United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

Proceedings

RMRS-P-38

November 2005

Sage-Grouse Habitat

Restoration Symposium

Proceedings

June 4–7 2001, Boise, ID
Abstract _______________________________________________

Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sage-grouse habitat restoration symposium proceedings;

2001 June 4–7, Boise, ID. Proc. RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station. 130 p.

Declines in habitat of greater sage-grouse and Gunnison sage-grouse across the western United States are related to degradation,

loss, and fragmentation of sagebrush ecosystems resulting from development of agricultural lands, grazing practices, changes in

wildfire regimes, increased spread of invasive species, gas and oil development, and other human impacts. These losses are

focusing management efforts on passive and active approaches to maintaining and restoring sagebrush rangelands. This series of

14 papers summarizes current knowledge and research gaps in sagebrush taxonomy and ecology, seasonal sage-grouse habitat

requirements, approaches to community and landscape restoration, and currently available plant materials and revegetation

technology to provide a basis for designing and implementing effective management prescriptions.

Keywords: Centrocercus urophasianus, Artemisia tridentata, big sagebrush, native species, biodiversity, ecology, revegetation,

rehabilitation, shrub steppe

Acknowledgments _________________________________________________________________________________

We wish to thank the many individuals and organizations who participated in planning and conducting the

symposium, field tour, and equipment demonstration day. Mike Rath assisted in organizing and coordinating all

phases of the symposium and served as overall meeting moderator. John Kinney provided logistical support in

planning and conducting the symposium and field tours. Marcia Wicklow-Howard of the Boise State University

Biology Department served as liaison with the University. Antonia Hedrick and Kim Buxton of the USDI Bureau

of Land Management, Idaho State Office designed and maintained the meeting webpage, prepared symposium

brochure graphics, and provided mailing lists. Former Governor Cecil Andrus presented the plenary address for

the symposium. Alan Sands, Dean Mitchell, and John Connelly moderated the sessions; Barry Rose facilitated

the discussion on restoring sagebrush communities. Stacey Daniel of Meetings, Inc., managed registration with

assistance from Jan Gurr, Stephanie Carlson, and Danielle Scholten of the Rocky Mountain Research Station and

Jamie Jukar of the Fish and Wildlife Service. Taylor Cox, representing the Idaho Section of the Society for Range

Management served as treasurer. Dale Turnipseed; Kelly Memmott, Rocky Mountain Research Station; Joe

Russell, USDI Bureau of Land Management, Shoshone District; and Scott Walker, Utah Division of Wildlife

Resources assisted in organizing the equipment demonstration program. The Boise State University Student

Union staff provided facility support. Finally, we wish to thank Ann Debolt and the Rocky Mountain Research

Station Publication Services, in particular Louise Kingsbury and Loa Collins, for technical editing and preparation of this proceedings. The Great Basin Native Plant Selection and Increase Project provided funding for

reprinting this document.

Nancy L. Shaw

Stephen B. Monsen

Mike Pellant

Cover photo by Bill Mullins, Boise, ID.

Shrub illustrations found throughout this publication are from Range Plant Handbook. 1937. Washington, DC: U.S. Department of Agriculture, Forest Service.
Compilers

Nancy L. Shaw, Research Botanist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research

Station, Boise, ID.

Mike Pellant, Great Basin Restoration Initiative Coordinator, U.S. Department of the Interior, Bureau of Land

Management, Boise, ID.

Stephen B. Monsen (retired), Botanist, U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station, Shrub Sciences Laboratory, Provo, UT.

Sage-Grouse Habitat Restoration

Symposium Proceedings

June 4–7 2001, Boise, ID
Publisher’s note: Papers in this report were reviewed by the compilers. Rocky Mountain Research Station Publishing Services reviewed papers for format and

style. Authors are responsible for content.
Special Acknowledgment

Organization of this symposium, like many before it, was guided by the insight and enthusiasm of our colleague,

Steve Monsen, who retired in 2002. The workshops, training sessions, and symposia he conducted during his

career provided hundreds of scientists and managers with opportunities to interact and discuss local and regional

management concerns, recent research results, and, most importantly, to generate new ideas. Although he has

not yet learned to slow down and take it easy, we wish him well in his retirement.

Nancy Shaw

Mike Pellant

Sponsors

USDI Bureau of Land Management

USDA Forest Service, Rocky Mountain Research Station

Boise State University

Idaho Department of Fish and Game

Society for Range Management, Idaho Section

Colorado Division of Wildlife

The Nature Conservancy

Nevada Division of Wildlife

North American Grouse Partnership

Oregon Department of Fish and Wildlife

Society for Ecological Restoration, Northwest Chapter

USDA Forest Service, Regions 1, 2, 4, 5 and 6

USDI Fish and Wildlife Service, Snake River Basin Office

Utah Division of Wildlife Resources

Line drawing of sage-grouse was provided by USDI-BLM Idaho State Office.
Breaks

DuPont Vegetation Management

Granite Seed Company

Pawnee Buttes Seed, Inc.

Rainier Seed, Inc.

Stevenson Intermountain Seed Company, Inc.

Line drawing of sagebrush was provided by USDI-BLM Idaho State Office.
Contents

Page

Nancy L. Shaw Sage-Grouse Habitat Restoration Symposium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Mike Pellant

Stephen B. Monsen

Roger Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse . . . . . . 3

Sherel Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western

North America With Emphasis on Sage-Grouse Habitat . . . . . . . . . . . . . . . . . . . . . 17

Clait E. Braun Seasonal Habitat Requirements for Sage-Grouse: Spring, Summer,

John W. Connelly Fall, and Winter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

Michael A. Schroeder

Bruce A. Roundy Plant Succession and Approaches to Community Restoration. . . . . . . . . . . . . . . . . 43

Thomas A. Jones Status and Use of Important Native Grasses Adapted to

Steven R. Larson Sagebrush Communities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Scott C. Walker Current and Potential Use of Broadleaf Herbs for Reestablishing

Nancy L. Shaw Native Communities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

Michael J. Wisdom Landscape Restoration for Greater Sage-Grouse: Implications for

Mary M. Rowland Multiscale Planning and Monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

Miles A. Hemstrom

Barbara C. Wales

John A. Fairchild Big Sagebrush Response to One-Way and Two-Way Chaining in

James N. Davis Southeastern Utah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

Jack D. Brotherson

Scott M. Lambert Seeding Considerations in Restoring Big Sagebrush Habitat. . . . . . . . . . . . . . . . . . 75

Mike Pellant Strategies to Enhance Plant Structure and Diversity in Crested

Cindy R. Lysne Wheatgrass Seedings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

Cindy R. Lysne Restoring Wyoming Big Sagebrush . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

Nancy L. Shaw Reseeding Big Sagebrush: Techniques and Issues . . . . . . . . . . . . . . . . . . . . . . . . . 99

Ann M. DeBolt

Roger Rosentreter

Carl L. Wambolt Sagebrush-Ungulate Relationships on the Northern

Yellowstone Winter Range . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

Nancy Hampton Insects of the Idaho National Laboratory: A Compilation and Review . . . . . . . . . . 116

USDA Forest Service Proceedings RMRS-P-38. 2005 1

Nancy L. Shaw is Research Botanist at the U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, ID 83702.

Mike Pellant is Great Basin Restoration Initiative Coordinator, U.S. Department of the Interior, Bureau of Land Management, Nevada State Office,

Boise, ID 83709. Stephen B. Monsen is Botanist (retired) at the U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station,

Provo, UT 84606.

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Sage-Grouse Habitat Restoration

Symposium

Nancy L. Shaw

Mike Pellant

Stephen B. Monsen

Sage-grouse (greater sage-grouse [Centrocercus urophasianus] and Gunnison sage-grouse [C. minimus]) were once

abundant over a range that approximated that of sagebrush

(Artemisia spp.) in 16 Western States and three Canadian

Provinces (Aldrich 1963; Connelly and others 2000; Johnsgard

1973). Although their specific requirements vary seasonally

and over their life cycle, sage-grouse are almost completely

reliant upon sagebrush habitats (Connelly and others 2000;

Crawford and others 2004). Some populations are migratory

and require ranges exceeding 1,300 km2 (Wambolt and

others 2002).

Sage-grouse are now among the 338 or more species whose

populations are considered at risk for persistence (Wisdom

and others 2003) and are dependent on sagebrush ecosystems. Connelly and Braun (1997) and Braun (1998) estimated that since European settlement, the distribution of

sage-grouse has been reduced by 50 percent, and breeding

populations have declined by 17 to 45 percent since 1985.

Four petitions for subspecies or populations and three rangewide petitions have been filed to list the greater sage-grouse

under the U.S. Endangered Species Act (Kritz 2004). The

Gunnison sage-grouse is currently listed as a candidate

species (U.S. Fish and Wildlife Service 2000).

Sagebrush communities earlier occupied about 63 million

ha in Western North America (West 1983; West and Young

2000). Degradation, loss, and fragmentation of sagebrush

habitat has occurred as a result of excessive livestock grazing, conversion to agricultural lands or seedings of introduced grasses, spread of invasive exotic plants and native

conifers, alterations of fire regimes, oil and gas development,

and other human-caused disturbances (Crawford and others 2004; Hann and others 1997; Knick 1999; Knick and

others 2003; Noss and others 1995). Many areas have been

degraded beyond the threshold where recovery is likely to

occur naturally (Laycock 1991; West and Young 2000). As a

result, some sagebrush ecosystems are among the most

imperiled in North America (Noss and Peters 1995; Noss and

others 1995). Conserving and protecting extant portions of

sagebrush communities, altering management to encourage

passive restoration of at-risk areas, and actively restoring

degraded lands incapable of recovering without intervention presents a major challenge for Western land managers.

This symposium was organized to provide an overview of

science and technology addressing this issue.

Invited papers discussed sagebrush systematics, communities, ecology, and distribution (Goodrich, this proceedings; Rosentreter; this proceedings). Habitat requirements

and movements of sage-grouse were described to indicate

specific seasonal requirements and to demonstrate the need

for planning restoration at the landscape level (Braun and

Connelly, this proceedings; Wisdom and others, this proceedings). Other papers examined the principles of ecological restoration (Roundy, this proceedings) and native plant

materials available for use on degraded sagebrush rangelands (Jones and Larson, this proceedings; Walker and

Shaw, this proceedings). Additional papers described techniques for reestablishing sagebrush and understory species

and managing woody vegetation (Shaw and others; Fairchild

and others; Lambert; Lysne; Pellant; all in this proceedings).

Sixteen posters added depth to the range of topics discussed during the meeting. Two of these, included here,

address wildlife-sagebrush relationships (Hampton, this

proceedings; Wambolt, this proceedings). A 2-day field tour

focused on successes and failures of local revegetation efforts

and concluded with a demonstration of restoration equipment. Although the challenge of restoring millions of acres

of sage-grouse habitat is formidable, the science and practical approaches presented during the symposium provided

attendees with an overview of the status of the sagebrush

ecosystem, sage-grouse habitat requirements, and the potential for restoring those habitats.

References _____________________

Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae.

Journal of Wildlife Management. 27: 529–545.

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. 67: 134-144.

Connelly, J. W.; Braun, C. E. 1997. Long-term changes in sagegrouse Centrocercus urophasianus populations in western North

America. Wildlife Biology. 3: 229-234.

Connelly, J. W.; Schroeder, M. A.; Sands, A. R.; Braun, C. E. 2000.

Guidelines for management of sage grouse populations and

habitats. Wildlife Society Bulletin. 28: 967-985.

Crawford, J. A.; Olson, R. A.; West, N. E.; Mosley, J. C.; [and others].

2004. Ecology and management of sage-grouse and sage-grouse

habitat. Journal of Range Management. 57: 2-19.
2 USDA Forest Service Proceedings RMRS-P-38. 2005

Shaw, Monsen, and Pellant Sage-Grouse Habitat Restoration Symposium

Hann, W. J.; Jones, J. L.; Karl, M. G.; Hessburg, P. F.; [and others].

1997. An assessment of ecosystem components in the Interior

Columbia Basin and portions of the Klamath and Great Basins.

Vol. II. Landscape dynamics of the basin. Gen. Tech. Rep. PNWGTR-405. Portland, OR: U.S. Department of Agriculture, Forest

Service, Pacific Northwest Research Station: 337-1055.

Johnsgard, P. A. 1973. Grouse and quails of North America. Lincoln:

University of Nebraska Press. 551 p.

Knick, S. T. 1999. Forum: requiem for a sagebrush ecosystem?

Northwest Science. 73: 47-51.

Knick, S. T.; Dobkin, D. S.; Rotenberry, J. T.; Schroeder, M. A.; [and

others]. 2003. Teetering on the edge or too late? Conservation and

research issues for avifauna of sagebrush habitats. The Condor.

105: 611-634.

Kritz, K. 2004. Summary of sage grouse petitions submitted to the

U.S. Fish and Wildlife Service (as of January 8, 2004) [Online].

Available: http://nevada.fws.gov/public/petitionsummaryJan04.

pdf [2004, April 5].

Laycock, W. A. 1991. Stable states and thresholds of range condition

on North American rangelands: a viewpoint. Journal of Range

Management. 44: 427-433.

Noss, R. F.; Peters, R. L. 1995. Endangered ecosystems. A status

report on America’s vanishing habitat and wildlife. Washington,

DC: Defenders of Wildlife. 132 p.

Noss, R. F.; LaRoe, E. T., III; Scott, J. M. 1995. Endangered ecosystems of the United States: a preliminary assessment of loss and

degradation. Biol. Rep. 28. Washington, DC: U.S. Department of

the Interior, National Biological Service. 58 p.

U.S. Fish and Wildlife Service. 2000. Endangered and threatened

wildlife and plants; notice of designation of Gunnison sage grouse

as a candidate species. Federal Register. 65: 82310-82312.

Wambolt, C. L.; Harp, A. J.; Welch, B. L.; Shaw, N.; [and others].

2002. Conservation of greater sage-grouse on public lands in the

Western U.S.: implications of recovery and management policies.

PACWPL Paper SG-02-02. Caldwell, ID: Policy Analysis Center

for Western Public Lands. 41 p.

West, N. E. 1983. Western intermountain sagebrush steppe. In:

West, N. E., ed. Temperate deserts and semi-deserts. Vol. 5.

Ecosystems of the world. Amsterdam: Elsevier: 351-374.

West, N. E.; Young, J. A. 2000. Intermountain valleys and lower

mountain slopes. In: Barbour, M. G.; Billings, W. D., eds. North

American terrestrial vegetation. 2nd ed. Cambridge: Cambridge

University Press: 255-284.

Wisdom, M. J.; Rowland, M. M.; Suring, L. H.; Schueck, L.; [and

others]. 2003. Procedures for regional assessment of habitats for

species of conservation concern in the sagebrush ecosystem.

March 2003 Report, Version 1. Pacific Northwest Research Station, La Grande, OR, [Online]. Available: http://sagemap.wr.usgs.gov/

Docs/SageHab_Assess_Procedures_March03.pdf

Yampa
USDA Forest Service Proceedings RMRS-P-38. 2005 3

Roger Rosentreter is State Botanist, USDI Bureau of Land Management,

1387 S. Vinnell Way, Boise, ID 83709, e-mail: Roger_Rosentreter@blm.gov

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Sagebrush Identification,

Ecology, and Palatability Relative

to Sage-Grouse

Roger Rosentreter

Abstract—Basic identification keys and comparison tables for 23

low and big sagebrush (Artemisia) taxa are presented. Differences

in sagebrush ecology, soil temperature regimes, geographic range,

palatability, mineralogy, and chemistry are discussed. Coumarin, a

chemical produced in the glands of some Artemisia species, causes

UV-light fluorescence of the leaves. Coumarin-containing taxa,

such as mountain, xeric, subalpine big, subalpine early, black, and

low sagebrush, each fluoresce a bright bluish-white color. These

taxa are also the most palatable. A table of UV-light fluorescence of

20 sagebrush taxa in water solution is provided. How plant chemicals, such as coumarin and methacrolein and their seasonal variation, relate to palatability and animal preference is discussed in

terms of sage-grouse. Restoration guidelines for some sagebrush

taxa are also presented.

Keywords: Sagebrush, Artemisia, sage-grouse, palatability, preference, UV-light fluorescence

Introduction ____________________

The woody sagebrushes (Artemisia) are a major food

source of and provide critical habitat for the declining sagegrouse (Centrocercus urophasiamus), icon of Western rangelands (Braun and others1977; Connelly and others 2000;

Drut and others 1994). Improved identification of the types

of sagebrush this species eats and uses for nesting and cover

will help in its management. To the biologist and general

public who are unfamiliar with the many different species

and subspecies of sagebrush, this ecosystem may appear to

be a bewildering array of variability. However, sagebrush

communities are actually repetitive and easily identifiable

(Beetle 1960; West 1988). Recognizing them is important

because they are indicators of a given local ecosystem composed of specific vegetation types, soil depth, climate, topography, and wildlife species. Each type of sagebrush has

different palatability and structural characteristics, which

influences its particular values for wildlife (Sheehy and

Winward 1981).

Woody sagebrush species have been of major interest and

concern to land managers. Additional taxonomic research on

western North America’s woody Artemisias is needed. Collections made in fall when sagebrush taxa flower and are

most distinctive are particularly valuable. This genus could

include more genetic and morphological groups than are

currently described. As more studies are conducted on the

taxonomy of Artemisia, many of the subspecies and varietylevel taxa will likely be raised to that of the species; new

subspecies and varieties can be expected as well. The sagebrushes have been successful, in large part, due to their

ability to exchange genetic material by hybridization and

introgression (Hanks and others 1973; McArthur and others

1988), thus maintaining genotypic variation with sufficient

plasticity to allow the development of ecotypes. This genetic

variability may have also helped minimize disease and

herbivory, which weaken and limit less genetically diverse

species.

Why bother determining sagebrush and other vegetation

to the species or even subspecies level? As former, and now

deceased, University of Montana Professor Mel Morris used

to say, “The better the plant is at indicating ecological

condition or palatability, the more one should learn to

identify that plant.” Winward and Tisdale (1977) state that

separation of big sagebrush into subspecies assists in the

recognition of (1) habitat types (fig. 1), (2) production potential, (3) chemical content, and (4) palatability preference.

When Nuttall described Artemisia tridentata in 1841, more

than 20 present-day taxa were included. This broad species

concept would not help us today in managing the 23 named

sagebrush taxa that comprise sage-grouse habitat.

Palatability is defined as “plant characteristics or conditions that stimulate a selective response by animals” (Heady

1964). Webster defines the word “palatable” as pleasing to

the taste. The term “preference” is reserved for selection by

the animal and is essentially behavioral. Relative preference or relative palatability is a proportional choice among

two or more foods. Items positively correlated with preference include (Heady 1964) (1) high protein content, (2) linolenic and butyric acids, (3) fat content, (4) sugar, and (5)

phosphate and potash. Food items negatively correlated

with preference include (1) high lignin content, (2) crude fiber,

(3) tannins, and (4) nitrates (Heady 1964). In general, sagebrush species and populations that are more palatable to

mule deer are also more palatable to sheep, cows, insects,

and sage-grouse (Kelsey and Shafizadeh 1978; Sheehy and

Winward 1981; Wambolt 2001; Wambolt and others 1991;

Welch and Davis 1984; Welch and others 1983).

It is well documented that some sagebrush species are

more palatable due to their chemical content (Morris and
4 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

Figure 1—Environmental distribution of woody Artemisia taxa and the related Tanacetum nuttallii

arranged by soil moisture, depth, texture, mineralogy, and soil temperature (modified from West 1988).

