, D.A.; Knick, S.T.; Chambers, J.C.; Pellant, M.; Miller, R.F.; Beck, J.L.; Doescher, 

P.S.; Schupp, E.W.; Roundy, B.A.; Brunson, M.; McIver, J.D., 2015. Restoration 

handbook for sagebrush steppe ecosystems with emphasis on greater sage-grouse 

habitat—Part 2. Landscape level restoration decisions: U.S. Geological Survey Circular 

1418. 21 p. http://dx.doi.org/10.3133/cir1418.

Pyke, D.A.; Shaff, S.E.; Lindgren, A.; Doescher, P.S.; Schupp, E.W.; Chambers, J.C.; 

Burnham, J.S.; Huso, M.M. 2014. Region-wide ecological responses of arid Wyoming 

big sagebrush communities to fuel treatments. Rangeland Ecology and Management. 67: 

455–467.

Raphael, M.G.; Wisdom, M.J.; Rowland, M.M.; Holthausen, R.S.; Wales, B.C.; Marcot, 

B.G.; Rich, T.D. 2001. Status and trends of habitats of terrestrial vertebrates in relation to 

land management in the Interior Columbia River Basin. Forest Ecology and Management.

153: 63–88.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 47

Rau, B.M.; Blank, R.R.; Chambers, J.C.; Johnson, D.W. 2007. Prescribed fire in a Great 

Basin sagebrush ecosystem: dynamics of soil extractable nitrogen and phosphorus. 

Journal of Arid Environments. 71: 362–375.

Rau, B.M.; Chambers, J.C.; Blank, R.R.; Johnson, D.W. 2008. Prescribed fire, soil, and 

plants: Burn effects and interactions in the central Great Basin. Range Ecology and 

Management. 61: 169–181.

Rau, B.R.; Chambers, J.C.; Roundy, B.A.; Pyke, D.A.; Schupp, E.W.; Doescher, P.S.; 

Caldwell, T.G. 2014. Soil resources influence vegetation and response to fire and 

fire-surrogate treatments in sagebrush-steppe ecosystems. Rangeland Ecology and 

Management. 67: 506–521.

Rehfeldt, G.E.; Crookston, N.L.; Warwell, M.V.; Evans, J.S. 2006. Empirical analyses of 

plant-climate relationships for the western United States. International Journal of Plant 

Sciences. 167:1123-1150. 

Rich, T.D.; Wisdom, M.J.; Saab, V.A. 2005. Conservation of priority birds in sagebrush 

ecosystems. In Ralph, C.J.; Rich, T.D., eds. 2005. Bird conservation implementation and 

integration in the Americas: Proceedings of the Third International Partners in Flight 

Conference. Gen. Tech. Rep. PSW-GTR-191 (Volume 1). Albany, CA: U.S. Department 

of Agriculture, Forest Service, Pacific Southwest Research Station: 589–606.

Richardson, B.; Shaw, N.; Germino, M.J. 2014. Ecological genetics of big sagebrush 

(Artemisia tridentata): Genetic structure and climate-based seed zone mapping. 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: 18–24. http://www.treesearch.fs.fed.us/pubs/46564.

Richardson, B.A.; Kitchen, S.G.; Pendleton, R.L.; Pendleton, B.K.; Germino, M.J.; Rehfeldt, 

G.E.; Meyer, S.E. 2014. Adaptive responses reveal contemporary and future ecotypes in a 

desert shrub. Ecological Applications. 24: 413–427.

Richardson, B.A.; Ortiz, H.G.; Carlson, S.L.; Jaeger, D.M.; Shaw, N.L. 2015. Genetic and 

environmental effects on seed weight in subspecies of big sagebrush: applications for 

restoration. Ecosphere. 6(10): Article 201.

Richardson, B.A.; Page, J.T.; Bajgain, P.; Sanderson, S.C.; Udall, J.A. 2012b. Deep 

sequencing of amplicons reveals widespread intraspecific hybridization and multiple 

origins of polyploidy in big sagebrush (Artemisia tridentata; Asteraceae). American 

Journal of Botany. 99: 1962–1975.

Richardson, B.A.; Shaw, N.L.; Pendleton, R.L. 2012a. Chapter 4: Plant vulnerabilities and 

genetic adaptation. In: Finch, D.M., ed. Climate change in grasslands, shrublands, and 

deserts of the interior American West: A review and needs assessment. Gen. Tech. Rep. 