Artemisia.

bigelovii

shallow

lithic

A. nova

shallow

gravelly

calcic

A. pgymaea

gravelly

calcic

A. arbuscula

ssp. lahontan

shallow, xeric

soil

 moisture

A. arbuscula

shallow

claypan

gravelly

A. rigida

shallow

stony

A. tridentata

ssp. wyomingensis

shallow to

moderately deep

A. tridentata

ssp. xericensis

moderately

deep, loamy

clay over basalt

A. tripartita

moderately

deep-deep

A. tridentata

ssp.

tridentata

deep, well

drained

A. tridentata

ssp. vaseyana

moderately

 deep,

well drained

A. tridentata

ssp.

spiciformis

Big Sagebrushes deep

Dwarf

sagebrush

of

warmer sites

Dwarf

sagebrush

of

cooler sites

Tanacetum

nuttallii gravelly

shallow calci shallow calcicc

A. longiloba

shallow

claypan,

ephemerally

flooded

A. cana

deep, poor

drainage

A. arbuscula

shallow

claypan

A. papposa

shallow lithic

basalt

Soil temperature gradient

Cryic Frigid Thermic

Dry Moist Seasonally wet

A. tripartita

ssp. rupicola

moderately

deep, lithic

Soil moisture gradient
USDA Forest Service Proceedings RMRS-P-38. 2005 5

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter

others 1976; Sheehy and Winward 1981; Wambolt 2001;

Welch and others 1983). The difference in palatability is

based on plant chemistry and the amount of volatile chemicals present in sagebrush leaf glands (Kelsey and others

1984; Striby and others 1987). Glands vary seasonally in the

amount and concentration of chemicals they contain, with

concentrations highest in spring and lowest in winter

(Cedarleaf and others 1983; Kelsey and others 1984). This is

due to the semievergreen nature of sagebrush and the

presence of persistent leaves, produced in the spring, with

glands full of volatile chemicals to discourage herbivory. In

fall and early winter, gland cell walls and neck cells age and

break open. These volatile chemicals are the distinctive

fragrance one smells after a rain in the sagebrush desert.

After releasing these chemicals, the sagebrush leaf becomes

more digestible. This difference has been shown through “in

vitro” digestibility of sagebrush leaves and alfalfa with the

addition of sagebrush-specific volatile compounds (Striby

and others 1987; Wambolt and others 1991). So, while some

sagebrush species’ high crude protein content encourages

herbivory, others contain chemicals, such as volatiles,

methacrolein, acetone, and 1-8 cineole, that discourage

feeding (Kelsey and others 1982; Wambolt 1996; Wambolt

and others 1991).

The amount of methacrolein in the three common subspecies of big sagebrush is consistent with their order of food

preference (Wambolt and others 1991). It might be reasonable to assume that the chemicals that mule deer, sheep, and

insects avoid will also be avoided by sage-grouse. The seasonal change in volatile oils also supports the premise for

greater wildlife use of sagebrush foliage in the winter, as

compared with spring and summer. However, in spite of

what is known, palatability information for the different

Artemisia species and subspecies is incomplete and somewhat based on assumption. This information gap is in part

due to the difficulty of distinguishing common sagebrush

species, as well as a lack of awareness of less common

species that may have seasonal importance. I hope this

paper helps improve this situation and stimulates research

and discussion about the seasonal vegetative needs and

preferences of the declining sage-grouse.

Methods _______________________

This paper summarizes current literature and the author’s

personal field and laboratory knowledge of woody sagebrush. Unfortunately, most of the sagebrush identification

and ecological literature has been treated on a State-byState rather than regional basis (Beetle 1960; Beetle and

Johnson 1982; Morris and others 1976; Winward and Tisdale

1977). Broader treatments using detailed flower characteristics for species divisions have been developed (Hall and

Clements 1923; McArthur 1979; Ward 1953); however, because they rely on the presence of the tiny sagebrush flowers,

they are impractical for most of the calendar year, or for the

biologist with no dissecting scope or herbarium reference

material. Most plant characteristics referred to in this paper

are visible with the naked eye or a 10x hand lens during any

season. All woody shrub and subshrub sagebrush utilized by

sage-grouse for food and habitat are included. The geographic

scope includes the Great Basin sagebrush steppe and adjacent portions of the Great Plains and Colorado Plateau that

have currently or historically supported sage-grouse (Connelly

and others 2000). The 23 sagebrush species and subspecies

treated are listed in table 1, arranged by their common and

scientific names. The table includes one non-Artemisiataxon,

Tanacetum nuttallii (chicken sage), a low-growing woody

species that vegetatively resembles Artemisia and is utilized

by sage-grouse.

Most palatability information does not come from sagegrouse use observations, since they are difficult to raise in

captivity, but are based on observations of other wildlife

species and on digestibility experiments such as those by

Barnett and Crawford (1994), Kelsey and others (1982),

Schwartz and others (1980), Sheehy and Winward (1981),

Wambolt (2001), Wambolt and others (1991), and Yabann

and others (1987). Much of the sagebrush chemistry literature is reported in highly technical chemistry-oriented journals and is in need of synthesis and interpretation for sagegrouse biologists and managers. Palatability of sagebrush

and other plants depends on the individual animal or population of animals feeding on it. In addition to the chemical

Table 1—Twenty-three sagebrush taxa (species and subspecies) are

listed in the order they are discussed. Nomenclature follows

McArthur (1983), with additional, newly described subspecies

following Goodrich and others (1985), Rosentreter and Kelsey

(1991), and Winward and McArthur (1995). The author

chose to exclude taxa that are either beyond the geographic

scope of this paper or that can be accounted for at a higher

taxonomic level.

Scientific name Common name

Dwarf sagebrush

Artemisia rigida Stiff sagebrush

A. spinescens Budsage

A. papposa Fuzzy sage

A. tripartita ssp. rupicola Wyoming threetip sagebrush

A. bigelovii Bigelow sagebrush

A. pygmaea Pygmy sagebrush

Tanacetum nuttallii Chicken sage

Artemisia longiloba Early sagebrush

A. arbuscula ssp. longicaulis Lahontan sagebrush

A. nova Black sagebrush

A. arbuscula Low sagebrush

Tall sagebrush

A. cana ssp. cana Plains silver sagebrush

A. cana ssp. bolanderi Bolander’s silver sagebrush

A. cana ssp. viscidula Mountain silver sagebrush

A. tripartita ssp. tripartita Threetip sagebrush

A. tridentata ssp. spiciformis Subalpine big sagebrush

A. tridentata ssp. vaseyana Mountain big sagebrush

A. tridentata ssp. vaseyana Few-flowered mountain

 var. pauciflora big sagebrush

A. tridentata ssp. wyomingensis Wyoming big sagebrush

A. tridentata ssp. tridentata Basin big sagebrush

A. tridentata ssp. xericensis Xeric big sagebrush

Subshrub sagebrush

Artemisia frigida Fringed sage

A. pedatifida Birdsfoot sage
6 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

content of food, learned behaviors may also dictate the food

choices animals make. Availability of the plant is also a

factor since hoofed animals may avoid, for example, a low

sagebrush site that is sloped and rocky, while sage-grouse

can readily use this type of terrain and the low sagebrush it

supports.

Results and Discussion __________

Taxonomy and the UV-Light Test

Several keys and comparison tables for field and lab identification of woody Artemisiaspecies are presented. The environmental distribution of these species is displayed by soil moisture, depth, texture, mineralogy, and soil temperature (fig. 1).

Field identification can be done year round; however, sagebrush specimens collected in the fall are much easier to identify

to species and subspecies. Ecological site knowledge and preferred soil mineralogy also help narrow down the possible taxa

that might occur at a given location (fig. 1).

It is easier to distinguish the different species and subspecies of sagebrush using both morphological and chemical

characteristics. Chemical analysis is a good tool to verify

field determinations and can help eliminate identification

problems due to morphological variation (Brunner 1972;

Scholl and others1977; Stevens and McArthur 1974). A

water extract of fresh or dried leaves of sagebrush can be

viewed under a long-wave ultraviolet (UV) light. Prior to

applying UV-light, several whole leaves are placed in a glass

vial with 10 ml or more of water and shaken. Leaves must be

from the same shrub rather than a composite sample since

one leaf with positive fluorescence will yield a false positive

result (Stevens and McArthur 1974). Table 2 contains the

UV-light response for each taxon.

Glass vials must be thoroughly cleaned between samples

to avoid contamination from previous tests. A voucher specimen of mountain big sagebrush should be the standard for

sample comparison. A positive test produces a bluish-white

fluorescence or glow, with the light held several inches from

the vial of leaf/water solution. Testing is best done in a dark

room or closet. Taxa cannot be distinguished solely by water

extract color differences, but the test is useful for taxa likely

to be confused based on morphology. This method can also be

applied to digested sagebrush from sage-grouse scats in the

field, using a portable UV-light and a dark-pigmented bag,

or the sample can be returned to the lab.

Palatability and the UV-Light Test

A positive test with blue fluorescence indicates the presence of coumarin, a chemical compound in certain sagebrush

species (Heywood and others 1977; McArthur and others

1988). These compounds, principally isocopoletin, scopoletin,

and esculentin, are water soluble and fluoresce under ultraviolet light. The higher the compound concentration in a plant,

the brighter the leaf/water fluorescence will be (Stevens and

McArthur 1974). Coumarin appears to correlate with increased palatability in most sagebrush taxa. Palatability

differences of individuals of the same taxa have even been

shown to correlate with UV-light fluorescence intensity

(Wambolt and others 1987, 1991; Welsh and others 1983).

Table 2—UV-light fluorescence of sagebrush taxa in water.

Fluorescence intensity is indicated as: (1) intense—very

bright bluish white that can be seen in a lighted room

indoors; (2) strong—bright bluish white that can serve

as a good standard for comparison in a dark location;

(3) moderate—bluish white in a dark location; (4) light—

very light blue and must be tested in complete darkness;

and (5) colorless—no fluorescence.

Bluish white Colorless

Early (intense) Basin big sagebrush

Subalpine big (intense) Wyoming big

Mountain big (strong) Bud

Xeric big (strong) Fuzzy

Low (strong) Stiff

Bigelow (moderate) Chicken sage

Lahontan (moderate) Black “type b”

Black “type a” (moderate)

Pygmy (moderate)

Silver (light)

Three-tip (light)

Wyoming three-tip (light)

Figure 2—Relative palatability gradient of sagebrush

for sage-grouse, from most to least palatable.

Palatability is defined as “plant characteristics or

conditions that stimulate a selective response by animals”

(Heady 1964). Webster’s New World Dictionary (Guralnik

1972) defines palatable as “pleasing to the taste.” The

term “preference” is reserved for selection by the animal

and is essentially behavioral. Relative preference or

relative palatability is a proportional choice among two

or more foods.

Species Palatability

Early

Low

Subalpine big

Mountain big

Wyoming big

Xeric big

Lahontan

Black “type a”

Mountain silver

Plains silver

Three tip

High

Chicken

Fringed

Bigelow

Bud

Pygmy

Bolander silver

Moderate

Black “type b”

Basin

Wyoming three-tip

Bird foot

Fuzzy

Stiff

Low
USDA Forest Service Proceedings RMRS-P-38. 2005 7

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter

Coumarin presence is a taxonomic indicator, separating

several of the sagebrush taxa (Kelsey and others 1982);

however, there are two exceptions to this. Wyoming big

sagebrush has little to no fluorescence, but is still highly

palatable. Bigelow sagebrush, which has a light-colored

fluorescence, contains volatile monoterpenes that discourage herbivory (fig. 2). Hybrids of taxa that brightly fluoresce

are intermediate in their response.

The UV-light test is an essential tool for sagebrush identification and palatability testing and should be used by

sagebrush botanists and sage-grouse wildlife biologists.

Palatability of sagebrush stands and individual plants can

be ranked based on their fluorescence intensity, without

even knowing the species.

Chemical Ecology

As previously mentioned, sagebrush chemicals are produced in glandular trichomes (Diettert 1938). These glands

cover 21 to 35 percent of both sides of a leaf ’s surface and are

hidden beneath a dense mat of hairs (Slone and Kelsey

1985). Glands contain coumarin as well as monoterpenes

and sesquiterpene lactones, all of which influence a plant’s

palatability. Presence of these volatile monoterpenes contributes to the characteristic smell of sagebrush. Because

these compounds are volatile, their concentration changes

seasonally, with lower concentrations in fall and winter.

The sesquiterpene lactones are the pasty, black material

found in sage-grouse scat, indicating that even sage-grouse

cannot digest these tar-like lactones. These chemicals are

probably deterrents to herbivory (Kelsey and others 1984;

Welch and others 1983). In laboratory experiments, a 10-

percent solution of lactones, extracted from big sagebrush

leaves and placed in potato dextrose agar (PDA), completely

inhibited growth of the common fungal mold, Alternaria sp.

A 5-percent solution of these lactones inhibited the growth

of Alternaria to as little as 25 to 61 percent of the control

(Rosentreter 1984).

Sagebrush Identification Guidelines

In order to identify sagebrush with a key, a few simple

rules must be followed. First, there are three types of leaves

on most sagebrush species (Miller and Shultz 1987; Winward

and Tisdale 1977). The “persistent” overwintering leaf is the

representative leaf shape and size used in the keys (Diettert

1938). The “ephemeral” leaf is generally larger and often

irregularly lobed. Ephemeral leaves are produced in spring

and shed in the summer when there is drought stress. These

odd-shaped leaves should be ignored, because they are fast

growing and atypical. The third leaf type is on the flowering

stalk. These leaves are often entire and lack the typical lobes

and shape of persistent leaves.

Comparison tables (tables 2, 4, and 5) and a dichotomous

key to all woody sagebrush species and subspecies are

provided. Leaf characteristics are based on overwintering

persistent leaves. Bell-shaped leaves have curved margins,

strap-shaped leaves have straight margins, and cleft-shaped

leaves are three parted. An “even crown” refers to flattopped shrubs with seedstalks originating at the same

height across a plant’s crown. A 10-power (10x) hand lens

can be used to examine leaf glands and hairs.

Individual Species Descriptions ____

Descriptions of each taxa are provided, including the

preferred mineralogy, palatability, ecology, distribution,

and management recommendations (figs. 1, 2). Dwarf sagebrush are discussed below as a group (also see table 3),

followed by tall sagebrush and subshrub taxa (see table 4).

Dwarf Sagebrush

A. Stiff Sagebrush (A. rigida)—Stiff sagebrush occurs

on very shallow skeletal basalt soils (Daubenmire 1982).

Stiff sagebrush has also been called scabland sage due to the

scabby, skeletal sites it prefers. Geographically, it grows in

the Pacific Northwest portion of the United States and

evades drought by being deciduous. Stiff sagebrush has

brittle or stiff branches and grows from 12 to16 inches tall.

Leaves are not reported to be palatable to any wildlife, but

sheep will eat the flowering stalks in late summer and fall

(Rosentreter 1992). Flower stalks are full of seeds that are

relatively high in protein. Stiff sagebrush has a large seed

(0.3 inch) that germinates quickly in 2 to 5 days (Rosentreter,

unpublished data). Sites are ephemerally saturated, and

contain a large diversity and cover of forbs when the sites are

not degraded (Rosentreter 1992; Rosentreter and McCune

1992). Sandberg bluegrass (Poa secunda) is the most common grass in these habitats due to the shallow soils. Stiff

sagebrush is not a resprouter as some authors have reported. It provides good spring and summer brood-rearing

habitat for sage-grouse. Suitable sites of stiff sagebrush

should be maintained and restored. The large seeds make

restoration of stiff sagebrush feasible and easier than many

other sagebrush species.

B. Budsage (A. spinescens)—Budsage grows on shallow, often saline soils at lower elevations, and is frequently

mixed with salt desert shrub vegetation. It flowers in the

spring (April to May). Budsage is geographically widespread, occurring from Montana to Arizona. It has palmately divided leaves that are deciduous. The leaves are

fragrant and smell different than the other species. Budsage

is considered to have low palatability, yet on degraded sites

it will be heavily used in the early spring by antelope, sheep,

and cattle. Budsage has a relatively large seed similar to stiff

and fuzzy sage, two other spring-flowering, deciduous species. Budsage has not been used in restoration projects, but

with its large seeds, it would appear to be feasible.

C. Fuzzy Sage (A. papposa)—Fuzzy sage occurs at mid

elevations (>5,000 ft) on shallow soils similar to low sagebrush sites (Rosentreter 1992). However, fuzzy sage is always on basalt bedrock, often with very shallow to almost no

soil over the skeletal basalt. Fuzzy sage is generally found on

large, flat basalt tables that ephemerally flood at the landscape level. It occurs in Idaho and Oregon (Rosentreter

1992). It is deciduous and has relatively large red or yellow

flowers in late spring. By late summer, plants are dried up

and domestic sheep, horses, and many wildlife species will
8 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

Key to the Woody Sagebrush of the Great Basin and Adjacent Areas

1. Short or tall shrubs with woody twigs..................................................................................................................................... 2

1. Short subshrubs with nonwoody twigs............................................................................................................................. Key A

2. Dwarf shrubs, mature plants generally <24 inches tall ......................................................................................... Key B

2. Tall to medium-sized shrubs, mature plants generally >24 inches tall ................................................................. Key C

Key A. Subshrubs with nonwoody twigs, woody at the base only

1. Leaf surface silvery, canescent .............................................................................................. Fringed sage, Artemisia frigida

1. Leaf greenish gray, pubescent ................................................................................................................................................. 2

2. Old flowering branches reduced to long spines, leaves dehiscent after spring, occurs at low elevations (also keyed

as a dwarf shrub in Key B)

............................................................................................................................................................ Budsage, A. spinescens

2. Plants without spines, leaves persistent, occurs at higher elevations in

Wyoming and Montana ............................................................................................................ Birdsfoot sage, A. pedatifida

Key B. Dwarf shrubs generally <24 inches tall

1. Plants deciduous, losing all their leaves in winter .................................................................................................................. 2

1. Plants semievergreen, retaining some leaves through winter................................................................................................ 4

2. Leaves three lobed and linear .................................................................................................. Stiff sagebrush, A. rigida

2. Leaves multilobed ............................................................................................................................................................. 3

3. Woody stems spiny, leaves light green ................................................................................................. Budsage, A. spinescens

3. Woody stems lacking spines, leaves palmately lobed, gray green and fuzzy

with many hairs on the surface ....................................................................................................... Fuzzy sage, A. papposa

4. Persistent leaves deeply cleft up to 1.5 inches, grows on

shallow soils on ridges at high elevations (7,500 to 9,000 ft)

......................................................................................................Wyoming threetip sagebrush, A. tripartita ssp. rupicola

4. Persistent leaves shallow lobed ........................................................................................................................................ 5

5. Pointed lobe tips, shallow lobes, and sharply three-toothed leaves

.............................................................................................................................................. Bigelow sagebrush, A. bigelovii

5. Rounded lobe tips ...................................................................................................................................................................... 6

6. Persistent leaves multilobed (>3 lobes), restricted to calcareous gravelly

soil in Utah ........................................................................................................................... Pygmy sagebrush, A. pygmaea

6. Persistent leaves three lobed ............................................................................................................................................ 7

7. Mature plants <4 inches tall, large flowered, growing only on windswept calcareous gravel ridges in Idaho and Montana

Chicken sage, Tanacetum nuttallii

7. Mature plants >4 inches tall, smaller flowers, growing on various habitats ......................................................................... 8

8. Early-maturing plants, flower in early summer; seed set by late August,

often layering, leaves broadly cuneate with relatively well-developed lobes,

large seeds (4 times the size of low sagebrush seeds) .......................................................... Early sagebrush, A. longiloba

8. Late-maturing plants, flower in late summer or fall with seed set in

October or November ............................................................................................................................................................ 9

9. Plant crown flat topped (even), flower stalks long and prominent, mostly above the plant, middle leaf lobe barely fits between the outside lobes

.............................................................................................................. Lahontan sagebrush, A. arbuscula ssp. longicaulis

9. Plant crown irregular not flat topped (uneven), flower stalks shorter and irregularly oriented, scattered throughout the

crown, leaves smaller with lobes of similar size ................................................................................................................ 10

10. Leaves with numerous glands on the surface best visible at 10x, leaves sticky to touch, when mashed between fingers, yields a glossy green to a black color, old flower stalks brown and persistent, flower heads with 3–5 florets

........................................................................................................................................................ Black sagebrush, A. nova

10. Leaves lack glands visible at 10x, gray green to silver-colored leaves, old flower stalks tan and nonpersistent,

flower heads with 5–11 florets

.................................................................................................................................................. Low sagebrush, A. arbuscula

Key C. Tall to medium sized shrubs (generally >24 inches tall)

1. Persistent leaves linear, layering .....................................................................................................Silver sagebrush, A. cana

1. Persistent leaves three lobed, layering absent or rare ........................................................................................................... 2

2. Persistent leaves deeply cleft

............................................................................................................ Three-tipped sagebrush, A. tripartita ssp. tripartita

2. Persistent leaves shallowly lobed ..................................................................................................................................... 3

3. Leaves large, 1.5 to 2.5 inches long and up to 0.75 inch wide, leaves dark green, broadly cuneate, often layering, only

found at high elevations

............................................................................................................. Subalpine sagebrush, A. tridentata ssp. spiciformis
USDA Forest Service Proceedings RMRS-P-38. 2005 9

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter

eat the mature flower stalks (Rosentreter 1992). Its leaves

are generally not palatable and are avoided except by horses

(personal observation by the author in Idaho and Oregon).