RMRS-GTR-285. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station: 48–59.

Roundy, B.A.; Miller, R.F.; Tausch, R.J.; Young, K.; Hulet, A.; Rau, B.; Jessop, B.;

Chambers, J.C.; Egget, D. 2014a. Understory cover responses to piñon and juniper 

removal treatments across tree cover gradients in the Great Basin. Rangeland Ecology and 

Management. 67: 482–494.
48 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Roundy, B.A.; Shaw, N.L.; Booth, D.T. 1997. Using native seeds on rangelands. In Shaw, 

N.L.; Roundy, B.A., comps. Proceedings: Using seeds of native species on rangelands. 

Gen. Tech. Rep. INT-GTR-372. Ogden, UT: U.S. Department of Agriculture, Forest 

Service, Intermountain Research Station: 1–8.

Roundy, B.A.; Young, K.; Cline, N.; Hulet, A.; Miller, R.F.; Tausch, R.J.; Chambers, J.C.;

Rau, B. 2014b. Piñon-juniper reduction effects on soil temperature and water availability 

of the resource growth pool. Rangeland Ecology and Management. 67: 495–505.

Roundy, Bruce A.; Hardegree, Stuart P.; Chambers, Jeane C.; Whittaker, Alison 2007. 

Prediction of cheatgrass field germination potential using wet thermal accumulation. 

Rangeland Ecology & Management: Vol. 60(6): 613-623.

Rowland, M.M. 2004. Effects of management practices on grassland birds: greater sagegrouse. Jamestown, ND: Northern Prairie Wildlife Research Center. Version August 12, 

2004. http://pubs.usgs.gov/unnumbered/70159591/report.pdf.

Rowland, M.M.; Vojta, C.D., tech. eds. 2013. A technical guide for monitoring wildlife 

habitat. Gen. Tech. Rep. WO-GTR-89. Washington, DC: U.S. Department of Agriculture, 

Forest Service. 400 p.

Rowland, M.M.; Wisdom, M.J. 2009. Chapter 19: Habitat networks for terrestrial wildlife: 

concepts and case studies. In Millspaugh, J.J.; Thompson, F.R., III, eds. Models for 

planning wildlife conservation in large landscapes. Cambridge, MA: Elsevier Science: 

501–531.

Rowland, M.M.; Wisdom, M.J.; Meinke, C.W.; Suring, L.H. 2005. Utility of greater 

sage-grouse as an umbrella species. In: Wisdom, M.J.; Rowland, M.M.; Suring, L.H., 

eds. Habitat threats in the sagebrush ecosystem: Methods of regional assessment and 

applications in the Great Basin. Lawrence, KS: Allen Press: 232–249.

Rowland, M.M.; Wisdom, M.J.; Suring, L.H.; Meinke, C.W. 2006. Greater sage-grouse as 

an umbrella species for sagebrush-associated vertebrates. Biological Conservation. 129: 

323–335.

Rowland, M.M.; Suring, L.H.; Tausch, R.J.; Geer, S.; Wisdom, M.J. 2010a. Dynamics of 

western juniper woodland expansion into sagebrush communities in central Oregon. 

Natural Resources and Environmental Issues. 16: article 13. 11 p.

Rowland, M.M.; Suring, L.H.; Wisdom, M.J. 2010b. Assessment of habitat threats to 

shrublands in the Great Basin: A case study. In Pye, J.M.; Rauscher, H.M.; Sands, Y.; 

Lee, D.C.; Beatty, J.S., tech. eds. Advances in threat assessment and their application to 

forest and rangeland management. Gen. Tech. Rep. PNW-GTR-802. Portland, OR: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest and Southern Research 

Stations: 673–685.

Rowland, M.M.; Suring, L.H.; Leu, M.; Knick, S.T.; Wisdom, M.J. 2011. Sagebrushassociated species of conservation concern. In: Hanser, S.E.; Leu, M.; Knick, S.T.; 

Aldridge, C.L., eds. Sagebrush ecosystem conservation and management: ecoregional 

assessment tools and models for the Wyoming Basins. Lawrence, KS: Allen Press: 46–68. 

Chapter 2.

Schlaepfer, D. R., Lauenroth, W. K. and Bradford, J. B. 2012. Ecohydrological niche of 

sagebrush ecosystems. Ecohydrology. 5: 453–466.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 49

Scholten, M.; Donahue, J.; Shaw, N.L.; Serpe, M.D. 2009. Environmental regulation 

of dormancy loss in Lomatium dissectum (Apiaceae) seeds. Annals of Botany. 103: 

1091–1101.

Schreiber, L. 2014. Greater sage-grouse nest site selection, brood-rearing site selection, and 

chick survival in Wyoming. Columbia, MO: University of Missouri. M.S. thesis. 112 p.

Schreiber, L.A.; Hansen, C.P.; Rumble, M.A.; Millspaugh, J.J.; Gamo, S.; Kehmeier, J.; 

Wojcik, N. 2015a. Microhabitat selection of brood-rearing sites by greater sage-grouse in 

Carbon County, Wyoming. Western North American Naturalist. 75: 348–363.

Schreiber, L.A.; Hansen, C.P.; Rumble, M.A.; Millspaugh, J.J.; Gamo, S.; Kehmeier, J.; 

Wojcik, N. 2015b. Greater sage-grouse apparent nest productivity and chick survival in 

Catron County, Wyoming. Wildlife Biology. doi 10.2981/wlb.00124.

Schroeder, M.A.; Young, J.R.; Braun, C.E. 1999. Sage-grouse (Centrocercus urophasianus). 

In: Poole, A.; Gill, F., eds. The birds of North America, no. 425. Philadelphia, PA: The 

Academy of Natural Sciences, and Washington, DC: The American Ornithologists’ 

Union.

Schroeder, M.A., Aldridge, C.L., Apa, A.D., Bohne, J.R., Braun, C.E., Bunnell, S.D., 

Connelly, J.W., Deibert, P.A., Gardner, S.C.,Hilliard, M.A., Kobriger, G.D., McCarthy, 

C.W., McCarthy, J.J.,Mitchell, D.L., Rickerson, E.V., Stiver, S.J., 2004. Distribution of 

sage-grouse in North America. Condor. 106: 363–373.

Schwartz, M.K. 2001. Estimating effects of landscape location on genetic variability in midsized carnivores. Missoula, MT: University of Montana. Ph.D. dissertation.

Schwartz, M.K.; Copeland, J.P.; Anderson, N.J.; Squires, J.R.; Inman, R.M.; McKelvey, 

K.S.; Pilgrim, K.L.; Waits, L.P.; Cushman, S.A. 2009. Wolverine gene flow across a 

narrow climatic niche. Ecology. 90: 3222–3232.

Shaw, N.; Jensen, S. 2014. Chapter 4.2: The challenge of using native plant materials for 

sagebrush steppe restoration in the Great Basin, USA. In Kiehl, K.; Kirmer, A.; Shaw, 

N.; Tischew, S., eds. Guidelines for native seed production and grassland restoration. 

Newcastle upon Tyne, UK: Cambridge Scholars: 141–159.

Shaw, N.; Pellant, M.; Fisk, M. Denney, E. 2012. A collaborative program to provide native 

plant materials for the Great Basin. Rangelands. 34: 11–16.

Shirk, A.J.; Schroeder, M.A.; Robb, L.A.; Cushman, S.A. 2015. Empirical validation 

of landscape resistance models: Insights from the greater sage-grouse (Centrocercus 

urophasianus). Landscape Ecology. 30: 1837–1850.

Shock, C.C.; Feibert, E.B.G.; Saunders, L.D. 2014. Irrigation requirements for native 

perennial wildflower seed production in a semi-arid environment. 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: 166–183. 

Oregon State University Agricultural Experimental Station, Corvallis, OR.

Shock, C.C.; Feibert, E.B.G.; Shaw, N.L.; Shock, M.P.; Saunders, L.D. 2015. Irrigation to 

enhance native seed production for Great Basin restoration. Natural Areas Journal. 35: 

74–82.

Shock, M.P.; Shock, C.C.; Feibert, E.G.B.; Shaw, N.L.; Saunders, L.D.; Sampangi, R.K. 

2012. Cultivation and irrigation of fernleaf biscuitroot (Lomatium dissectum) for seed 

production. HortScience. 47: 1525–1528.
50 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Smith, B.N.; Monaco, T.A.; Jones, C.; Holmes, R.A. 2002. Stress-induced metabolic 

differences between populations and subspecies of Artemisia tridentata (sagebrush) from 

a single hillside. Thermochimica Acta. 394: 205–210.