D. Wyoming Threetip Sagebrush (A. tripartita ssp.

rupicola)—Wyoming threetip sagebrush is a dwarf shrub

rarely more than 7 inches tall, with relatively long (1–1.2

inch), deeply cleft leaves with narrow (1-mm wide) linear

lobes. It occurs only in cold sites at high elevations greater

than 7,200 ft, east of the Continental Divide in Wyoming and

Montana. It is chemically similar to tall threetip sagebrush

(A. tripartita ssp. tripartita) and is not very palatable. It

will resprout weakly following physical disturbance or fire,

unlike the tall and more common threetip sagebrush that

readily resprouts. Because of their high elevation, most

Wyoming threetip sagebrush sites are not heavily impacted

by livestock, but the shallow rocky soils along ridgelines can

be impacted by off-highway trail proliferation.

E. Bigelow Sagebrush (A. bigelovii)—Bigelow sagebrush can be confused with both low and Wyoming big

sagebrush; however, Bigelow sagebrush leaves are more

shallowly lobed and sharply pointed. The pointed leaf tips

make identification of this species easy, as long as biologists

and managers are aware of its potential presence. It occurs

on arid and mesic calcareous soils and on highly decomposed

granite. It grows throughout the Southwest from California

to west Texas and north to northwest Colorado. Bigelow

sagebrush is one of the sagebrush taxa that fluoresces, but

is not considered highly palatable (silver sagebrush is another). Increased awareness of Bigelow sagebrush by the

wildlife community, particularly in Colorado, may provide

additional information on its palatability in the future.

F. Pygmy Sagebrush (A. pygmaea)—Pygmy sagebrush

grows on dry alluvial fans at elevations from 5,000-7,000 ft.

It occurs in shallow soils less than 20 inches deep with a

gravelly soil surface texture and is often mixed with black

sagebrush. It is found only in Utah, Nevada, and northern

Arizona. Pygmy sagebrush has moderate palatability and

may be utilized by wildlife in the winter, due to its availability at moderately low-elevation sites.

G. Chicken Sage (Tanacetum nuttallii)—Taxonomically, chicken sage has been treated as either Tanacetum

nuttallii or Sphaeromeria nuttalli. Both of these genera are

closely related to the genus Artemisia. Chicken sage grows

on windswept benches and large flat areas on very shallow,

calcareous gravels in Idaho, Montana, and Wyoming. Chicken

sage looks like a diminutive low sagebrush, but it has

smaller leaves and flowers in early summer. Flower heads

are relatively large with a small pappus forming around the

achene. Due to the presence of the small scalelike pappus, it

has not been classified as an Artemisia, despite its other

similar morphological characteristics. Chicken sage is woody

and has three-lobed leaves like many Artemisias. Palatability of this species is moderate, based on its use by antelope

(Brent 1976; Thomas and Rosentreter 1992). Sage-grouse

are found where this species is common, but it is unknown

if they utilize it for food. Its growth form is low spreading to

almost creeping, and its branches are less than 4 inches tall;

thus, it does not provide structural or hiding cover for sagegrouse. Brent (1976) recorded antelope spending large

amounts of time in windswept, normally snow-free chicken

sage sites. This suggests their availability for sage-grouse in

winter as well.

H. Early Sagebrush (A. longiloba)—Early sagebrush

grows on shallow, ephemerally flooded soils, often with a

claypan or skeletal rock layer near the surface (Robertson

and others 1966). It is frequently found in low-drainage

Key to the Woody Sagebrush of the Great Basin and Adjacent Areas (con.)

3. Leaves smaller, gray green, not layering ................................................................................................................................. 4

4. Plant crown flat topped (even), flower stalks long and prominent,

mostly above the plant, leaf margins tapered, leaves widest just below the

lobes, leaves in water fluoresce bright bluish white under UV light

........................................................................................................... Mountain big sagebrush, A. tridentata ssp. vaseyana

A. Persistent leaves widest at base of lobes. Inflorescence a spike or raceme with relatively few heads, plants occasionally layering ....................................................................................................................................... A. t. var. vaseyana

B. Persistent leaves widest slightly below the base of the lobes.

Inflorescence a panicle with numerous heads, plants do not layer

.................................................................................................................................................................. A. t. var. pauciflora

4. Plant crown irregular (uneven), not flat topped, flower stalks smaller

and irregularly oriented, scattered throughout the crown ................................................................................................. 5

5. Mature shrubs short, <3 ft tall, leaves bell shaped, grows at lower elevations, plant is U shaped Wyoming big sagebrush,

A. tridentata ssp. wyomingensis

5. Mature shrubs taller, generally >3 ft tall, leaves not bell-shaped, grows at

low to high elevations, plant is Yshaped with a central stalk, rather than

U shaped or with cupped growth form of many other species ............................................................................................ 6

6. Leaf margins straight, leaves long and strap shaped, leaves in water

do not fluoresce, prefers deep well-drained soils, widespread geographically and ecologically

................................................................................................................ Basin big sagebrush, A. tridentata ssp. tridentata

6. Leaf margins tapered, leaves widest just below the lobes, leaves fluoresce bright blue under UV light, grows only in

loamy clay soils in western Idaho ..........................................................Xeric big sagebrush, A. tridentata ssp. xericensis
10 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Shallow claypan, granitic or basalt; aridic-cryic Species or subspecies Crown Leaf shape and color Leaf margina Layering Preferred soil mineralogy and soil temperature Color of sage/water solution under UV lightc Flowering begins Low sagebrush Uneven-even Lahontan sagebrush Even, due to the long flower stalks No Strongly bluish white August Black sagebrush Uneven-even Early sagebrush Shallow claypan, often in ephemerally flooded but not always alkaline areas; frigid-cryic Even Yes Intense bluish white Plant architectureb August Moderately bluish white No Dwarf spreading, U shaped Slightly bell shaped Gray August June-July Shallow clay soil; aridic-mesic Calcareous; shallow, aridic-frigid No U shaped, often taller than the often low sages Dwarf, spreading, U shaped Dwarf, spreading, U shaped Elongated bell shape, ephemeral leaves often very large Bell shaped Broadly bell shaped with an obviously cleft central lobe Gray, larger leaf than low sage Dark green Dark gray green Moderately bluish white; some colorless populations exist Table 3—Morphological and chemical characteristics that distinguish the low sagebrushes. a Leaf margin refers to the overall shape of the leaf, the curvature of its margin, and how wide it is at its apex. b Plant architecture refers to a species’ basal branching pattern—multiple stems bestow an overall “U shape,” and one to few stems produce a wider appearance at the top or a “Y shape.” c Consider all taxonomic characteristics in combination when making a determination. A long-wave UV light should be used to test the sagebrush leaf/water solution in a dark location.
USDA Forest Service Proceedings RMRS-P-38. 2005 11

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter Deep, well drained; aridic-mesic Species or subspecies Crown Leaf shape Leaf margina Layering Preferred soil mineralogy and soil temperature Color of sage/water solution under UV lightc Flowering begins Basin big sagebrush Uneven Mountain big sagebrush Even No Strong bluish white July, (<6 flowers/ inflorescense) Wyoming big sagebrush Uneven Colorless Xeric big sagebrush Uneven Loamy clay over basalt, mesic-xeric Strong bluish white Subalpine big sagebrush Even Yes Deep, noncalcar- eous, often a clay layer; frigid Intense bluish white Plant architectureb Late August Colorless No Y shaped, erect Straight Long and narrow August Late August July, (>6 flowers/ inflorescense) Well drained, frigid-cryic Shallow to moderately deep soil, aridic No No U shaped, basal branching U shaped Y shaped, radiate Spreading, multiple stemmed Tapered Bell shaped Tapered Tapered to elongated bell shape Broadly cuneate Belled Broadly cuneate Large leaves Table 4—Morphological and chemical characteristics that distinguish the big sagebrushes. a Leaf margin refers to the overall shape of the leaf, the curvature of its margin, and how wide it is at its apex. b Plant architecture refers to a species’ basal branching pattern—multiple stems bestow an overall “U shape,” and one to few stems produce a wider appearance at the top or a “Y shape.” c Consider all taxonomic characteristics in combination when making a determination. A long-wave UV light should be used to test the sagebrush leaf/water solution in a dark location.
12 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

areas of flats, plateaus, or tables. Early sagebrush is a

prolific seed producer and could be used for restoration in

appropriate, shallow soil sites (Beetle and Johnson 1982;

Monsen and Shaw 1986). It layers and can resprout after

cool fires. Early sagebrush is one of the most valuable taxa

for sage-grouse, and many of the largest leks in Idaho are in

areas dominated by this species (Camas Prairie, south of

Fairfield, ID) (fig. 1) (Robertson and others 1966). It flowers

very early in the summer, in contrast to other low-statured

species. Early sagebrush has been confused with low-growing Wyoming big sagebrush because of its broadly cuneate

3-lobed leaves, and with low sage because of its dwarf size.

Early sagebrush is palatable to sheep and, historically,

stands were commonly used as lambing areas (Beetle and

Johnson 1982). These areas should be monitored to prevent

heavy spring grazing by domestic livestock. Early sagebrush has also been referred to as “alkali sagebrush,”

although sites may or may not be alkaline (Robertson and

others 1966).

I. Lahontan Sagebrush (A. arbuscula ssp. longicaulis)—Lahontan sagebrush is a type of low sagebrush that

grows on shallow clay soils formed on the shore of Pleistocene Lake Lahontan. It grows in northwest Nevada and

adjacent California and Oregon at elevations from 3,400-

6,600 ft. It differs from low sagebrush chemically and by its

longer floral stalks and larger leaves. Lahontan sagebrush

occurs on soils similar to low sagebrush, but in areas that

receive less precipitation (5-12 inches). It is moderately to

highly palatable (Winward and McArthur 1995).

J. Black Sagebrush (A. nova)—There appears to be at

least two chemical races of black sagebrush in the West

(Kelsey 2002, personal communication; McArthur and

Plummer 1978). One race with gray leaves is highly palatable, while the green-leafed race has low palatability (fig. 2)

(McArthur and Plummer 1978). This latter form does not

fluoresce under UV light. Additional studies are needed to

determine the geographic ranges and correlation with physical characteristics for these two races. Black sagebrush has

been greatly reduced or eliminated on some ranges where

sheep graze in winter (Clary 1986). The best feature to

identify this species is its flower stalks. The stiff, erect stalks

dry to brown and persist through the following year. Most

populations have leaf glands visible with a 10x hand lens

(Kelsey and Shafizadeh 1980). Black sagebrush grows well

on very shallow stony soils, often on windswept slopes and

ridges at mid- to high elevations where annual precipitation

is more than 10 inches (Behan and Welch 1985). It prefers

calcareous or well-decomposed granitic soils that seem to

mimic calcareous sites due to weathering of calcium feldspars. Black sagebrush is a widespread species, second only

in its geographical distribution to basin big sagebrush.

K. Low sagebrush (A. arbuscula)—Low sagebrush

grows on shallow soils with a restrictive layer of bedrock or

clay pan. This species is usually found where annual precipitation is greater than 12 inches. Parent material is

noncalcareous. Low sagebrush is one of the most palatable

sagebrushes for sage-grouse. It is a wide-ranging species,

found throughout the Great Basin. Black, early, Bigelow and

Lahontan sagebrush, and chicken sage are often misidentified

as low sagebrush.

Tall Sagebrush

L. Silver Sagebrush (A. cana)—Silver sagebrush is a

tall shrub with three subspecies that grow in distinctly

different habitats. All three subspecies are root-sprouters

and layer vegetatively. The three subspecies are distinguished as:

1. Mature plants 3 to 5 ft tall, leaves mostly >0.8 inch

long and strongly pubescent, a plant of arid riparian

drainages

.................. Plains silver sagebrush, A. cana ssp. cana

1. Mature plants <40 inches tall, leaves mostly <0.8

inch long...................................................................... 2

2. Leaves pubescent and silver gray, plant of playas

(internally drained basins)

.... Bolander silver sagebrush, A. cana ssp. bolanderi

2. Leaves sparsely pubescent and dark green, plant

of high elevations

.... Mountain silver sagebrush, A. cana ssp. viscidula

Mountain and plains silver sagebrush are considered

highly palatable (Wambolt 2001), while Bolander silver

sagebrush is only moderately palatable. The former two

species generally grow where they receive additional moisture from the surrounding vegetation. All three subspecies

are within the range of sage-grouse. Plains silver sagebrush

is often the only Artemisia used by grouse on the flat plains

of central and eastern Montana.

M. Threetip Sagebrush (A. tripartita ssp.

tripartita)—Threetip sagebrush is a fairly tall, erect shrub

(4 to 6 ft). It grows on deep, well-drained soils, often mixed

with basin or mountain big sagebrush. It will seldom layer

without disturbance, but will vigorously stump sprout and

layer after burning. It is considered highly palatable to

wildlife (Wambolt 2001); however, there is high seasonal

variation in its utilization. Livestock, including sheep, appear to avoid utilization of this species. Beware of control or

prescribed burning in threetip sagebrush habitat, as it can

increase well beyond the site’s preburn density. It is common

in Washington, Idaho, Montana, Wyoming, Utah, and Colorado.

N. Subalpine Big Sagebrush (A. tridentata ssp.

spiciformis)—Subalpine big sagebrush grows on deep, cryic

soils and is highly palatable. Sage-grouse reportedly use this

species; however, it probably becomes unavailable in late

winter due to snow cover. It frequently grows where large

snowdrifts form, unlike dwarf sagebrush types that grow in

windswept areas. Subalpine big sagebrush can occur on

ridgelines, similar to some of the dwarf and low-stature

sagebrushes. These ridgelines are frequently used by sagegrouse. Chemically, subalpine big sagebrush appears to be

a choice food for sage-grouse and other wildlife species. It

layers vegetatively and resprouts following defoliation from

heavy snow. It occurs in Idaho, Wyoming, Montana, Utah,

and Colorado (Goodrich and others 1985; McArthur and

Goodrich 1986).

O. Mountain Big Sagebrush (A. tridentata ssp.

vaseyana)—This includes varieties paucifloraand vaseyana.

Mountain big sagebrush is a flat-topped shrub that grows to

3 ft tall (Tisdale and Hironaka 1981). It has a U-shaped
USDA Forest Service Proceedings RMRS-P-38. 2005 13

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter

crown and is found on moderate to deep, well-drained, frigid

soils, generally above 5,000 ft. Mountain big sagebrush can

grow as low as 3,000 ft, and when it does, soils are typically

very well drained. It is highly palatable to most wildlife;

however, limited access in the winter and the chemical

content in spring and summer may discourage herbivory

(Kelsey and Shafizadeh 1978; Kelsey and others 1984).

Mountain big sagebrush is a major food source for sagegrouse in the winter months. Sage-grouse scats collected

from Wyoming big sagebrush-dominated sites in Idaho and

Colorado tested positive for UV light, indicating that grouse

were feeding nearby on UV positive sagebrush (mountain

big sagebrush or low sagebrush) (Vasquez 2002). In the

Gunnison Basin of Colorado, sage-grouse utilize a hybrid of

A. t. ssp. vaseyana and A. t. ssp. wyomingensis (Vasquez

2002).

Compared to other sagebrush taxa, mountain big sagebrush has a greater potential to increase its density due to

its general ecology and the higher moisture its habitat

receives. Stands can become so dense they are difficult for

humans to walk through. In much of the West, heavy

livestock use, both historic and current, has reduced forb,

perennial grass, and biological soil crust components, allowing sagebrush and exotic annual grasses to become dense

(Billings 1994; Rosentreter and Eldridge 2002). Mechanical

control, burning, or seeding followed by rest from grazing, is

necessary in many areas to restore the vegetative and

structural diversity needed for optimal wildlife habitat.

“Hobble Creek” mountain big sagebrush, a highly palatable

cultivar of A. t. ssp. vaseyana, is recommended for restoration projects with the goal of improving wildlife winter range

(Welch and others 1990).

P. Wyoming Big Sagebrush (A. tridentata ssp.

wyomingensis)—Wyoming big sagebrush is a medium sized

shrub from 1 to 3 ft tall. It branches from the base, giving it

a U-shaped architecture. Wyoming big sagebrush grows at

warmer, lower elevations and is more available as forage in

winter and early spring (Wambolt 1998). It occurs at sites

receiving from 8 to 12 inches of precipitation. This species is

generally palatable, though its palatability is highly variable. Many Wyoming big sagebrush sites have been severely

degraded and converted to exotic annual grasslands; thus, in

harsh winters, they are no longer available for sage-grouse

use (Hilty and others 2003).

Nondegraded, lightly grazed Wyoming big sagebrush sites

have a high percent cover of biological soil crusts and low

percentage of cheatgrass cover (Kaltenecker and others

1999; Rosentreter 1986; Rosentreter and Eldridge 2002).

Due to their susceptibility to invasion and domination by

cheatgrass and other exotic annuals, use of fire to manage

them must be approached with caution. Wyoming big sagebrush sites should be managed for retention of the biological

soil crust component. Late fall, winter, and early spring is

the most appropriate season of use for this low-elevation

vegetation type. Four to 6 weeks of moist soil conditions in

late spring facilitates regrowth of biological soil crusts disturbed by trampling (Memmott and others 1998; Rosentreter

and Eldridge 2002). “Gordon Creek” Wyoming big sagebrush,

a highly palatable cultivar, is recommended for restoration

projects aimed at improving wildlife winter range (Welch

and others 1992).

Q. Basin Big Sagebrush (A. tridentata ssp. tridentata)—Basin big sagebrush is the least palatable of the big

sagebrushes (Wambolt 1998), though it is chemically and

genetically (2n = 18 or 36) highly variable. It is considered of

low palatability relative to other sagebrush taxa (fig. 2), and

it is also the tallest. Its architecture is somewhat single

trunked (tree like) or Y shaped, with lateral branches diverging from the main stem at a different angle than those of

either Wyoming or mountain big sagebrush. This prolific

seed producer grows on deep, well-drained soil (Daubenmire

1975). The extra moisture runoff from roads can create

artificial sites for this subspecies, even in soils normally

occupied by Wyoming big sagebrush. Large areas dominated

by Wyoming big sagebrush will frequently have basin big

sagebrush adjacent to the road ditch. Seed of the more

prolific, larger statured basin big sagebrush is often harvested along with seed of Wyoming big sagebrush. Dalzell

(2004) found that 8 percent of shrubs in Wyoming big

sagebrush seedings in southern Idaho were basin big sagebrush. Basin big sagebrush leaves have rarely been identified in sage-grouse scats (Rosentreter 2001, unpublished

data; Vasquez 2002). However, even small dense stands of

this shrub can provide good nesting habitat for sage-grouse

in Colorado’s Gunnison Basin.

R. Xeric Big Sagebrush (Artemisia tridentata ssp.

xericensis)—Xeric big sagebrush is a tall shrub (>3 ft) with

Y-shaped architecture similar to basin big sagebrush. However, its chemistry, leaf shape, and palatability are most

similar to mountain big sagebrush. At lower elevations

(2,500–4,500 ft), this Idaho subspecies is restricted to heavy

clay-loam and drier, xeric soils than mountain big sagebrush. In Idaho, mountain big sagebrush grows between

4,000 and 9,500 ft, in moister Udic soils. Xeric big sagebrush

is heavily utilized in winter by mule deer and, based on its

chemistry (high crude protein) (Rosentreter and Kelsey

1991), is likely preferred by sage-grouse. It can increase in

density similar to mountain big sagebrush, with heavy

spring, summer, or early fall cattle grazing.

Subshrub Sagebrush

S. Fringed Sage (A. frigida)—Fringed sage is a small

subshrub, woody only at the base. It is the most widespread

species treated in this paper, extending into other North

American and Asian biomes, such as alpine meadows, the

Great Plains, and mountain meadows. It was described from

Siberia before being identified in North America. Fringed

sage occurs in a variety of soil types and depths, but prefers

shallow soils with frigid soil temperatures (Morris and

others 1976). Some sites are windswept and are readily

available to wildlife in the winter. Fringed sage is moderately palatable. In the Gunnison Basin, sage-grouse have

been observed eating fringed sage seedlings in early spring

(Young 2001).

T. Birdsfoot Sage (A. pedatifida)—Birdsfoot sage is a

small subshrub, weakly woody at the base. It occurs in dry
14 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

shallow soils at high elevations with frigid soil temperatures

in Wyoming, Montana, and Idaho. This subshrub is most

commonly found in montane grasslands and on windswept

sites. It is 1 to 5 inches tall with finely canescent, basal

leaves. Leaves are once or twice ternately divided into linear

divisions with finely white-tomentulose hairs. Flowers are

brownish. Birdsfoot sage has low palatability.

Conclusion_____________________

Coumarin-containing taxa such as mountain, xeric, subalpine big, subalpine early, black, and low sagebrush all

fluoresce a bright bluish-white color. These taxa are also the

most palatable. Plant chemicals such as coumarin and

methacrolein and their seasonal variation affect shrub palatability and animal, including sage-grouse, preference.

Because sagebrush species also differ vastly in their structural characteristics and habitat requirements, knowledge

of Artemisia ecology will enhance our ability to improve and

manage habitat for sage-grouse.

Acknowledgments ______________

I would like to thank the many people that have taught me

about sagebrush ecology and chemistry over the years. Field

trips with range scientists, many now deceased, such as Mel

Morris, Ed Tisdale, Chuck Wellner, and Doug Henderson,

were an important part of this education. I would also like to

thank Alma Winward and Durant McArthur, great educators and accomplished sagebrush ecologists and taxonomists. Soil scientist Al Harkness, natural product chemist

Rick G. Kelsey, and botanist Steve Monsen have also been

instrumental. My former supervisor and mentor, Alan Sands,

along with Vicki Saab, taught me how to examine bird scats

and bird crops, and to view the landscape from a grouse’s

perspective. Ann DeBolt provided extensive comments, and

reviewers Cindy Dalzell, Carl Wambolt, and Gay Austin also

significantly improved this paper.

References _____________________

Barnett, J. K.; Crawford, J. A. 1994. Pre-laying nutrition of sagegrouse hens in Oregon. Journal of Range Management. 47:

114–118.

Beetle, A. A. 1960. A study of sagebrush, the section Tridentatae of

Artemisia. Bull. 368. Laramie: University of Wyoming, Agricultural Experiment Station. 83 p.

Beetle, A. A.; Johnson, K. L. 1982. Sagebrush in Wyoming. Bull. 779.

Laramie: University of Wyoming, Agricultural Experimental

Station. 68 p.

Behan, B.; Welch, B. L. 1985. Black sagebrush: mule deer winter

preference and monoterpenoid content. Journal of Range Management. 38: 278–279.

Billings, W. D. 1994. Ecological impacts of cheatgrass and resultant

fire on ecosystems in the western Great Basin. In: Monsen, S. B.;

Kitchen, S. G., comps. Ecology and management of annual rangelands symposium: proceedings; 1992 May 18–22; Boise, ID. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 22–30.

Braun, C. E.; Britt, T.; Wallestad, R. O. 1977. Guidelines for

maintenance of sage-grouse habitats. Wildlife Society Bulletin. 5:

99–106.

Brent, W. C. 1976. Observations on a pronghorn antelope winter

range. Pocatello, ID: Idaho State University. Thesis.

Brunner, J. R. 1972. Observations on Artemisia in Nevada. Journal

of Range Management. 25: 205–208.

Cedarleaf, J. D.; Welch, B. L.; Brotherson, J. D. 1983. Seasonal

variation of monoterpenoids in big sagebrush (Artemisia

tridentata). Journal of Range Management. 36: 492–494.

Clary, W. P. 1986. Black sagebrush response to grazing in the eastcentral Great Basin. In: McArthur, E. D.; Welch, B. L., comps.

Proceedings—symposium on the biology of Artemisia and

Chrysothamnus; 1984 July 9–13; Provo, UT. Gen Tech. Rep. INT200. Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 181–185.

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.

Dalzell, C. R. 2004. Post-fire establishment of vegetation communities following reseeding on southern Idaho’s Snake River Plain.

Boise, ID: Boise State University. 112 p. Thesis.

Daubenmire, R. F. 1975. Ecology of Artemisia tridentata subsp.

tridentata in the State of Washington. Northwest Science. 49:

24–34.

Daubenmire, R. F. 1982. The distribution of Artemisia rigida in

Washington: a challenge to ecology and geology. Northwest Science. 56: 162–164.

Diettert, R. A. 1938. The morphology of Artemisia tridentata Nutt.

Lloydia. 1: 3–74.

Drut, M. S.; Pyle, W. H.; Crawford, J. A. 1994. Technical note: diets

and food selection of sage-grouse chicks in Oregon. Journal of

Range Management. 47: 90–93.

Goodrich, S.; McArthur, E. D.; Winward, A. H. 1985. A new combination and a new variety in Artemisia tridentata. Great Basin

Naturalist. 45: 99–104.

Guralnik, D. B. 1971. Webster’s new world dictionary of the American language. New York, NY: The World Publishing Company.

882 p.

Hanks, D. L.; McArthur, E. D.; Stevens, R.; Plummer, P. A. 1973.

Chromatographic characteristics and phylogenetic relationships

of Artemisia section Tridentatae. Res. Pap. INT-141. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 1–23.

Heady, H. 1964. Palatability of herbage and animal preference.

Journal of Range Management. 17: 76–82.

Heywood V. H.; Harborne J. B; Turner B. L. 1977. The biology and

chemistry of the Compositae. In: A joint symposium sponsored by

the Linnean Society of London and the Phytochemical Society.

Vol. 1. New York: Academic Press: 338–357.

Hilty, J. H.; Eldridge, D. J.; Rosentreter, R.; Wicklow-Howard, M. C.

2003. Burning and seeding influence soil surface morphology in

an Artemisia shrubland in southern Idaho. Arid Land Research

and Management. 17: 1–11.

Kaltenecker, J. K.; Wicklow-Howard, M. C.; Rosentreter, R. 1999.

Biological soil crusts in three sagebrush communities recovering

from a century of livestock trampling. In: McArthur, E. D.; Ostler,

K. W.; Wambolt, C. L., comps. Shrubland ecotones symposium:

proceedings; 1998 August 12–14; Ephraim, UT. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 222–226.

Kelsey, R. G. 2002. Natural Products Chemist. [Personal communication]. On file at: U.S. Department of Agriculture, Forest Service,

Pacific Northwest Research Station, Forestry Science Laboratory,

Corvallis, OR.

Kelsey, R. G.; Shafizadeh, F. 1978. Sesquiterpene lactones and

systematics of the genus Artemisia. Photochemistry. 18: 1591–1611.

Kelsey, R. G.; Shafizadeh, F. 1980. Glandular trichomes and sesquiterpene lactones of Artemisia nova (Asteraceae). Biochemical

Systematics and Ecology. 8: 371–377.

Kelsey, R. G.; Stevens, J. R.; Shafizadeh, F. 1982. The chemical

constituents of sagebrush foliage and their isolation. Journal of

Range Management. 35: 617–622.

Kelsey, R. G.; Reynolds, G. W.; Rodriguez, E. 1984. The chemistry

of biologically active constituents secreted and stored in plant

glandular trichomes. In: Rodriguez, E.; Healey, P. L.; Mehta, I.,
USDA Forest Service Proceedings RMRS-P-38. 2005 15

Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse Rosentreter

eds. Biology and chemistry of plant trichomes. New York, NY:

Plenum Press: 187–241.

Lowe-Dalzell, C.; Eldridge, D.; Pellant, M.; Wicklow-Howard, M. C.

2003. Post-fire establishment of sagebrush on Idaho rangelands

in the Western United States. African Journal of Range and

Forage Science 20: 370–372.

McArthur, E. D. 1979. Sagebrush systematics and evolution. In:

The sagebrush ecosystem: a symposium proceedings; 1978 April

27–28; Logan: Utah State University, College of Natural Resources: 14–22.

McArthur, E. D. 1983. Taxonomy, origin, and distribution of big

sagebrush (Artemisia tridentata) and allies (subgenus

Tridentatae). In: Johnson, K. L., ed. Proceedings of the first

Utah shrub ecology workshop; 1981 September 9–10; Ephraim,

UT. Logan: Utah State University, College of Natural Resources: 3–13.

McArthur, E. D.; Goodrich, S. K. 1986. Artemisia tridentata ssp.

spiciformis: distribution and taxonomic placement. In: McArthur,

E. D., Welch, B. L., comps. Proceedings—symposium on the

biology of Artemisia and Chrysothamnus; 1984 July 9–13; Provo,

UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Research Station:

55–57.

McArthur, E. D.; Plummer, P. 1978. Biogeography and management of native western shrubs: a case study, section Tridentatae

of Artemisia. Great Basin Naturalist Memoirs. 2: 229–243.

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.

Memmott, K. L.; Anderson, V. J.; Monsen, S. B. 1998. Seasonal

grazing impact on cryptogamic crusts in a cold desert system.

Journal of Range Management. 51: 547–550.

Miller, R. F.; Shultz, L. M. 1987. Development and longevity of

ephemeral and perennial leaves on Artemisia tridentata Nutt.

ssp. wyomingensis. Great Basin Naturalist. 47: 227–231.

Monsen, S. B.; Shaw, N. L. 1986. Response of an alkali sagebrush/

fescue site to restoration treatments. In: McArthur, E. D.; Welch,

B. L., comps. Proceedings—symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9–13; Provo, UT. Gen. Tech.

Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 126–133.

Morris, M. S.; Kelsey, R. G.; Griggs, D. 1976. The geographic and

ecological distribution of big sagebrush and other woody Artemisias in Montana. Proceedings of the Montana Academy of Sciences. 36: 56–79.

Robertson, D. R.; Nielsen, J. L.; Bare, N. H. 1966. Vegetation and

soils of alkali sagebrush and adjacent big sagebrush ranges in

North Park, Colorado. Journal of Range Management. 19: 17–20.

Rosentreter, R. 1984. The synecology of the epiphytic lichen,

Xanthoria fallax (Hepp) Arn. occurring on the three subspecies of

Artemisia tridentata Nutt. Missoula, MT: University of Montana.

173 p. Dissertation.

Rosentreter, R. 1986. Compositional patterns within a rabbitbrush

(Chrysothamnus) community of the Idaho Snake River Plain. In:

McArthur, E. D., Welch, B. L., comps. Proceedings—symposium

on the biology of Artemisia and Chrysothamnus; 1984 July 9–13;

Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research

Station. 273–277.

Rosentreter, R. 1992. Camas Prairie and possible evolutionary links

with old world Artemisia species. In: Clary, W. P.; McArthur, E. D.;

Bedunah, D.; Wambolt, C. L., comps. Proceedings—symposium

on ecology and management of riparian shrub communities; 1991

May 29–31; Sun Valley, ID. Gen. Tech. Rep. INT-289. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 223–227.

Rosentreter, R. 2001. Unpublished data on file at: Bureau of Land

Management, Boise, ID.

Rosentreter, R.; Eldridge, D. J. 2002. Monitoring biodiversity and

ecosystem function: grasslands, deserts, and steppe. In: Nimis, P. L.;

Scheidegger, C.; Wolseley, P. A., eds. Monitoring with lichens—

monitoring lichens. Dordrecht, The Netherlands: Kluwer Academic Publishers: 233–237.

Rosentreter, R.; Kelsey, R. G. 1991. Xeric big sagebrush, a new

subspecies in the Artemisia tridentata complex. Journal of Range

Management. 44: 330–335.

Rosentreter, R.; McCune, B. 1992. Vagrant Dermatocarpon in

Western North America. The Bryologist. 95: 15–19.

Scholl, J. P.; Kelsey, R. G.; Shafizadeh, F. 1977. Involvement of

volatile compounds of Artemisia in browse preference by mule

deer. Biochemical and Systematic Ecology. 5: 291–295.

Schwartz, C. C.; Regelin, W. L.; Nagy, J. G. 1980. Deer preference

for juniper forage and volatile oil treated foods. Journal of Wildlife Management. 44: 114–120.

Sheehy, D. P.; Winward, A. H. 1981. Relative palatability of seven

Artemisiataxa to mule deer and sheep. Journal of Range Management. 34: 397–399.

Slone, J. H.; Kelsey, R. G. 1985. Isolation and purification of

glandular secretory cells from Artemisia tridentata ssp. vaseyana

by percoll density gradient centrifugation. American Journal of

Botany. 72: 1445–1451.

Stevens, R.; McArthur, E. D. 1974. A simple field technique for the

identification of some sagebrush taxa. Journal of Range Management. 27: 325–326.

Striby, K. D.; Wambolt, C. L.; Kelsey, R. G.; Havstad, K. M. 1987.

Crude terpenoid influence on in vitro digestibility of sagebrush.

Journal of Range Management. 40: 244–247.

Thomas, A. E.; Rosentreter, R. 1992. Utilization of lichens by

pronghorn antelope in three valleys in east-central Idaho. Tech.

Bull. 92-3. Boise, ID: U.S. Department of the Interior, Bureau of

Land Management, Idaho State Office. 10 p.

Tisdale, E. W.; Hironaka, M. 1981. The sagebrush-grass region: a

review of the ecological literature. Bull. 33. Moscow: University

of Idaho, Forest, Wildlife and Range Experiment Station. 31 p.

TNC. 1997. Columbia Plateau ecoregional plan. Arlington, VA: The

Nature Conservancy. 365 p.

Vasquez, M. 2002. Determination of winter use of sagebrush subspecies by Gunnison sage-grouse on Flat Top Mountain: assessment for prescribed fire planning. Unpublished Report on file at:

U.S. Department of the Interior, Bureau of Land Management,

Boise, ID. 16 p.

Wambolt, C. L. 1996. Mule deer and elk foraging preference for 4

sagebrush taxa. Journal of Range Management. 49: 499–503.

Wambolt, C. L. 1998. Sagebrush and ungulate relationships on

Yellowstone’s northern range. Wildlife Society Bulletin. 26:

429–437.

Wambolt, C. L. 2001. Mule deer foraging preference among five

sagebrush (Artemisia L.) taxa. Western North American Naturalist. 64: 490–494.

Wambolt, C. L.; Kelsey, R. G.; Bray, R. O. 1991. Wildlife reactions

to sagebrush crude terpenoids. In: Fisser, H. G., ed. Proceedings

of the seventeenth Wyoming shrub ecology workshop; 1988 June

21–22; Jackson, WY. Laramie: Wyoming Shrub Ecology Workshop. 17: 37–40.

Wambolt, C. L.; Kelsey, R. G.; Personius, T. L.; [and others]. 1987.

Preference and digestibility of three big sagebrush subspecies

and black sagebrush as related to crude terpenoid chemistry. In:

Provenza, F. D.; Flinders, J. T.; McArthur, E. D., comps. Proceedings—symposium on plant herbivore interactions; 1985 August

7–9; Snowbird, UT. Gen. Tech. Rep. INT-222. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 71–73.

Wambolt, C. L.; Walhof, K. S.; Frisina, M. R. 2001. Recovery of big

sagebrush communities after burning in southwest Montana.

Journal of Environmental Management. 61: 243–252.

Ward, G. H. 1953. Artemisia, section Seriphidiumin North America:

a cytotaxonomic study. Contributions from the Dudley Herbarium. 4: 155–205.

Welch, B. L.; Davis, J. N. 1984. In vitro digestibility of Kochia

prostrata (L.) Schrad. Great Basin Naturalist. 44: 296–298.

Welch, B. L.; McArthur, E. D.; Davis, J. N. 1983. Mule deer

preference and monoterpoids (essential oils). Journal of Range

Management. 34: 380–384.

Welch, B. L.; Nelson, E. D.; Young, S. A.; Sands, A. R.; Wagstaff, F. J.

1992. Gordon Creek—a superior, tested germplasm of Wyoming

big sagebrush. Res. Pap. INT-461. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Research Station. 7 p.
16 USDA Forest Service Proceedings RMRS-P-38. 2005

Rosentreter Sagebrush Identification, Ecology, and Palatability Relative to Sage-Grouse

Welch, B. L.; Wagstaff, F. J.; Jorgensen, G. L. 1990. Hobble Creek

mountain big sagebrush seed production. In: McArthur, E. D.;

Romney, E. M.; Smith, S. D.; Tueller, P. T., comps. Proceedings—

symposium on cheatgrass invasion, shrub die-off and other aspects of shrub biology and management; 1989 April 5–7; Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 167–170

West, N. E. 1988. Intermountain deserts, shrub steppes, and woodlands. In: Barbour, B. G.; Billings, W. D., eds. North American

terrestrial vegetation. New York, NY: Cambridge University

Press: 210–230.

Winward A. H.; McArthur, E. D. 1995. Lahontan sagebrush (Artemisia arbuscula ssp. longicaulis): a new taxon. Great Basin

Naturalist. 55: 151–157.

Winward A. H.; Tisdale, E. W. 1977. Taxonomy of the Artemisia

tridentata complex in Idaho. Bull. 19. Moscow: University of

Idaho, Forest, Wildlife and Range Experiment Station. 15 p.

Yabann, W. K.; Burritt, E. A.; Malechek, J. C. 1987. Sagebrush

(Artemisia tridentata) monoterpenoid concentrations as factors

in diet selection by free-ranging sheep. In: Provenza, J. T.;

Flinders, J. T.; McArthur, E. D., comps. Proceedings—symposium on plant herbivore interactions; 1985 August 7–9; Snowbird, UT. Gen. Tech. Rep. INT-222. Ogden, UT: U.S. Department

of Agriculture, Forest Service, Intermountain Research Station:

64–70.

Young, J. 2001. Biologist. [Personal communication]. Gunnison,

CO: Western State College.

Arrowleaf balsamroot
USDA Forest Service Proceedings RMRS-P-38. 2005 17

Sherel Goodrich is an Ecologist, Ashley National Forest, Vernal, UT 84078.

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Classification and Capabilities of

Woody Sagebrush Communities

of Western North America With

Emphasis on Sage-Grouse

Habitat

Sherel Goodrich

Abstract—This paper deals with diversity, classification, and

capabilities of different sagebrush (Artemisia spp.) communities. Capabilities of sagebrush communities in terms of production, plant

diversity, potential for ground cover and sage-grouse (Centrocercus

urophasianus) habitat are discussed. Reaction to fire and relationships

with introduced annuals are also discussed for some communities.

Introduction ____________________

There are about 30 taxa of woody and semiwoody sagebrush (Artemisia spp.) in western North America. Many of

these sagebrush taxa are community dominants that are

highly specialized. They are high-resolution indicators of

climate, geomorphic settings, soils, elevations, and other

features of the environment. Ecotones between stands of

different species of sagebrush are often narrow and even

sharply abrupt. Ecotones between taxa of the same species

are often wider, but these are also sometimes quite narrow.

Sagebrush taxa are often more site specific than many

associated indicator species. Bluebunch wheatgrass (Elymus

spicatus) and Idaho fescue (Festuca ovina var. ingrata)

provide vivid examples. These are useful indicator plants in

the sagebrush ecosystem. However, they are generalists

compared to some of the sagebrush taxa. Respectively, these

grasses are listed as indicator species in 9 and 11 sagebrush

types, and they extend beyond the amplitude of sagebrush

into grassland (Mueggler and Stewart 1980; Tisdale 1986),

mountain brush, and some forested communities.

Hybrids and hybrid zones of sagebrush often indicate

intermediate environments (Freeman and others 1999; Graham and others 1999; McArthur and Sanderson 1999b).