Smith, J.T.; Flake, L.D.; Higgins, K.F.; Kobriger, G.D.; Homer, C.G. 2005. Evaluating lek 

occupancy of greater sage-grouse in relation to landscape cultivation in the Dakotas. 

Western North American Naturalist. 65: 310–320.

Snodgrass, S.A.; Jensen, T.C. 2015. Four questions you may have about the sagegrouse “Not Warranted” Decision. The National Law Review: October 5, 2015. 

http://www.natlawreview.com/article/four-questions-you-may-have-about-sagegrouse-not-warranted-decision. [Accessed October 5, 2015].

Soberon, J.; Nakamura, M. 2009. Niches and distributional areas: concepts, methods, and 

assumptions. Proceedings of the National Academy of Science U.S.A. 106:19644–19650.

Soliai, M.M.; Meyer, S.E.; Udall, J.A.; Elzinga, D.E.; Hermansen, R.A.; Bodily, P.M.; Hart, 

A.A.; Coleman, C.E. 2014. De novo genome assembly of the fungal plant pathogen 

Pyrenophora semeniperda. PLoS ONE. 9(1): e87045.

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.

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.

Swanson, C.C.; Rumble, M.A.; Grovenburg, T.W.; Kaczor, N.W.; Klaver, R.W.; HermanBrunson, K.M.; Jenks, J.A.; Jensen, K.C. 2013. Greater sage-grouse winter habitat use on 

the eastern edge of their range. Journal of Wildlife Management. 77: 486–494.

Tausch, R.J.; Miller, R.F.; Roundy, B.A.; Chambers, J.C. 2009. Piñon and juniper field 

guide: Asking the right questions to select appropriate management actions. 

Circular 1335. Reston, VA: U.S. Department of the Interior, U.S. Geological Survey.

http://pubs.usgs.gov/circ/1335/.

Taylor, M.M.; Hild, A.L.; Shaw, N.L.; Norton, U.; Collier, T.R. 2014. Plant recruitment and 

soil microbial characteristics of rehabilitation seedings following wildfire in northern 

Utah. Restoration Ecology. 22: 598–607.

Thompson, T.W.; Roundy, B.A.; McArthur, E.D.; Jessop, B.D.; Waldron, B.; Davis, 

J.N. 2006. Fire rehabilitation using native and introduced species: A landscape trial. 

Rangeland Ecology and Management. 59: 237–248.

Turesson, G. 1925. The plant species in relation to habitat and climate. Hereditas. 6: 

147–236.

USDA Forest Service. 2012. National Forest System Land Management Planning. Federal 

Register. 77(68): 21162–21276.

USDA Forest Service. 2015. Fire Effects Information System. Missoula, MT: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula 

Fire Sciences Laboratory. http://www.feis-crs.org/feis/. [Accessed January 28, 2016].
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 51

U.S. Department of Energy. 2008. 20% Wind energy by 2030: Increasing wind energy’s 

contribution to U.S. electricity supply. DOE/GO-102008-2567. Washington, DC: 

U.S. Department of Energy, Office of Energy Efficiency and Renewable Energy.

http://www.nrel.gov/docs/fy08osti/41869.pdf. [Accessed December 13, 2014].

U.S. Department of the Interior, Bureau of Land Management, and U.S. Department of 

Agriculture, Forest Service [US Department of the Interior, BLM and USDA Forest 

Service]. 2014. The greater sage-grouse monitoring framework. Interagency Greater SageGrouse Disturbance and Monitoring Subteam. https://eplanning.blm.gov/epl-front-office/

projects/lup/21152/48421/52584/GRSG-FINAL-Monitoring_Framework_20140530.pdf . 

[Accessed January 31, 2016].

U.S. Department of the Interior, Fish and Wildlife Service [US Fish and Wildlife Service]. 

2014. Endangered and Threatened Wildlife and Plants; Threatened Status for Gunnison 

Sage-Grouse; Final Rule. 50 CFR 17. Federal Register. 79(224): 69192–69310.

U.S. Department of the Interior, Fish and Wildlife Service [US Fish and Wildlife Service]. 

2015a. Endangered and Threatened Wildlife and Plants; Withdrawal of the Proposed Rule 

To List the Bi-State Distinct Population Segment of Greater Sage-Grouse and Designate 

Critical Habitat. 80 FR 22827. 50 CFR 17. Federal Register. 78(208): 22827–22866.

U.S. Department of the Interior, Fish and Wildlife Service [US Fish and Wildlife Service]. 

2015b. Endangered and Threatened Wildlife and Plants; 12-Month Finding on a 

Petition to List Greater Sage-Grouse (Centrocercus urophasianus) as an Endangered or 

Threatened Species; Proposed Rule. 80 FR 59857. 50 CFR 17. Federal Register. 80(191): 

59857–59942.

Wang, H.; McArthur, E.D.; Sanderson, S.C.; Graham, J.H.; Freeman, D.C. 1997. Narrow 

hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae). 

IV. Reciprocal transplant experiments. Evolution. 51: 95–102.

Welch, B.L.; McArthur, E.D. 1981. Variation of monoterpenoid content among subspecies 

and accessions of Artemisia tridentata grown in a uniform garden. Journal of Range

Management. 34: 380–384.

Whittaker, A.; Roundy, B.; Chambers, J.; Meyer, S.; Blank, R.; Kitchen, S.; Korfmacher, J. 

2008. Effects of herbaceous species removal, fire and cheatgrass (Bromus tectorum) on 

soil water availability in sagebrush steppe. In: Kitchen, S.G.; Pendleton, R.L.; Monaco, 

T.A.; Vernon, J. comps. Proceedings―Shrublands under fire: Disturbance and recovery in 

a changing world. Proc. RMRS-P-52. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station: 49–56.

Williams, M.I.; Dumroese, R.K. 2013. Growing assisted migration: Synthesis of a climate 

change adaptation strategy. In: Haase, D.L.; Pinto, J.R.; Wilkinson, K.M., tech. coords. 

National proceedings: Forest and Conservation Nursery Associations - 2012. Proc. 

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

Mountain Research Station: 90–96.

Wisdom, M.J.; Chambers, J.C. 2009. A landscape approach for ecologically-based 

management of Great Basin shrublands. Restoration Ecology. 17: 740–749.
52 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

Wisdom, M.J.; Holthausen, R.S.; Wales, B.C.; Hargis, C.D.; Saab, V.A.; Lee, D.C.; Hann, 

W.J.; Rich, T.D.; Rowland, M.M.; Murphy, W.J.; Eames, M.A. 2000. Source habitats 

for terrestrial vertebrates of focus in the interior Columbia basin: Broad-scale trends and 

management implications. Vol. 1—Overview. Gen. Tech. Rep. PNW-GTR-485. Portland, 

OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 

176 p.

Wisdom, M.J.; Meinke, C.W.; Knick, S.T.; Schroeder, M.J. 2011. Factors associated with 

extirpation of sage-grouse. Studies in Avian Biology. 38: 451–472.

Wisdom, M.J.; Rowland, M.M.; Hemstrom, M.A.; Wales, B.C. 2005a. Landscape restoration 

for greater sage-grouse: Implications for multiscale planning and monitoring. In: Shaw, 

N.L.; Pellant, M.; Monsen, S.B., comps. Sage-grouse Habitat Restoration Symposium 

proceedings. Proc. RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station: 62–69.

Wisdom, M.J.; Rowland, M.M.; Suring, L.K. eds. 2005b. Habitat threats in the sagebrush 

ecosystem: Methods of regional assessment and applications in the Great Basin. 

Lawrence, KS: Allen Press. 301 p.

Wisdom, M.J.; Rowland, M.M.; Suring, L.H.; Schueck, L.; Meinke, C.W.; Knick, S.T.;

Wales, B.C. 2005c. Chapter 7: Habitats for groups of species. In Wisdom, M.J.; Rowland, 

M.M.; Suring, L.H., eds. Habitat threats in the sagebrush ecosystem: Methods of regional 

assessment and applications in the Great Basin. Part II: Regional assessment of habitats 

for species of conservation concern in the Great Basin. Lawrence, KS: Allen Press: 

205–231.

Wisdom, M.J.; Rowland, M.M.; Wales, B.C.; Hemstrom, M.A.; Hann, W.J.; Raphael, 

M.G.; Holthausen, R.S.; Gravenmier, R.A.; Rich, T.D. 2002a. Modeled effects of 

sagebrush-steppe restoration on greater sage-grouse in the Interior Columbia Basin, USA. 