Hybrid zones coupled with a propensity for development of

polyploidy further increase the complexity of the sagebrush

group (McArthur and Sanderson 1999a; McArthur and others 1981). With many highly specialized taxa, hybrid zones,

and polyploidy, the sagebrush complex is highly diverse.

Classification of vegetation types inherently includes information useful in understanding capabilities of the land,

and many classification works include management implications based on estimates or measurements of capabilities.

Most classification of sagebrush vegetation types has been

at local scales and based on different methods of data

collection and ordination. Although a comprehensive and

uniform classification would be useful, the work of the past

provides considerable information about the sagebrush ecosystem. Classification works were relied upon heavily in

preparing this paper. Units of sagebrush classification found

in the literature include community types, plant associations, habitat types, and ecological units. Distinction of

classification types used by different workers is not made in

this paper. All methods of classification seem to be useful in

helping to define the variability and capability of sagebrush

systems. Recognizing differences in capabilities of sagebrush communities is important for restoration projects,

management for desired condition, and other aspects of land

management.

Evaluations of values for sage-grouse (Centrocercus

urophasianus) habitat made in this paper are based on

indicated capabilities of various sagebrush types in comparison with tables 1, 2, and 3 in Connelly and others (2000). In

general Connelly and others (2000) indicated sage-grouse

nest sites are associated with sagebrush with heights of 29

to 80 cm and crown cover of 15 to 38 percent. Characteristics

needed for productive sage-grouse winter habitat include

sagebrush with heights of 25 to 35 cm and crown cover of 10

to 30 percent. Characteristics needed for productive breeding and brood-rearing habitat include sagebrush with heights

of 30 to 80 cm and crown cover of 15 to 25 percent for breeding

habitat and 10 to 25 percent for brood-rearing habitat.

The majority of this paper is focused on Wyoming big

sagebrush (A. tridentata ssp. wyomingensis), Vasey big

sagebrush, and mountain big sagebrush (A. t. ssp. vaseyana)

communities. These taxa are selected for greater discussion

because of their comparatively wide distribution, the large

area they cover, and their greater importance to sage-grouse

than many of the other taxa.

In this paper common names of plants are not used in

reference to classification. However, they are used in the

text, and a list of scientific names including authors and

corresponding common names is provided in tables 1 and 2.

Symbols of plant taxa as listed by athe U.S. Department of
18 USDA Forest Service Proceedings RMRS-P-38. 2005

Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Agriculture (2001) are also included in tables 1 and 2. Only

indicator species of vegetation types are listed. Expanded

lists are found in some of the works listed in the references

section. For the most part, plant nomenclature in this paper

follows that of Shultz (1986) for Artemisia and Welsh and

others (1993) for other plant species.

References to production are generally based on standing

crop (a measurement at one time of the year). Thus, plants

that grow in spring and early summer then dry and wither

away are often not well represented in production values. As

used in this paper, ground cover includes the percentage of

the ground surface covered by live plants, litter or plant

residue, and rock. Bare soil and pavement of less than about

2 cm diameter are generally not considered ground cover.

For the cited references, however, definitions of bare ground

may vary among authors.

Artemisia arbuscula _____________

This plant of western North America has been treated as

a subspecies and as a variety of A. tridentata. However, it

has been accepted at the species level by many Intermountain authors. Shultz (1986) recognized three subspecies in

this complex and provided distribution maps for each.

Winward and McArthur (1995) described a fourth subspecies.

These are listed below. All plants of the complex are low,

nonsprouting, wintergreen shrubs.

Artemisia arbuscula ssp. arbuscula

Low Sagebrush _________________

This plant is the most widespread taxon of the complex. It

ranges from Washington to Wyoming and south to California and Utah.

Classification

Artemisia arbuscula ssp. arbuscula/Bunchgrass (ElymusFestuca) (Hall 1973)

Artemisia arbuscula ssp. arbuscula/Elymus spicatus

(Hironaka and others 1983: Jensen 1989; Mueggler and

Stewart 1980; Nelson and Jensen 1987; Tew 1988; Zamora

and Tueller 1973)

Artemisia arbuscula ssp. arbuscula/Festuca idahoensis

(Hironaka and others 1983; Jensen 1989; Mueggler and

Table 1—Scientific name, codes, and common names of sagebrush type indicator plants: woody plants.

Scientific name Code Common name

Artemisia arbuscula Nutt. ssp. arbuscula ARARA Low sagebrush

Artemisia arbuscula ssp. longicaulis Winward & McArthur ARARL3 Lahontan sagebrush

Artemisia arbuscula ssp. thermopola Beetle ARART Cleftleaf sagebrush, hotsprings sagebrush

Artemisia bigelovii Gray ARBI3 Bigelow sagebrush

Artemisia cana Pursh ssp. cana ARCAC5 Plains silver sagebrush

Artemisia cana ssp. bolanderi (A. Gray) Ward ARCAB3 Sierra silver sagebrush

Artemisia cana ssp. viscidula (Osterh.) Beetle ARCAV2 Mountain silver sagebrush

Artemisia filifolia Torrey ARFI2 Sand sagebrush

Artemisia frigida Willd. ARFR4 Fringed sagebrush

Artemisia longiloba (Osterh.) Beetle ARLO9 Alkali sagebrush, early sagebrush

Artemisia nova A. Nels. var. nova ARNO4 Black sagebrush

Artemisia nova var. duchesnicola Welsh & Goodrich ARNOD Red clay sagebrush

Artemisia papposa S. F. Blake & Cronquist ARPA16 Owyhee sagebrush

Artemisia pedatifida Nutt. ARPE6 Birdsfoot sagebrush

Artemisia porteri Cronq. ARPO5 Porter sagebrush

Artemisia pygmaea A. Gray ARPY2 Pygmy sagebrush

Artemisia rigida (Nutt.) A. Gray ARRI2 Scabland sagebrush; stiff sagebrush

Artemisia rothrockii A. Gray ARRO4 Rothrock sagebrush

Artemisia spinescens D. C. Eaton ARSP5 Bud sagebrush, budsage

Artemisia tridentata ssp. parishii (A. Gray) Hall & Clements ARTRP2 Parish big sagebrush

Artemisia tridentata ssp. spiciformis (Osterh.) Kartesz & Gandhi ARTRS2 Spiked big sagebrush, subalpine big sagebrush

Artemisia tridentata ssp. tridentata ARTRT Basin big sagebrush

Artemisia tridentata ssp. vaseyana var. pauciflora Winward & Goodrich ARTRP4 Mountain big sagebrush

Artemisia tridentata ssp. vaseyana (Rydb.) Beetle var. vaseyana ARTRV Vasey big sagebrush

Artemisia tridentata ssp. wyomingensis Beetle & Young ARTRW8 Wyoming big sagebrush

Artemisia tridentata ssp. xericensis Rosentreter and Kelsey ARTRX Xeric big sagebrush

Artemisia tripartita Rydb. ssp. tripartita ARTRT2 Threetip sagebrush

Artemisia tripartita ssp. rupicola Beetle ARTRR4 Wyoming threetip sagebrush

Atriplex confertifolia (Torr. & Frem) Wats. ATCO Shadscale

Chrysothamnus nauseosus (Pallas) Britt. CHNA2 Rubber rabbitbrush

Grayia spinosa (Hook.) Moq. GRSP Spiny hopsage

Purshia tridentata (Pursh) DC. PUTR2 Bitterbrush

Sarcobatus vermiculatus (Hook.) Torr. SAVE4 Greasewood

Symphoricarpos occidentalis Hook. SYOC Wolfberry

Symphoricarpos oreophilus Gray SYOR Mountain snowberry
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

Stewart 1980; Nelson and Jensen 1987; Volland 1976;

Zamora and Tueller 1973)

Artemisia arbuscula ssp. arbuscula/Festuca idahoensis/

Poa secunda (Nelson and Jensen 1987)

Artemisia arbuscula ssp. arbuscula/Stipa thurberiana

(Zamora and Tueller 1973)

Artemisia arbuscula ssp. arbuscula/Purshia tridentata/

Agropyron spicatum (Zamora and Tueller 1973)

Of the plants recognized above as indicator species of

classification units within the low sagebrush type, all likely

have broader ecological amplitude than does low sagebrush.

This taxon appears to be the most specialized of any of the

associates listed above.

Habitat and Capabilities

Low sagebrush forms stands below the pinyon-juniper

(Pinus-Juniperus) belt, within this belt, and well above it.

Stands in the Toiyabe Range of Nevada extend up to 3,200 m

elevation or higher (Goodrich 1981) and up to 3,900 m in the

White Mountains of California (Mozingo 1987). Perhaps low

sagebrush is less common below the pinyon-juniper belt

than is black sagebrush (A. nova).

Low sagebrush is often an indicator of soils with clay

subsurface horizons or bedrock within 8 to 33 cm of the

surface (Fosberg 1963; Summerfield and Peterson 1971).

This plant is sometimes found where restrictive soil horizons cause water to stand on the surface in late winter and

spring, during which time the soils are poorly aerated

(Hironaka 1963). The restrictive soil layers also tend to

reduce water-holding capacity in summer. The soil features

associated with low sagebrush indicate a habitat with

potential for soil saturation in spring and drier conditions

in summer than are typical for big sagebrush.

Sabinske and Knight (1978) found low sagebrush on

gravelly soils without a restrictive horizon in Wyoming. In

the mountain ranges of central Nevada, low sagebrush

occupies shallow, rocky, well-drained soils as well as those

with heavy clay horizons (Goodrich 1981).

Low sagebrush is highly competitive on sites where it is

well adapted. It will reestablish from seed after disturbance,

but it does not sprout. Comparatively low productivity and

other site features indicate difficulty and low economic

return for restoration and forage improvement projects in

this type. Such projects should be planned and conducted

carefully (Johnson 1987; Winward 1980). Low-elevation

stands can be expected to be vulnerable to displacement by

cheatgrass (Bromus tectorum). High-elevation stands are

likely to be more resistant.

Annual production of between 392 to 728 kg/ha (350 to 650

lbs/acre) has been reported for low sagebrush communities

(Hall 1973; Jensen 1989; Nelson and Jensen 1987; Tew

1988). Volland (1976) listed a mean of 200 kg/ha (179 lbs/

acre) for sites in fair condition in the pumice zone of Oregon.

As implied by its common name, low sagebrush is short in

stature. Height of low sagebrush is commonly 20 to 40 cm

and less commonly from 10 to 50 cm (Cronquist 1994;

Table 2—Scientific name, symbols, and common names of sagebrush type indicator plants: graminoids.

Scientific name Symbol Common name

Bouteloua gracilis (H.B.K.) Lag. ex Steudel BOGR Blue gramma

Bromus carinatus H. & A. BRCA5 Mountain brome

Carex geyeri F. Boott CAGE2 Elk sedge

Deschampsia cespitosa (L.) Beauv. DECE Tufted hairgrass

Elymus cinereus Scribn. & Merr. ELCI2 Basin wildrye

Elymus elymoides (Raf.) Swezey ELEL5 Bottlebrush squirreltail

Elymus lanceolatus (Scribn. & Sm.) Gould ELLA3 Thickspike wheatgrass

Elymus salinus Jones ELSA Salina wildrye

Elymus smithii (Rydb.) Gould ELWM3 Western wheatgrass

Elymus spicatus (Pursh) Gould ELSP3 Bluebunch wheatgrass

Elymus trachycaulus (Link.) Gould ex Shinn. ELTR7 Slender wheatgrass

Festuca ovina L. var. ingrata Hackel ex Beal (Festuca idahoensis Elmer) FEOVI Idaho fescue

Festuca ovina var. rydbergii St. Yves FEOVR Sheep fescue

Festuca scabrella Torr. FESC Rough fescue

Festuca thurberi Vasey FETH Thurber fescue

Koeleria macrantha (Ledeb.) Schultes KOMA June grass

Leucopoa kingii (Wats.) W. A. Weber LEKI2 Spike fescue

Poa fendleriana (Stuedel) Vasey POFE Muttongrass

Poa pratensis L. POPR Kentucky bluegrass

Poa secunda Presl POSE Sandberg bluegrass

Sporobolus cryptandrus (Torr.) Gray SPCR Sand dropseed

Stipa comata Trin. & Rupr. var. comata STCOC2 Needle-and-thread grass

Stipa comata var. intermedia Scribn. & Tweedy STCOI Mountain needle-and-thread grass

Stipa hymenoides R. & S. STHY6 Indian ricegrass

Stipa lettermanii Vasey STLE4 Letterman needlegrass

Stipa nelsonii Scribn. STNE3 Columbia needlegrass

Stipa richardsonii Link STRI2 Richardson needlegrass

Stipa thurberiana Piper STTH2 Thurber needlegrass

Trisetum spicatum (L.) Richter TRSP2 Spike trisetum
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Harrington 1954; Welsh and others 1993). In four habitat

types in Nevada, maximum height of low sagebrush is

between 11 to 19 cm and crown cover is between 12 to 16

percent (Zamora and Tueller 1973).

Hall (1973) listed 13 percent surface rock, 10 percent

erosion pavement, and 16 percent bare ground for a low

sagebrush/bunchgrass type in Oregon and Washington.

Tables in Tew (1988) indicate ground surface with an average of 21 percent bare soil, 42 percent vegetation and litter,

and the remaining 37 percent rock and pavement. Volland

(1976) listed 11 percent rock and 42 percent bare ground and

pavement for a type in fair condition in the pumice zone of

Oregon.

Low sagebrush sites of low and moderate productivity are

indicated to lack the capability to meet the requirements of

shrub height and percent crown cover for sage-grouse nesting habitat. Some of the more productive sites might provide

moderate value nesting habitat. However, the crowns of low

sagebrush are often in contact with the ground or close

enough to the ground to obstruct sage-grouse nesting.

Low sagebrush sites dry rapidly in early summer. This

indicates moderate-value brooding habitat in the spring and

low value in summer and fall. Small stands of low productivity likely provide high-value strutting habitat especially

where they are mixed with stands of big sagebrush (Artemisia tridentata). On windswept areas low sagebrush might

provide high-value winter forage. However, snow can be

expected to cover shrubs of this community on flat and

concave areas before shrubs of big sagebrush are covered.

Artemisia arbuscula ssp. longicaulis

Lahontan Sagebrush ____________

This plant is known from western Nevada, southern

Idaho, and eastern California where the old shorelines of

Pleistocene Lake Lahontan are one of the centers of its

current distribution (Winward and McArthur 1995).

Apparently this taxon has not been included in published

vegetation type classifications. It is interspersed with salt

desert shrub species including shadscale. It is also associated with Wyoming big sagebrush (Artemisia tridentata ssp.

wyomingensis), black sagebrush, and low sagebrush. The

most common understory species include bluebunch wheatgrass, Thurber needlegrass (Stipa thurberiana), desert

needlegrass (Stipa speciosa), Indian ricegrass (S.

hymenoides), bottlebrush squirreltail (Elymus elymoides),

and Sandberg bluegrass (Poa sandbergii) (Winward and

McArthur 1995). The value of Lahontan sagebrush to

sage-grouse is likely similar to that of other taxa of the

A. arbuscula complex.

Artemisia arbuscula ssp. thermopola

Cutleaf Sagebrush ______________

This plant ranges from central Idaho to western Wyoming

and south to northern Utah. It is restricted to areas adjacent

to those occupied by A. tripartita, which perhaps hybridized

with A. arbuscula to form A. a. thermopola (Beetle 1960;

Shultz 1984).

Classification

Artemisia arbuscula thermopola/Festuca idahoensis

(Hironaka and others 1983)

Habitat and Capabilities

Habitat includes ridgetops and glacial outwash areas that

are thinly mantled, poorly drained, and usually within

forest openings. Soils are shallow and have a strongly

developed clay horizon (Hironaka and others 1983). Low

stature of shrubs and herbaceous species can be expected to

be inherent to stands of cutleaf sagebrush. K. Johnson

(1987) listed this as a spreading, lax shrub up to 30 cm tall.

Value for sage-grouse likely depends upon the size of

forest openings in which stands are found. Smaller openings

can be expected to have lower value than larger openings.

Low site productivity indicates that stature and crown cover

of cutleaf sagebrush will be of low value for nesting habitat.

Soil and habitat features listed by Hironaka and others

(1983) indicate early drying of herbaceous species and thus

low value for brooding habitat. Windswept knolls might

provide moderate- to high-value breeding and winter habitat

where these are adjacent to stands of big sagebrush.

Shallow soils and poor drainage indicate low productivity.

Restoration projects can be expected to be of low success due

to the short growing season and water stress occurring

during the seedling establishment stage (Hironaka and

others 1983).

Artemisia bigelovii

Bigelow Sagebrush ______________

Bigelow sagebrush is known from California east to Colorado and Texas. Habitat includes rimrock, cliff faces, canyons, and ravines within the desert shrub and pinyonjuniper belts. Shrubs are commonly 10 to 30 cm tall with the

flower stalks often equal to or greater than the length of the

woody stems. The flower stalks die back each year.

The stature and growth form of Bigelow sagebrush is not

conducive to sage-grouse nesting. Sage-grouse generally

show low selectivity for the habitat of this plant. Where

stands of Bigelow sagebrush are adjacent to big sagebrush

stands, it might be used by sage-grouse as winter forage or

sparingly at other times of the year.

Artemisia cana ssp. bolanderi

Bolander Silver Sagebrush _______

Bolander silver sagebrush is primarily a plant of the

Sierra Nevada Mountains of California. It extends into

Oregon and barely into Nevada.

Habitat and Capabilities

This shrub occurs along streams and in snow catchment

basins on granitic soils (Shultz 1986). Its value as sagegrouse habitat is likely similar to that of mountain silver

sagebrush (A. cana ssp. visicidula). However, prolonged

flooding of some sites likely limits the values of this type for
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

sage-grouse habitat. The plant is known from the margin of

the range of sage-grouse.

Artemisia cana ssp. cana

Plains Silver Sagebrush __________

Plains silver sagebrush ranges from Montana and North

Dakota south to northern Colorado and western Nebraska.

As indicated by its common name, this is a plant of the plains

and it is mostly found east of the Continental Divide.

However, outlier populations have been found in Sweetwater

County, Wyoming; Daggett County, Utah; and in Moffat County

(Yampa River Valley) and Rio Blanco County, Colorado, on the

west side of the Continental Divide.

Classification

Artemisia cana ssp. cana/Elymus smithii (Johnston [1987]

listed several references for the Plains.)

Artemisia cana ssp. cana/Symphoricarpos occidentalis/E.

smithii (Johnston [1987] listed three references for North

Dakota.)

Artemisia cana ssp. cana/Elymus cinereus (Thatcher 1959)

Habitat and Capabilities

Plains silver sagebrush is most common on deep, welldrained soils on alluvial flats and terraces along watercourses (Walton and others 1986). This is well demonstrated

in the distribution map by Beetle (1960) where the distribution of this plant is represented by linear lines following

watercourses. It extends beyond watercourses especially on

sandy soils (Thatcher 1959). Although not requiring a high

water table, plains silver sagebrush can tolerate a water

table within the rooting zone. Thatcher (1959) found it on

soils with pH between 6.6 and 8.5 with most sites having a

pH of above 7.4; he did not find it on heavy-textured soils

regardless of topographic position. In Albany County, Wyoming, Thatcher (1959) found plains silver sagebrush from

about 18 to 102 cm tall with shorter plants on upland sites.

Within its range, it is capable of growing up to 1.5 m tall

(Walton and others 1986) on better sites.

Stature of plains silver sagebrush indicates high value for

sage-grouse for nesting and for winter habitat. This shrub is

capable of forming dense stands (Walton and others 1986),

which indicates high value for nesting habitat. However, the

crown-form of plains silver sagebrush might not be of equal

value to that of Vasey big sagebrush or Wyoming big sagebrush. The plains habitat indicates productive understory

communities. Where stands occur with sufficient shrub

crown cover, plains silver sagebrush might be expected to

provide yearlong, high-value habitat for sage-grouse. However, linear stands of this plant might not have as high a

value as do broad areas covered by Wyoming big sagebrush.

Artemisia cana Pursh ssp. viscidula

Mountain Silver Sagebrush _______

Mountain silver sagebrush ranges from western Montana

south to Nevada and Colorado.