Conservation Biology. 16: 1223–1231.

Wisdom, M.J.; Wales, B.C.; Rowland, M.M.; Raphael, M.G.; Holthausen, R.S.; Rich, 

T.D.; Saab, V.A. 2002b. Performance of Greater sage-grouse models for conservation 

assessment in the Interior Columbia Basin, USA. Conservation Biology. 16: 1232–1242.

Young, S.; Kitchen, S.; Armstrong, J.; Watson, V. 1995. AOSCA approves certification 

guidelines for wild land collected seed. Seed World. 133: 20–21.
USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 53

Appendix: Results of Sage-grouse/Sagebrush 

Questionnaire

Respondents

•  126 Forest Service (FS) employees completed the questionnaire

•  Most respondents were from middle management (GS9–GS13)

•  Senior managers (GS14–GS15) accounted for 12 percent of respondents

•  Respondents represented a wide array of research and management disciplines

•  Among disciplines, wildlife biologists were the dominant responders (40 percent)

•  About 75 percent of the respondents were from the National Forest System; 25 percent 

were from FS Research & Development

•  Nearly 30 percent of respondents work at the District level, versus 21 percent at the 

Supervisor’s Office level

Selected Key Findings by Subject Area

1. Ecosystem dynamics of sagebrush communities on National Forest System (NFS) lands

•  Strong agreement among respondents about the importance of understanding system 

dynamics of sagebrush communities, including effects of climate change, conifer 

encroachment, invasive species, and wildfire

•  Emphasis on the importance of better understanding of ecological relationships in 

high-elevation mountain big sagebrush communities

•  Disagreement about how much is known about effects of management, including livestock grazing and fire, on sagebrush maintenance and restoration

•  Strong disagreement among respondents that sagebrush will reestablish naturally under 

changing climate regimes

2. Contribution of sagebrush communities on NFS lands in supporting broader species/

system conservation

•  Strong agreement that more attention should be given to managing sagebrush plant and 

animal communities and identifying species found disproportionately on NFS lands

•  Strong agreement on the importance of understanding the effects of disturbance processes on sagebrush on NFS lands

3. Sage-grouse populations

•  Agreement on the need for knowledge about seasonal habitats of sage-grouse on NFS 

lands, effects of habitat management and restoration on sage-grouse, habitat connectivity for sage-grouse, and effects of energy development

•  Strong agreement about the need to better understand the contribution of sage-grouse 

that use NFS lands to larger populations of this species, contributions of seasonal 

habitats to overall species persistence, and landscape and community characteristics 

where sage-grouse populations are robust
54 USDA Forest Service, Gen. Tech. Rep. RMRS-GTR-348. 2016. 

•  Disagreement among respondents about whether sage-grouse are an appropriate 

indicator of sagebrush community condition, but general agreement that sage-grouse 

could be useful in assessments of other terrestrial species dependent on sagebrush 

habitats

•  Disagreement that seasonal habitats for sage-grouse are used disproportionately on 

NFS lands compared to those on other land ownerships

•  Strong disagreement among respondents that sage-grouse will adapt to rapid climate 

change

4. Habitat management and restoration

•  Strong agreement that new research is needed about effects of management on sagebrush systems, such as vegetation composition and structure, on NFS lands

•  Strong agreement that monitoring is important to ascertain effectiveness of management, and that disturbances should be monitored to understand recovery timeframes

•  Agreement on the need to synthesize research findings to provide management 

guidance

•  Strong agreement on the need for more information on restoration of sagebrush communities, especially at the periphery of current sage-grouse range, and in systems 

invaded by non-native species

•  Disagreement about our current understanding of effects of livestock grazing on sagegrouse habitats

5. Conservation guidance for sage-grouse

•  General (though not unanimous) agreement that the FS lacks sufficient conservation 

guidance for sage-grouse and sagebrush systems in current forest plans, and agreement 

that a synthesis of the literature would be useful in providing that guidance

•  A variety of opinions on the sufficiency of information to address threats and risks to 

sage-grouse habitats and whether sage-grouse are an appropriate surrogate for monitoring other species

To learn more about RMRS publications or search our online titles:

www.fs.fed.us/rm/publications

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