Classification

Artemisia canaPursh ssp. viscidula/Deschampsia cespitosa

(Padgett and others 1989)

Artemisia cana Pursh ssp. viscidula/Elymus trachycaulus

(Bramble-Brodahl 1978; Tew 1988)

Artemisia cana Pursh ssp. viscidula/Festuca idahoensis

(Bramble-Brodahl 1978; Hironaka and others 1983;

Youngblood and others 1985)

Artemisia canaPursh ssp. viscidula/Festuca ovina (Padgett

and others 1989; Tew 1988)

Artemisia cana Pursh ssp. viscidula/Festuca thurberi

(Francis 1983; Johnston [1987] listed six references for

Colorado; Tew 1988)

Artemisia canaPursh ssp. viscidula/Poa pratensis (Padgett

and others 1989; Youngblood and others 1985)

Artemisia cana Pursh ssp. viscidula/Symphoricarpos

oreophilus/Elymus trachycaulus (Tew 1988)

Habitat and Capabilities

This is a plant of montane valleys where it is commonly

associated with riparian communities. Soils of some communities show evidence of seasonally high water tables (Padgett

and others 1989). Communities of mountain silver sagebrush often form between wetlands of sedges (Carex spp.)

and willows (Salix spp.) and uplands of big sagebrush, aspen

(Populus tremuloides), and coniferous forests. Occasionally,

mountain silver sagebrush forms stands on upland sites well

removed from riparian communities.

In the cool and moist setting of mountain silver sagebrush

communities, cool season graminoids are highly productive.

Tables from Tew (1988) from central Utah show total herbage production ranges from 858 kg/ha (855 lbs/acre) for the

low end of an A. c. viscidula/Festuca ovina type to 1,919 kg/

ha (1,713 lbs/acre) for an A. c. viscidula/Festuca thurberi

type. With high production, potential for ground cover is also

high. Tables from Tew (1988) indicate potential ground

cover ranges from 84 to 97 percent with an average of 92

percent.

Mountain silver sagebrush is a multiple-stemmed shrub

with capacity to sprout from the root crown. Although the

leaves of mountain silver sagebrush persist though some or

all of the winter, they dry upon freezing and many fall.

Common grasses include those listed above as vegetation

type indicators. Succulent forbs including dandelion (Taraxacum officinale), longfoot clover (Trifolium longipes), yarrow (Achillea millefolium), and everywhere aster (Aster

chilensis) are often present, and they likely increase with

heavy, persistent livestock grazing.

Many stands of mountain silver sagebrush are likely too

wet or snow covered to be of more than low-value nesting

habitat for sage-grouse. Many are at elevations of deep snow

accumulations in winter where low value for winter habitat

is indicated. Drying and deciduous leaves indicate mountain

silver sagebrush has low value for winter forage for sagegrouse. Crowns of mountain silver sagebrush are typically

thin and ascending compared to the dense, spreading crowns

of Vasey big sagebrush and Wyoming big sagebrush. This is

likely a limiting factor in value for sage-grouse nesting

habitat.
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Presence of succulent forbs and proximity to water and

meadows indicate high value for brood-rearing habitat for

sage-grouse. This seems especially so where mountain

silver sagebrush stands are adjacent to, or near to, stands

of big sagebrush. Where mountain silver sagebrush communities are adjacent to stands of big sagebrush, value for

sage-grouse is indicated to be higher than where they are

surrounded by coniferous forests.

High elevation, cool climate, relatively high annual precipitation, and high productivity of aggressive perennials

indicate high resistance to invasion of cheatgrass.

Artemisia filifolia

Sand Sagebrush ________________

This is a plant of the western Great Plains and Colorado

Basin from eastern Wyoming and South Dakota, south to

Arizona and New Mexico and extending into Texas and

Mexico.

Classification

Artemisia filifolia/Andropogon hallii(Johnston [1987] listed

two references for the Comanche and Cimarron National

Grasslands.)

Artemisia filifolia/Sporobolus cryptandrus-Bouteloua gracilis (Johnston [1987] listed a reference for the Comanche

National Grasslands.)

Habitat and Capabilities

Sand sagebrush is most common on sandy soil including

dunes. The plants are from 50 to 150 cm tall, and they have

a stature that would indicate favorable cover for sagegrouse. However, due to the sparse, filiform leaves, the

crowns of sand sagebrush provide a thin cover. Shrub density in communities of sand sagebrush is also sometimes

rather low.

Features of this shrub and its communities indicate low

value for sage-grouse at any time of the year. The distribution

of sand sagebrush is mostly at the margin and well beyond the

current and historic range of sage-grouse. This could indicate

sage-grouse did not find suitable habitat in sand sagebrush

communities during their evolutionary history.

Artemisia frigida

Fringed Sagebrush ______________

This subshrub ranges from Alaska to the Atlantic and

south to Arizona and Kansas. It is a characteristic species of

the high plains of central North America, and it is disjunct

in Siberia (Cronquist 1994; Welsh and others 1993).

Classification

Artemisia filifolia/Elymus spicatus (Francis 1983; Tew 1988)

Although Baker and Kennedy (1985) did not formally

recognize a fringed sagebrush association, they did note the

presence of this plant in a bluebunch wheatgrass/cushion

plant community of rocky, wind-exposed summits.

Habitat and Capabilities

Fringed sagebrush is a subshrub that dies back to near the

base in winter in most locations. Where it is a dominant or

codominant, it often indicates windswept sites. It often

grows as a minor component in many communities in more

protected sites. Fringed sagebrush can readily pioneer disturbed sites, and it occurs through all successional stages

including climax (Wambolt and Frisina 2002).

Stature and crown cover of fringed sagebrush rarely, if

ever, meet the needs of sage-grouse for nesting, broodrearing, or wintering. Where stands of fringed sagebrush are

adjacent to those of big sagebrush, they might be used as

strutting grounds. Windswept areas might provide winter

forage. As fringed sagebrush dies back in winter, availability of active leaves can be expected to decrease. However, the

lower leaves remain green throughout winter. Where these

leaves are not covered by snow, they can be expected to be of

some value to sage-grouse in winter.

Artemisia longiloba

Early Sagebrush, Longleaf

Sagebrush, Alkali Sagebrush______

Early sagebrush ranges from extreme southeastern Oregon to western Wyoming and south to Nevada and Colorado. Early sagebrush has been treated as a part of the A.

arbusculacomplex. However, unlike taxa of the A. arbuscula

complex, this plant flowers in late spring and early summer

instead of fall. The difference in flowering period seems to

indicate considerable phylogenetic distance from A. arbuscula.

Artemisia longiloba also differs from the A. arbuscula complex in habitat and, to some extent, in geography. Recognition at the species level is consistent with treatments by

Welsh and others (1993) and Wambolt and Frisina (2002).

Classification

Artemisia longiloba/Elymus spicatus (Bramble-Brodahl 1978;

Robertson and others 1966)

Artemisia longiloba/Festuca idahoensis (Bramble-Brodahl

1978; Hironaka and others 1983; Nelson and Jensen 1987;

Zamora and Tueller 1973)

Artemisia longiloba/Poa secunda (Francis 1983)

Habitat and Capabilities

Habitat includes fine-textured soils derived from alkaline

shales in clay basins and in mountains (Schultz 1986;

Wambolt and Frisina 2002). It was found only on claypan

soils in Idaho (Hironaka and others 1983), on the Humboldt

National Forest, Nevada (Nelson and Jensen 1987), and at

North Park, Colorado (Robertson and others 1966). Soil

features indicate curing of herbaceous plants in late spring

or early summer.

Francis (1983) recorded only 24 plant species per stand in

an early sagebrush type in Colorado compared to 30 to 40

species in Wyoming big sagebrush types and 55 to 67 species

in Vasey big sagebrush types. Zamora and Tueller (1973)

listed maximum shrub height at 24 cm and shrub crown
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

cover at 19 percent for a habitat type in northern Nevada.

Average annual production of 504 kg/ha (450 pounds/acre)

(Nelson and Jensen (1987) and 572 kg/ha (510 pounds/acre)

(Robertson and others 1966) have been reported for northeastern Nevada and northern Colorado, respectively.

Productivity, succulence of summer herbage, shrub height,

and crown cover of most early sagebrush stands indicate

relatively low value for sage-grouse nesting, brooding, and

wintering habitat. Small stands might provide moderate to

high-value strutting habitat, especially where they are adjacent to larger stands of big sagebrush. Windswept sites

might provide some forage in winter. Heavy clay subsoil and

shallow depth to bedrock limit the potential for management

(Nelson and Jensen 1987).

Artemisia nova

Black Sagebrush ________________

This nonsprouting, evergreen, low shrub ranges from

Oregon to Montana and south to eastern California and

north-central New Mexico.

Classification

Artemisia nova/Bouteloua gracilis (Tew 1988)

Artemisia nova/Elymus spicatus (Baker and Kennedy 1985;

Hironaka and others 1983; Nelson and Jensen 1987; Tew

1988; Thatcher 1959; Zamora and Tueller 1973)

Artemisia nova/Festuca idahoensis (Hironaka and others

1983)

Artemisia nova/Stipa comata (Baker and Kennedy 1985;

Zamora and Tueller 1973)

Artemisia nova/Stipa hymenoides (Nelson and Jensen 1987;

Zamora and Tueller 1973)

Artemisia nova/Atriplex confertifolia/Elymus elymoides

(Nelson and Jensen 1987)

Habitat and Capabilities

Black sagebrush has a rather wide elevational range that

includes valleys within the cold desert shrub belt to slopes

and ridges of mountains up into the lower end of the coniferous forest belt. It is noted to extend from 1,524 to 3,353 m in

California (Munz 1959). Its extension upslope into montane

areas is often facilitated by calcareous substrates.

Soils of black sagebrush communities are often basic and

shallow or rocky or both. Compared to soils of big sagebrush

taxa, soils of black sagebrush communities often have the

lowest water-holding capacity, organic matter, and nitrogen

content, and the thinnest mollic epipedons (Jensen 1990). In

cold desert areas, soils can be expected to be without mollic

epipedons.

Cold Desert Stands—Stands of black sagebrush in the

cold desert shrub belt appear to be highly competitive with

other native plants, and they appear to be long-persistent

with little potential for displacement by other shrubs. However, in some areas black sagebrush is highly selected by

ungulates, and heavily browsed stands are thinned and

replaced by shrubs more resistant to grazing (Clary 1986;

Holmgren and Hutchings 1972; Hutchings and Stewart

1953). With the introduction of cheatgrass, cold desert

stands of black sagebrush are highly subject to conversion to

cheatgrass systems. Replacement or displacement of black

sagebrush stands in cold desert shrub systems can be expected to be long-persistent with low potential for recovery

of sagebrush.

Crown cover of black sagebrush in stands of the cold desert

belt is often limited to less than 15 percent by the low

capacity of these sites to support vegetation. Height of black

sagebrush in cold desert areas is mostly less than 25 cm.

Cold desert shrub communities inherently have low potential for perennial forbs. Limitations are mostly a function of

precipitation, which is commonly as low as 15 to 18 cm/year.

Depending on amount and timing of precipitation, forbs

might flourish in some years and be essentially absent in

others.

Canopy cover and height of black sagebrush in cold desert

areas and low presence of succulent forbs indicate low (or no)

value for sage-grouse for nesting and brooding. Some stands

might have moderate value in winter, or other times of the

year, if they are adjacent to stands of Wyoming big sagebrush.

Artemisia nova var. duchesnicola (red clay sagebrush) is

endemic to the Duchesne River Formation in the Uinta

Basin in northeastern Utah (Welsh and Goodrich 1995).

Plants of this taxon are restricted to the cold desert shrub

environment. The discussion of values for the cold desert

shrub belt of black sagebrush are applicable to communities

of red clay sagebrush.

Disturbances that reduce black sagebrush in cold desert

systems can be expected to be long-persistent, and recovery

of black sagebrush can be expected to be slow. Risk of longterm loss of black sagebrush is high in cold desert areas

compared to montane stands.

Stands Within the Pinyon-Juniper Belt—Stands of

black sagebrush are common within the pinyon pine (Pinus

spp.) and juniper (Juniperus spp.) belt where they are

clearly seral to pinyon-juniper. Pinyon-juniper trees have

advanced into many black sagebrush communities in the

past several decades. Degree of displacement depends on

time since disturbance and other factors. Stands within the

pinyon-juniper belt are also highly vulnerable to conversion

to cheatgrass-driven communities. Although black sagebrush is highly sensitive to fire, the long-term maintenance

of black sagebrush within the pinyon-juniper belt requires

periodic fires to reduce conifer encroachment. The potential

for cheatgrass invasion greatly complicates management of

black sagebrush communities within the pinyon-juniper

belt in many areas.

Value of black sagebrush stands within the pinyon-juniper belt for sage-grouse is dependent on fire or other disturbance that reduces pinyon-juniper cover. Height and crown

cover of black sagebrush on better sites in the pinyon-juniper

belt are often within the limits indicated by Connelly and

others (2000) for at least winter habitat. However, capability

for height and crown cover decreases as site potential decreases. Sites of low to moderate potential for sagebrush

cover have low value for sage-grouse nesting. These limitations also indicate low value for winter habitat except where

stands are adjacent to more productive sagebrush sites.

Early drying of herbaceous plants in summer indicates low

value for brood rearing.
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Montane Stands—Stands of black sagebrush above the

pinyon-juniper belt are often quite stable but they are

sometimes vulnerable to displacement by Douglas-fir

(Pseudotsuga menziesii) and other conifers. These stands

appear to be more resistant to displacement by cheatgrass

than those of lower elevations.

Black sagebrush can be highly competitive in montane

stands. At two sites on the Tavaputs Plateau (Ashley National Forest [ASNF] 67-26, 68-1) stands of black sagebrush

were plowed and seeded with crested wheatgrass (Agropyron cristatum) and other aggressive introduced species.

Where protected from livestock and lagomophs, black sagebrush returned to pretreatment status within 20 years, and

essentially displaced the seeded grasses.

At another site (ASNF 45-6) on the Uinta Mountains,

Ashley National Forest, black sagebrush was sprayed with

2,4-D. Percent sagebrush kill was high and grass production

increased 3.2 times over what it was prior to treatment and

in the control. At 14, 19, and 23 years post treatment, crown

cover of black sagebrush was measured at 5, 12, and 17

percent, respectively. At 23 years post treatment, crown

cover in the treated area equaled that of the control.

Black sagebrush does not sprout after fire. However it is

capable of vigorous seedling establishment following fire.

Montane stands of black sagebrush on the Tavaputs Plateau

have returned to pretreatment crown cover within about 20

years after burning.

Crown cover of black sagebrush in stands at the upper

edge of the pinyon-juniper belt and above have been measured at 27 percent (ASNF 68-1) and 28 percent (ASNF 68-69)

at sites where use by wild ungulates has been minor. In the

absence of livestock or at low levels of livestock use, but with

mule deer (Odocoileus hemionus) concentrations in winter,

crown cover has been measured at 15 to 18 percent (ASNF 4-

4; 45-6U). At high concentrations of mule deer and elk (Cervus

elaphus), crown cover was 11 percent (ASNF 32-64).

Shrubs of black sagebrush in the pinyon-juniper belt and

above are 20 to 30 cm, and less commonly to 50 cm tall. Forbs

are more common in montane stands than in cold desert

stands. However, number of forbs and abundance of forbs

are commonly much lower than in mountain big sagebrush

communities.

The short stature of black sagebrush and lower abundance

of forbs indicates that stands of this shrub have lower value

than stands of mountain big sagebrush for sage-grouse

nesting and winter habitat. The crowns are often in contact

with the ground or close enough to the ground to obstruct

sage-grouse nesting. However, some stands of black sagebrush do have height and crown cover features that meet the

requirements indicated by Connelly and others (2000) for

nesting and winter habitat for sage-grouse. Some windswept ridges supporting short and scattered plants of sagebrush have been found to be highly selected by sage-grouse

as winter-feeding areas (Gullion 1964).

Artemisia papposa

Owyhee Sagebrush ______________

This half-shrub ranges from the northern margin of the

Snake River Plain in Idaho to adjacent Nevada and Oregon.

It is limited in height to 5 to 15 cm, and the woody stems are

commonly only half this height. Features of this plant

indicate little, if any, value for sage-grouse nesting, brood

rearing, or wintering. Where stands of Owyhee sagebrush

are adjacent to stands of big sagebrush, they might provide

strutting habitat and some forage. Herbaceous plants in

Owyhee sagebrush communities can be expected to dry in

early summer, and they likely have low value for sagegrouse in summer.

Artemisia pedatifida

Birdsfoot Sagebrush_____________

This dwarf half-shrub ranges from Wyoming and southwestern Montana to Colorado. It is limited in height to 5 to

15 cm. Value for sage-grouse habitat is likely similar to that

of Owyhee sagebrush discussed above.

Artemisia porteri

Porter Sagebrush _______________

This is another dwarf half-shrub limited to less than 20 cm

tall. It is a narrow endemic of barren clay hills in Fremont

County, Wyoming (Dorn 1992). The habit and habitat of this

plant indicate low value for sage-grouse except where stands

are adjacent to stands of big sagebrush.

Artemisia pygmaea

Pygmy Sagebrush _______________

Pygmy sagebrush is mostly limited to the Great Basin and

the Uinta Basin, mostly in Nevada and Utah, with the

margins of its range reaching into Arizona and Colorado. It

is mostly confined to geologic strata that weather to

semibarrens. It is an indicator of inherently semibarren,

low productive sites. It is sometimes associated with narrowly endemic plants.

Stature of this plant is short enough that it has little, if

any, value as cover for sage-grouse at any season of the year.

It is indicative of sites of inherently low, scattered plants

where rock, pavement, and bare ground dominate the soil

surface. It is often associated with pinyon and juniper where

the presence of trees also limits its value for sage-grouse.

Forage value of pygmy sagebrush for sage-grouse is apparently unknown.

Sites occupied by pygmy sagebrush are not recommended

for forage improvement projects. Reclamation of disturbed

stands can be expected to be slow and difficult. The badland

habitat indicates low potential for cheatgrass to form closed

stands. However, cheatgrass can be expected to grow in the

habitat of pygmy sagebrush.

Artemisia rigida

Scabland Sagebrush, Rigid

Sagebrush, Stiff Sagebrush _______

This sprouting, deciduous shrub ranges from southeastern Washington and northeastern Oregon to western Idaho
USDA Forest Service Proceedings RMRS-P-38. 2005 25

Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

and northwestern Montana where it grows on basalt scablands of the Columbia Basin (Schultz 1986; Wambolt and

Frisina 2002).

Classification

Artemisia rigida/Poa sandbergii (Hall 1973; Hironaka and

others 1983; Tisdale 1986)

Habitat and Capabilities

Scabland sagebrush is the primary woody Artemisia of the

Palouse grasslands (Hironaka and others 1983). It typically

grows on a thin mantle of soil, underlain by basalt bedrock.

The soil is commonly saturated in winter and spring, frequently enough to preclude establishment of other woody

sagebrush species. The scabland habitat supports a relatively sparse cover of vegetation (Hironaka and others 1983).

Hall (1973) listed 232 kg/ha (207 lbs/acre) herbage production, 5 percent surface rock, 18 percent erosion pavement, 20

percent bare ground, and soil depth of 8 to 25 cm with

stoniness commonly of 25 to 60 percent for a community

type in Oregon and Washington.

Cronquist (1994) described scabland sagebrush with a

height of (20) 30 to 60 cm and with capability to sprout from

the roots. This shrub is deciduous and drops all of its leaves

in winter (Hironaka and others 1983).

Stature of the plant indicates moderate value for sagegrouse nesting. However, the scabland habitat indicates

relatively low percent crown cover of sagebrush. Hall (1973)

reported that cover for this sagebrush ranged from 5 to 20

percent. Tisdale (1986) found that foliar cover averaged 12

percent with no other shrubs in high-elevation communities

of scabland sagebrush. He also found 11 species of perennial

forbs occurring on more than one-half of the sites. On lowelevation sites with soils only 7 to 10 cm deep, Tisdale (1986)

found sparse cover of scabland sagebrush and poor representation of forbs. The crowns of scabland sagebrush seem to be

somewhat less spreading and not as dense as mountain big

sagebrush or Wyoming big sagebrush. This also indicates

lower value for nesting habitat for sage-grouse.

The scabland habitat indicates early summer drying of

succulent plants. This feature suggests low value for nesting

and brood rearing of sage-grouse. The deciduous feature of

the plant makes this sagebrush of little value as winter

forage for sage-grouse. In Idaho, scabland sagebrush forms

a mosaic with other sagebrush types (Hironaka and others

1983). In this setting stands of this shrub likely provide

strutting habitat.

The scabland habitat indicates difficulty and low returns

for forage improvement and reclamation projects. Features

of the type that indicate management limitations include

severe moisture saturation during winter and severe frost

heaving (Hall 1973; Hironaka and others 1983).

The sprouting capability of scabland sagebrush indicates

high value of this shrub for reclamation where cheatgrass

and high fire frequencies have altered structure and

function of plant communities. However, high fidelity of

this plant for basalt scablands indicates seedings beyond

its rather narrow ecological amplitude are not likely to

succeed.

Artemisia rothrockii

Rothrock Sagebrush _____________

As treated by Shultz (1986), Rothrock sagebrush is endemic to the southern Sierra Nevada and San Bernardino

Mountains of California where it is found in high-elevation

silt basins and on rocky slopes. This places Rothrock sagebrush, in a strict sense, beyond the range of sage-grouse.

References to Rothrock sagebrush in the Intermountain

West are likely based on specimens of spiked big sagebrush

(A. tridentata ssp. spiciformis), because habit and habitat

features of Rothrock sagebrush are somewhat similar to

those of spiked big sagebrush.

Adaptation to moist conditions, a strong tendency to rootsprout, large heads, and small amounts of pubescence on the

involucres suggest a close relationship to A. cana (Ward

1953). The origin of Rothrock sagebrush is perhaps similar

to that of spiked big sagebrush, but perhaps with Bolander

silver sagebrush, rather than mountain silver sagebrush,

being one of the parent taxa.

Artemisia spinescens

Bud Sagebrush _________________

This is a plant of cold desert areas from Oregon to Montana and south to California and New Mexico. The shrubs

are commonly 5 to 30 cm tall and rarely taller. Bud sagebrush is commonly associated with other desert shrubs, but

occasionally it forms small stands where it is the dominant

shrub. Cold desert shrub communities typically have relatively few forb species, and these are mostly annuals that

flourish in some years and are essentially absent in others,

depending on the amount and season of precipitation. These

communities are commonly of low stature and plant density.

The height and form of bud sagebrush and the communities in which it is found indicate low value for sage-grouse.

Where bud sagebrush is adjacent to stands of big sagebrush,

it likely provides high-value forage for sage-grouse in spring

and early summer. Stands of bud sagebrush might be used

as strutting grounds, particularly where they are adjacent to

stands of big sagebrush.

Artemisia tridentata ssp. parishii

Parish Big Sagebrush ____________

Parish big sagebrush is restricted to the costal ranges and

cismontane region of California. Features of this taxon and

its communities are quite similar to those of basin big

sagebrush. Features ascribed to Parish big sagebrush might

be found at random over much of the range of basin big

sagebrush. Perhaps additional work will indicate A. t. ssp.

parishii should be reduced to synonymy. If a distinct taxon,

Parish big sagebrush is known from outside the range

indicated for sage-grouse.
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Artemisia tridentata ssp. spiciformis

Spiked Big Sagebrush, Snowfield

Big Sagebrush __________________

Plants of this taxon are of apparent hybrid origin with

mountain silver sagebrush and montane taxa of big sagebrush

being the apparent parents. Some variation in populations

of spiked big sagebrush could be a function of different

parent taxa on the big sagebrush side. Features of silver

sagebrush include numerous fine stems, large heads, and

ability to sprout. Features of big sagebrush include lobed

leaves and upland habitat.

Generally there are fewer flower heads per inflorescence

in spiked big sagebrush than is common in the suggested

parent taxa. Flower heads of spiked big sagebrush are often

larger than those of either mountain silver sagebrush or

other taxa of the big sagebrush complex, and in most populations there are more flowers per head. When transplanted

at low elevations, plants of spiked big sagebrush have come

to full flower by the end of May and first of June (McArthur

and Goodrich 1984). This is a feature not shared by either

silver sagebrush or other taxa of big sagebrush.

This taxon has been confused with Rothrock sagebrush

that Shultz (1986) considered endemic to California. References to Rothrock sagebrush outside California, including

Bramble-Brodahl (1978), are based on spiked big sagebrush.

Vegetation types outside California listed under Rothrock

sagebrush are included in the following list for spiked big

sagebrush. Spiked big sagebrush has also been confused

with Vasey big sagebrush. Osterhout’s Artemisiaspiciformis,

based on his type specimen, clearly describes plants with a

few large heasds per inflorescence. Beetle’s concept of Artemisia tridentata ssp. vaseyana forma spiciformis is based, at

least in part, on Vasey or subalpine big sagebrush. Vegetation types of Hironaka and others (1983) listed under spiked

big sagebrush follow Beetle’s concept and are treated in this

paper under Vasey or subalpine big sagebrush.

Spiked big sagebrush is a sprouting, evergreen shrub that

ranges from southern Idaho to western Wyoming and southwestern Montana and south to California, Nevada (Ruby

Mountains), and Colorado. In Utah it is mostly confined to

basic substrates.

Classification

Artemisia tridentata ssp. spiciformis/Elymus trachycaulus

(Tart 1996)

Artemisia tridentatassp. spiciformis/Trisetum spicatum(Tart

1996)

Artemisia tridentata ssp. spiciformis/mountain forb

(Bramble-Brodahl 1978)

Habitat and Capabilities

Habitat of spiked big sagebrush typically includes open

slopes and openings within the coniferous forest belt. Populations of this plant extend upward to treeline and approach

alpine conditions. It is known from 2,680 to 3,350 m elevation in Utah (Welsh and others 1993). It is capable of

withstanding longer duration of snow cover than are other

taxa of big sagebrush. The common name of snowfield big

sagebrush is comparatively well applied to many populations

of this taxon. The capacity to sprout appears to be a factor in

the ability to occupy areas of deeper snow cover.

Some associated species also indicate high elevation, snowpersistent sites. These include slender wheatgrass, mountain brome (Bromus carinatus), and yellowbrush (Chrysothamnus viscidiflorus var. lanceolatus). High elevation and

cold climate indicate high resistance of these communities to

invasion by cheatgrass.

Although height and crown cover of spiked big sagebrush

and some features of spiked big sagebrush communities

indicate high-value habitat for sage-grouse, the elevation of

many populations of this taxon are such that they are not

snow-free until after the nesting season of sage-grouse.

Stands of this sagebrush are typically covered with too much

snow in winter to provide winter habitat. Thus, their value

for sage-grouse can be limited primarily to the summer

season. The abundance of succulent vegetation during much

of the summer suggests potential for high-value summer

habitat. However, sage-grouse are not likely to reach some

of the higher elevation populations of spiked big sagebrush

at any season of the year.

Artemisia tridentata ssp. tridentata

Basin Big Sagebrush ____________

Basin big sagebrush ranges from southern British Columbia to southwestern North Dakota and south to Baja California and northern New Mexico. This taxon rivals, or perhaps

equals, Wyoming big sagebrush in its geographic range of

distribution. However, the extent of area covered is much

less than that covered by Wyoming big sagebrush.

Classification

Artemisia tridentata ssp. tridentata/Bouteloua gracilis

(Johnston 1987)

Artemisia tridentata ssp. tridentata/Chrysothamnus

nauseosus (Johnston 1987)

Artemisia tridentatassp. tridentata/Elymus cinereus (Baker

and Kennedy 1985; Johnston [1987] listed six references

for Colorado and Wyoming.)

Artemisia tridentata ssp. tridentata/Elymus lanceolatus (An

apparent type unpublished, Ashley National Forest.)

Artemisia tridentatassp. tridentata/Elymus smithii (Francis

1983)

Artemisia tridentata ssp. tridentata/Elymus spicatus

(Hironaka and others 1983; Jensen 1989; Nelson and

Jensen 1987)

Artemisia tridentata ssp. tridentata/Festuca idahoensis

(Hironaka and others 1983; Nelson and Jensen 1987)

Artemisia tridentatassp. tridentata/Stipa comata (Hironaka

and others 1983)

Artemisia tridentata ssp. tridentata/Purshia tridentata/

Elymus lanceolatus (Johnston 1987)

Artemisia tridentata ssp. tridentata/Sarcobatus vermiculatus/Elymus smithii (Francis 1983)
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

Habitat and Capabilities

Stands of this shrub are most highly developed on deep

alluvial soils of canyon bottoms and drainage ways of valleys.

Although of alluvial settings, basin big sagebrush is highly

sensitive to flooding and high water tables, and it does better

on well-drained areas, such as old terraces, than it does on

current flood plains with wet-meadow vegetation. Soil texture is often sandy, gravelly, or loamy. Height of basin big

sagebrush plants is commonly 1 to 3 m and occasional

specimens exceed 3 m.

Jensen (1989) listed annual production in one basin big

sagebrush type at 665 kg/ha (593 lbs/acre). Although this

indicates lower production than for many mountain big

sagebrush communities, potential production under irrigation is high. This tall sagebrush was used as an indicator of

productive sites for agriculture during settlement of the

West. Due to the deep alluvial, productive soils and valley

locations, much of the original habitat of basin big sagebrush

has been converted to agricultural land. Many populations

are highly vulnerable to cheatgrass invasion and subsequent displacement by cheatgrass via high fire frequency.

Based on observations and studies on the Tavaputs Plateau of Utah, it appears that basin big sagebrush might take

4 or more decades to achieve 20 percent crown cover following fire where livestock grazing is limited to winter. In this

respect it appears to function more like Wyoming big sagebrush than like Vasey big sagebrush. However, in other

parts of its range, it might recover more rapidly, and especially under spring and summer use by livestock.

Tall stature indicates low value for strutting habitat.

Although stature of this shrub meets the height needs for

sage-grouse nesting, the crowns of the shrubs are often

elevated above the ground as much as 60 to 100 cm or more.

Sage-grouse are likely to avoid stands with this feature

when selecting nest sites. Stands with succulent forbs might

provide moderate value brood-rearing habitat.

Tall stature and valley bottom habitat indicate foliage of

basin big sagebrush would be available to sage-grouse in

most locations in most winters. However, in very tall stands,

the leaves are difficult for sage-grouse to reach. Other

features of some basin big sagebrush habitats appear to be

of low preference by sage-grouse. This includes the bottoms

of narrow stream canyons where the physical features of the

canyon perhaps discourage sage-grouse. Stands in wide

valleys are indicated to be of much higher value to sagegrouse than those confined to narrow canyons.

Artemisia tridentata ssp. vaseyana

Vasey Big Sagebrush, Subalpine

Big Sagebrush, Mountain Big

Sagebrush _____________________

In addition to Artemisia tridentata. ssp. vaseyana, A. t.

var. pauciflora has been recognized as part of the Vasey big

sagebrush complex based on fewer flowers per head and on

distribution and habitat (Goodrich and others 1985).

In general, populations of Vasey big sagebrush are more

common from Washington to Montana and south to Oregon

and Wyoming. Stands of mountain big sagebrush are more

common and more highly developed in the Great Basin,

Colorado Plateau, and central Rocky Mountains. In Wyoming, communities of Vasey big sagebrush are generally at

higher elevation than those of mountain big sagebrush

(Tart 1996). Even at high elevations in Utah, very few stands

of sagebrush are clearly identifiable as typical Vasey big

sagebrush. Critical review of numerous populations is needed

to more fully understand the distribution and to clarify separation of these two taxa. Intermediate populations might be

encountered throughout much of the range of these two taxa.

The Vasey/mountain big sagebrush complex is a landscape dominant in mountains of much of the West from

British Columbia and western Montana south to California

and Colorado. Within the big sagebrush group, the Vasey

sagebrush complex is likely second only to Wyoming big

sagebrush in extent of area covered.

Classification

Distinction between Artemisia tridentata ssp. vaseyana

and A. t. var. pauciflora is rare in literature dealing with

vegetation type classification. This is because publication of

A. t. var. pauciflora (Goodrich and others 1985) followed

much of the sagebrush classification work listed below, and

likely because of reluctance to accept the distinction. Of the

works listed below, only Tart (1996) recognized A. t. var.

pauciflora. Since the distinction was not made in most

classification works, A. t. var. pauciflora is not treated

separately in the following list of vegetation types or in the

discussion that follows. All references to the complex are

lumped under A. t. ssp. vaseyana or Vasey big sagebrush.

Artemisia tridentata ssp. vaseyana/Bouteloua gracilis (Tew

1988)

Artemisia tridentata ssp. vaseyana/Bromus carinatus

(Hironaka and others 1983; Tew 1988)

Artemisia tridentata ssp. vaseyana/Carex geyeri (Hironaka

and others 1983)

Artemisia tridentata ssp. vaseyana/Elymus cinereus (Jensen

1989; Nelson and Jensen 1987)

Artemisia tridentata ssp. vaseyana/Elymus cinereus/Bromus

carinatus (Mooney 1985; Tew 1988)

Artemisia tridentata ssp. vaseyana/Elymus lanceolatus (An

apparent type unpublished, Ashley National Forest.)

Artemisia tridentata ssp. vaseyana/Elymus smithii (Tew

1988)

Artemisia tridentata ssp. vaseyana/Elymus spicatus (Baker

and Kennedy 1985; Bramble-Brodahl 1978; Hironaka and

others 1983; Jensen 1989; Mooney 1985; Mueggler and

Stewart 1980; Nelson and Jensen 1987)

Artemisia tridentata ssp. vaseyana/Elymus spicatus/Poa

fendleriana (Mooney 1985)

Artemisia tridentata ssp. vaseyana/Elymus trachycaulus

(Tart 1996)

Artemisia tridentata ssp. vaseyana/Festuca idahoensis

(Bramble-Brodahl 1978; Francis 1983; Hironaka and others 1983; Jensen 1989; Mooney 1985; Mueggler and Stewart

1980; Nelson and Jensen 1987)

Artemisia tridentata ssp. vaseyana/Festuca idahoensis/E.

spicatus (Tart 1996)

Artemisia tridentata ssp. vaseyana/Festuca thurberi (Francis

1983; Johnston[1987] listed nine references for Colorado.)
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Artemisia tridentata ssp. vaseyana/Festuca scabrella

(Mueggler and Stewart 1980)

Artemisia tridentata ssp. vaseyana/Leucopoa kingii (Johnston

[1987] listed two references for Colorado; this is also found

in Utah.)

Artemisia tridentata ssp. vaseyana/Leucopoa kingii /Koeleria

macrantha (Mooney 1985)

Artemisia tridentata ssp. vaseyana/Stipa comata (BrambleBrodahl 1978; Hironaka and others 1983)

Artemisia tridentata ssp. vaseyana/Stipa richardsonii (Tart

1996)

Artemisia tridentata ssp. vaseyana/Trisetum spicatum (Tart

1996)

Artemisia tridentata ssp. vaseyana/Purshia tridentata (Tart

1996)

Artemisia tridentata ssp. vaseyana/Purshia tridentata/E.

smithii (Johnston [1987] listed three references for this

association in Colorado.)

Artemisia tridentata ssp. vaseyana/Purshia tridentata/F.

idahoensis (Francis 1983)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus (Tart 1996)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus/Bromus carinatus (Jensen 1989; Nelson and

Jensen 1987)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus/E. spicatus (Bramble-Brodahl 1978; Hironaka

and others 1983; Jensen 1989; Nelson and Jensen 1987;

Tew 1988; Tueller and Eckert 1987)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus/E. trachycaulus (Tew 1988; Tueller and Eckert

1987)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus/F. idahoensis (Bramble-Brodahl 1978;

Hironaka and others 1983; Tueller and Eckert 1987)

Artemisia tridentata ssp. vaseyana/Symphoricarpos

oreophilus/Carex geyeri (Hironaka and others 1983)

Habitat and Capabilities

The lengthy list of vegetation types listed above for the

Vasey big sagebrush complex is a reflection of the high

diversity of habitats occupied by this complex. It also reflects

the favorable climate for plant growth within these habitats.

Paralleling the diversity of vegetation types is the number of

associated species, as discussed below and shown in table 3.

Stands of this complex are most highly developed in areas

with about 35 to 75 cm of annual precipitation. Stands on

sites receiving more than 75 cm annual precipitation are

mostly confined to southerly exposures within the aspen and

coniferous forest belts. Within the coniferous belt, stands are

found on sites receiving as much as 149 cm annual precipitation. On drier sites receiving less than 35 cm of annual

precipitation, they often grade into Wyoming big sagebrush

communities.

Florescence tests in Utah show a rather broad hybrid or

mixing zone between Wyoming big sagebrush and Vasey big

sagebrush in the 30 to 36 cm precipitation belt, which in

Utah corresponds rather well with the pinyon-juniper belt

(Goodrich and others 1999a). Where stands of Vasey big

sagebrush occur within the distribution of pinyon-juniper,

they are often found in and mostly above the pinyon-juniper

thermal belt (Goodrich 1981; Goodrich and others 1999a;

Hodgkinson 1989). Within the mixing zone, bitterbrush is

sometimes an associate. Below the pinyon-juniper belt and

within well-marked Wyoming big sagebrush communities,

bitterbrush is rare or lacking.

Big sagebrush of this complex is often indicative of Mollisols

or mollic intergrades with mollic epipedons. In Oregon,

Swanson and Buckhouse (1984) found it on Mollisols and

Wyoming big sagebrush on Aridisols. Jensen (1989, 1990)

listed five community types for Vasey big sagebrush in

northeastern Nevada, all of which were found principally on

Cryoboralls (Mollisols with cold temperature regime). Tart

(1996) found communities of Vasey big sagebrush common

on Cryoboralls on the west flank of the Wind River Mountains, Wyoming.

Plants of the Vasey big sagebrush complex frequently

grow where depth and duration of snow pack is considerably

greater than what is typical for Wyoming big sagebrush

(Meyer and Monsen 1992). Sturges and Nelson (1986) found

Vasey big sagebrush grew where snow depth was greater

than 38 cm, and Wyoming big sagebrush was more common

where snow depth was less than 40 cm.

Compared to those of Wyoming big sagebrush, the ecological settings of Vasey big sagebrush are more productive.

Higher production is indicated as inherent over at least the

past few hundred to thousands of years by the dominance of

Mollisols instead of Aridisols in Vasey big sagebrush communities. Higher production of Vasey big sagebrush compared to Wyoming big sagebrush at present is verified by

comparison of annual growth per unit area. It is also indicated by greater ground cover potential, higher diversity of

understory species, and the potential for greater crown cover

of sagebrush.

Table 3—Comparison of features of Wyoming big sagebrush

communities (ARTRW) from pediments associated with the

Green River and mountain big sagebrush communities

(ARTRP) on the Bishop Conglomerate Formation (from

Goodrich and Huber 2001).

Community feature ARTRW ARTRP

n = 9 n = 6 a

Crown cover of sagebrush (percent) 0–22 0–38

Total number of vascular plant taxa 51 93

Total number of shrub taxa 7 7

Total number of graminoid taxa 13 20

Total number of forb taxa 26 66

Average number taxa/site (alpha diversity) 18 47

Average number of taxa/quadrat (beta diversity) 2.92 11.35

Average number of shrub taxa/site 4 4

Average number of graminoid taxa/site 6 12

Average number of forb taxa/site 7 31

Shrubs with 100 percent consistency 1 3

Shrubs with >49 percent consistency 1 4

Graminoids with 100 percent consistency 1 7

Graminoids with >49 percent consistency 7 12

Forbs with 100 percent site consistency 0 8

Forbs with >49 percent site consistency 4 31

Average ground cover (percent) 58 92

a

n = 6 except for crown cover of sagebrush for which n = 42.
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

A range of annual production of 418 to 2,354 kg/ha (373 to

2,100 lbs/acre) is indicated for Vasey big sagebrush communities (Goodrich and Huber 2001; Harniss and Murray 1973;

Jensen 1989; Tart 1996; Tew 1988). The low end of the range

(418 kg/ha) was reported from near Dubois, Idaho, where

sample sites were likely near the ecotone with Wyoming big

sagebrush. Sites where bluebunch wheatgrass and blue

gramma (Bouteloua gracilis) were indicators generally had

lower production than sites with snowberry (Symphoricarpos

spp.) and slender wheatgrass (Agropyron trachycaulum) as

indicator species.

Ground cover is rarely less than 65 percent, and it is

commonly above 85 percent even under moderate livestock

grazing. At sites in the Uinta Mountains with a history of

about 100 years of livestock grazing, ground cover has been

found at 87 to 95 percent with an average of 92 percent

(Goodrich and Huber 2001). For the Fishlake National

Forest, Tew (1988) considered Vasey big sagebrush communities at potential to have between 7 and 22 percent bare

soil. This indicates ground cover potential of between 78

and 93 percent for that National Forest. Tart (1996) listed

percent bare soil at 6 to 15 percent for 8 plant associations

at late seral condition on the West Flank of the Wind River

Mountains. This indicates potential for ground cover of

about 85 to 94 percent.

Potential for recovery of ground cover following fire is

high. Ground cover was found at potential (90 percent or

higher) within 5 to 7 years postfire on the Ashley National

Forest. This recovery was concurrent with light to moderate

livestock grazing. At a site without livestock (ASNF 5-29),

ground cover increased from 55 to 97 percent at 2 and 6 years

postfire, respectively. However, it is apparent that not all

Vasey big sagebrush sites have potential for ground cover of

over 90 percent. At two adjacent sites on Bare Top Mountain,

Uinta Mountains (ASNF 5-13), ground cover was found at 62

and 85 percent in an area closed to livestock grazing for over

20 years. The site with 62 percent ground cover supported

moderate frequency of rock goldenrod (Petradoria pumila)

and low pussytoes (Antennaria dimorpha). These plants were

lacking on the site with 85 percent ground cover. These and

other plants might be used as indicators of lower potential for

ground cover in Vasey big sagebrush communities.

On the west flank of the Wind River Mountains, Wyoming,

Tart (1996) generally found 10 to 44 vascular plant taxa per

375 m2 plot in associations of the Vasey big sagebrush

complex. He also indicated potential for ground cover was

greater than 80 percent in these associations.

Fisser (1962) found average percent basal area was 12.4

for graminoids and 6.4 for forbs. Although he found 43 forbs

in his study area, only silver lupine (Lupinus argenteus) and

11 other forbs had greater than trace amounts of cover, and

the lupine accounted for 70.1 percent of the forb cover.

Diversity of vascular plants is relatively high in Vasey big

sagebrush communities compared to Wyoming big sagebrush communities. Table 3 compares Vasey big sagebrush

communities on plateau lands of the Bishop Conglomerate

Formation in the eastern Uinta Mountains with Wyoming big

sagebrush communities on gravel pediments at the base of the

Uinta Mountains. This comparison includes a contrast of an

average of 47 and 18 vascular plant taxa per site for Vasey big

sagebrush and Wyoming big sagebrush communities, respectively (Goodrich and Huber 2001).

Average crown cover of Wyoming big sagebrush rarely exceeds 25 percent in areas of over 0.5 ha, while values of greater

than 25 percent are common in Vasey big sagebrush communities. Without fire or other crown-reducing disturbance for

greater than 30 years, crown cover values between 25 and 40

percent are common for Vasey big sagebrush communities in

the Uinta Mountains. On the west flank of the Wind River

Mountains, Tart (1996) found that canopy cover of all shrubs

in mid and late seral communities of Vasey big sagebrush

was between 23 to 40 percent. Crown cover of Vasey big

sagebrush of nearly 50 percent was recorded in recent

studies in Strawberry Valley and West Tavaputs Plateau,

Utah, where values of between 35 and 45 percent are not

uncommon (Goodrich and Huber 2004, unpublished data).

Higher productivity, including greater crown closure of

Vasey big sagebrush communities, supports a higher fire

frequency than for Wyoming big sagebrush communities.

Although Houston (1973) did not specify subspecies of sagebrush for his study area, the location and associated species

indicate Vasey big sagebrush where he suggested a mean

fire frequency of 20 to 25 years.

Miller and Rose (1999) found mean fire intervals of 12 to 15

years for a sagebrush steppe basin in Oregon. Citing four

papers, Miller and others (1999) indicated fire return intervals

usually ranged between 10 and 25 years. This is well within the

range of 10 to 40 years suggested by Winward (1991).

Shorter recovery time for Vasey big sagebrush, compared

to Wyoming big sagebrush, following fire is also indicative of

an ecological history of higher fire frequency. Recovery from

a burn to 20 percent canopy cover can range from 12 years in

the Vasey big sagebrush type to over 40 years in the drier

Wyoming big sagebrush type (Winward 1991). Annual herbage production of sagebrush has returned to preburn levels

within 30 years or less following fire (Harniss and Murray

1973). Review of data from over 40 sites on the Ashley

National Forest shows Vasey big sagebrush returning to

over 20 percent crown cover within 20 years of burning

(Goodrich and others 2004, unpublished data). History of

some of these sites included previous burns or herbicide

applications from which sagebrush recovered prior to the

most recent fire.

Although some species are reduced in density and production for a few years, essentially all plant species of Vasey big

sagebrush ecosystems return to burned areas by seedbanks,

sprouting, or mobile seeds. Big sagebrush is among the least

responsive species to fire, but it generally returns to burned

areas.

The general adaptation of plant species to fire within

Vasey big sagebrush ecosystems is additional evidence of an

ecological history of fire. Also indicative of a high fire

frequency for Vasey big sagebrush communities is the ease

with which these communities are burned under prescribed

conditions. On the Ashley National Forest, fire has spread

though these communities when most other communities

will not carry fire.

In general, Vasey big sagebrush communities demonstrate greater capacity to recover from fire and to resist

cheatgrass dominance or turn into cheatgrass-driven communities than do Wyoming big sagebrush communities. In

the Uinta Mountains, cheatgrass has made little entry into

Vasey big sagebrush communities of plateau lands above

2,450 m elevation. It has made conspicuous advances into
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

these communities on southerly facing aspects of these mountains, and particularly on gradients exceeding 40 percent.

However, even on southerly aspects, native plants have

demonstrated high capacity to recover in the presence of

cheatgrass following fire.

Height and crown cover of Vasey big sagebrush in midand late-seral communities provide high-value nesting habitat for sage-grouse. Distance between the ground and lower

level of the dense, spreading crowns of Vasey big sagebrush

is often highly favorable for concealment of sage-grouse

nests. The high capacity of these communities to produce an

abundance of understory herbage is highly conducive to nest

concealment and for brood-rearing habitat. Abundance and

diversity of forbs and graminoids and succulence of herbaceous plants in summer indicate higher value for sage-grouse

brooding habitat than for many Wyoming big sagebrush

communities.

Deeper snow accumulations than in Wyoming big sagebrush communities indicate lower value for winter habitat.

However, in the presence of tall shrubs or on windswept

sites, high-value winter habitat is provided within some

stands of Vasey big sagebrush. In some cases sage-grouse

are known to summer in Vasey big sagebrush areas (Strawberry Valley, Utah), and migrate to Wyoming big sagebrush

areas for winter (Current Creek near Fruitland, Utah)

(Bambrough 2002).

The differences between communities of Wyoming big

sagebrush and those of Vasey big sagebrush are major.

These differences have implications for restoration projects

and habitat values for sage-grouse and other wildlife. Where

expectations exceed the capability of the land, restoration

projects and other management activities are marked for

disappointment.

Artemisia tridentata ssp. wyomingensis

Wyoming Big Sagebrush _________

Wyoming big sagebrush is a landscape dominant of plains

and basins from eastern Washington to western North

Dakota and south to California and northern Arizona and

New Mexico (McArthur and Sanderson 1999a). This range

includes the Snake River Plain of Idaho, basins of the Great

Basin, and the Intermountain basins and high plains of

Wyoming that grade into the mixed grass prairie.

Classification

Artemisia tridentatassp. wyomingensis/Atriplex confertifolia/

Grayia spinosa/Stipa comata (Baker and Kennedy 1985)

Artemisia tridentata ssp. wyomingensis/Elymus elymoides

(Hironaka and others 1983; Nelson and Jensen 1987)

Artemisia tridentata ssp. wyomingensis/Elymus spicatus

(Baker and Kennedy 1985; Francis 1983; Hironaka and

others 1983; Mueggler and Stewart 1980)

Artemisia tridentata ssp. wyomingensis/Poa secunda

(Hironaka and others 1983)

Artemisia tridentata ssp. wyomingensis/Elymus lanceolatus

(An apparent type unpublished, Ashley National Forest.)

Artemisia tridentata ssp. wyomingensis/Elymus smithii

(Baker and Kennedy 1985; Johnston [1987] listed nine

references from Colorado, Idaho, Montana, North Dakota,

and Wyoming for this association. Although both A. t.

vaseyana and A. t. wyomingensis were listed by Johnston

[1987] for this association, elevation and soils strongly

indicate A. t. wyomingensis is the sagebrush of this association in most areas.)

Artemisia tridentata ssp. wyomingensis/Stipa comata

(Hironaka and others 1983)

Artemisia tridentata ssp. wyomingensis/Stipa hymenoides

(Johnston [1987] listed three references for Colorado.)

Artemisia tridentata ssp. wyomingensis/Stipa thurberiana

(Hironaka and others 1983)

Hilaria jamesii/Artemisia tridentata ssp. wyomingensis

(Everett and others 1980)

Habitat and Capabilities

The list of vegetation types for Wyoming big sagebrush is

only about one-third as long as that for the Vasey big

sagebrush complex. This shorter list is an apparent reflection of the lower diversity of habitats occupied by Wyoming

big sagebrush. It also reflects the more restrictive climate for

plant growth within the habitat of this shrub. Parallel to the

reduced diversity of vegetation types is the reduced number

of associated species, as discussed below and as shown in

table 3.

Soils under Wyoming big sagebrush communities are

typically Aridisols (Barker and McKell 1986; Swanson and

Buckhouse 1984) with restrictive horizons at about 30 to 56

cm (Goodrich 1981; Winward 1983). However, communities

of this plant are also found on soils with alluvial features in

which deposition exceeds horizon development where the

restrictive horizon, if present, is much deeper than indicated

above.

Stands of Wyoming big sagebrush are most highly developed in areas with about 22 to 30 cm of annual precipitation.

They do develop with only 17 to 22 cm of annual precipitation, but in this case they are often confined to drainageways and other local conditions where they grade into cold

desert shrub communities. With about 30 to 36 cm of annual

precipitation they often grade into stands of the Vasey big

sagebrush complex.

Florescence tests in Utah show a rather broad hybrid or

mixing zone between Wyoming big sagebrush and Vasey big

sagebrush within the 30 to 36 cm precipitation belt, which in

Utah corresponds rather well with the pinyon-juniper belt

(Goodrich and others 1999a). Where Wyoming big sagebrush is found within the distribution of pinyon-juniper, it is

often found in and below the pinyon-juniper thermal belt

(Goodrich 1981; Goodrich and others 1999a; Hodgkinson

1989).

Lower annual precipitation indicates less annual production for Wyoming big sagebrush communities than for the

Vasey big sagebrush complex. Lower production potential is

reflected in a lower potential for ground cover. Winward

(1983) noted as much as 25 percent bare ground for Wyoming

big sagebrush communities even under undisturbed conditions. Kindschy (1994) reported 38 percent bare soil and 21

percent rock for a pristine site with 5 to 7.5 percent crown

cover of Wyoming big sagebrush in the Jordan Crater Kipukas

of southeastern Oregon. In Daggett County, Utah, a mean of

55 percent ground cover and 45 percent bare soil was found

for six sites that had been rested from livestock use for
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

greater than 10 years (Goodrich and others 1999b). Rowlands

and Brian (2001) indicated as high as 70 percent bare ground

was found in a relic site in northern Arizona in what

appeared to be a Wyoming big sagebrush community.

Lower potential for production compared to Vasey big

sagebrush communities is also reflected in the lower potential for crown cover of Wyoming big sagebrush. Average

crown cover of Wyoming big sagebrush rarely exceeds 25

percent in areas of over 0.5 ha, while values of greater than 25

percent are common in Vasey big sagebrush communities.

Fisser (1962) found average percent crown cover of big

sagebrush ranged from 17.8 to 27 with an overall average of

22.4 on arid, low-elevation sites on the Owl Creek Range in

Wyoming. In contrast, he found average percent crown cover

of big sagebrush ranged from 13 to 60 with an overall

average of 35.6 on high-elevation, moist sites. The work of

Fisser (1962) preceded the published taxonomic distinction

of Wyoming big sagebrush by Beetle and Young (1965);

however, his description of sites and community features

leaves little doubt that he was distinguishing between Wyoming big sagebrush and the Vasey big sagebrush complex.

On low-elevation, semiarid sites, average crown cover of

big sagebrush of up to 23 percent has been reported by

Anderson and Holte (1981), Goodrich and others (1999b),

Rowlands and Brian (2001), Tuller and Blackburn (1974),

and Winward (1991). Average values of higher than 23

percent were not found in the literature reviewed for this

paper. The high of 27 percent reported by Fisser (1962)

appears to represent the high end for Wyoming big sagebrush. Higher values might be found under some conditions

including alluvial soils without restrictive horizons and

where some moisture is funneled; however, these appear to

be exceptions limited to localized areas.

Winward (1991) suggested Wyoming big sagebrush sites

with the least disturbance had sagebrush crown cover values

of between 8 and 11 percent while those with highest grazing

impacts had sagebrush crown cover that exceeded 20 percent.

Sturges and Nelson (1986) found Wyoming big sagebrush

grew in locations where the snow pack was less than 40 cm

deep and Vasey big sagebrush grew where snow pack was

greater than 38 cm. In this respect, Wyoming big sagebrush

communities are indicted to provide a larger base of winter

habitat for sage-grouse than communities of Vasey big

sagebrush.

In addition to sagebrush-grass communities, Wyoming

big sagebrush forms communities with sparse understory at

lower elevations and especially at the lower end of its

precipitation range. Such communities are particularly common in some geomorphic settings. They are common on the

Green River Formation in Sweetwater County, Wyoming,

where annual precipitation is about 18 to 20 cm, and where

they intergrade into cold desert shrub communities as they

do in other parts of the West. Striking contrasts in understory composition and abundance are common at the interface of materials of the Green River Formation and areas

where this formation is covered with pediment materials

deposited by the Green River. The pediment materials

support grass-rich communities, and the Green River Formation is grass-poor. Wyoming big sagebrush dominates the

overstory on both substrates.

Volland (1976) provided another example of low-producing sagebrush communities of specific geomorphic settings

for the Central Oregon Pumice Zone. Rhyolite was recognized as a major abiotic feature of this community. Although

he did not specify subspecies of big sagebrush, Volland (1976)

recognized a big sagebrush/needlegrass plant community for

which, under good condition, herbage production was approximately168 kg/ha (150 lbs/acre) and bare ground and

pavement ranged from 60 to 85 percent. Sagebrush crown

cover was limited to 10 to 20 percent.

Sparse understory is indicated for some areas of the

Wyoming big sagebrush belt by early explorers of the West.

One of the examples cited by Peters and Bunting (1994) is

from John C. Fremont and illustrates this point. The report,

made in 1843, includes a note made near the mouth of Goose

Creek on the Snake River Plain in which Fremont reported

“the country has a barren appearance, sandy, and densely

covered with the artemisias from the banks of the river to the

foot of the mountains.” Several days later, near the present

site of Twin Falls, Fremont added “there was no grass here,

the soil among the sage being entirely naked.”

Current relic sites also indicate low potential for diversity

and production. In a study of a relic site that included big

sagebrush communities on Fishtail Mesa in northern Arizona, Rowlands and Brian (2001) found crown cover of big

sagebrush and muttongrass averaged 19.2 and 2.6 percent,

respectively. All other species including forbs contributed

less than 1 percent cover each, and comparison of relative

cover with absolute cover in the Rowlands and Brian (2001)

data indicates total absolute cover of all other species was

only 2.4 percent. They listed only 14 forbs for their sagebrush

sites. Eight of the 14 forbs were found in only 12 percent of

their sample sites. Number of forbs/sample site varied from

1 to 6 with an average of 3.75 forbs/site. The sample size in

this study was a line intercept 121.9 m long. Although they

did not specify subspecies of big sagebrush, their site seems

to fit within the climatic zone for Wyoming big sagebrush for

northern Arizona. That their site was considered a relic

without livestock grazing strongly suggests low inherent

capability to produce forbs.

The Arizona location is toward the margin of the range of

Wyoming big sagebrush, and this could be a factor in the

sparse understory found in the relic site of Rowlands and

Brian (2001). However, Marquiss and Lang (1959) reported

similar conditions for two relic sites in Sweetwater County,

Wyoming. On the two relic sites, they found only five forbs

with greater than 2 percent frequency. Common forbs included Hoods phlox, cushion buckwheat (Eriogonum

ovalifolium), and Hooker sandwort (Arenaria hooker). Hooker

sandwort had 74 percent frequency for the two sites. This

cushion plant is an indicator of harsh sites with inherently

high percent bare soil and exposed gravel. Marquiss and

Lang (1959) also noted forbs were more abundant in adjacent areas grazed by livestock than in the relic sites.

Winward (1983) noted relatively few perennial forbs for

Wyoming big sagebrush communities even in undisturbed

conditions. Bunting (1985) reported that Wyoming big sagebrush habitat types have low perennial forb components in

any successional stage. Although Fisser (1962) found 38

forbs in a study area, the total combined average percent

basal area was 2.6, and only Hoods phlox (Phlox hoodii) and

six other forbs were found with greater than trace amounts.

Hoods phlox alone accounted for 69.5 percent of forb cover.

A contrast of Wyoming big sagebrush and Vasey big sage-
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Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

brush communities in the Uinta Mountains indicated 7 and

31 forbs per site, respectively, for these communities

(Goodrich and Huber 2001). This and other comparisons are

made in table 3.

In a relic site in Oregon, two forbs were found at 5.9

percent and 1.7 percent crown cover. However, Wyoming big

sagebrush crown cover was only 5.2 percent at this site

(Kindschy 1994). Tueller and Blackburn (1974) defined an

overstory/understory inverse relationship in which needleand-thread grass decreased from 7.5 percent to 1.4 percent

basal area as big sagebrush increased from 1.3 percent to

13.5 percent crown cover. Similar overstory/understory relationships have been discussed by a number of workers

(Fisser 1986; Rittenhouse and Sneva 1976; Sturges 1983,

1986; Tanaka and Workman 1988; Wambolt and Payne

1986; West and Hassan 1985; Winward 1983). Some Wyoming big sagebrush sites might not have the capability to

provide high percent crown cover of herbaceous plants concurrent with high percent crown cover of sagebrush.

A low perennial forb component is often characteristic for

salt-desert shrub communities (Goodrich 1986). Proximity

of many Wyoming big sagebrush communities to salt-desert

shrub communities is also an indicator that a low forb

component is inherent to Wyoming big sagebrush communities. As Wyoming big sagebrush communities intergrade

into desert shrub communities, it is reasonable to expected

fewer forbs, decreasing percent ground cover, and lower

production. As they grade toward Vasey big sagebrush

communities, it seems reasonable to expect more forbs,

higher percent ground cover, and greater production.

The preceding discussion of overstory/understory relationships could be misleading if it is read to imply that a

vigorous understory, including forbs, is necessary for Wyoming big sagebrush communities to function as sage-grouse

habitat. Sage-grouse populations have been sustained in

forb-poor Wyoming big sagebrush and mountain big sagebrush stands in Sweetwater County, Wyoming, Parker

Mountain, Utah, and other locations. These populations

strongly indicated that the structure and forage provided by

big sagebrush is far more important than any other habitat

component. Features of Wyoming big sagebrush, including

spreading, dense crowns in close proximity to the ground,

appear to be major compensating factors for low presence of

herbaceous plants. This is not to imply that succulent forbs

are not beneficial. However, an abundance of herbaceous

plants will not compensate for lack of sagebrush structure

and sagebrush forage.

The discussion of overstory/understory relationships is

directed to the need for management to be based on inherent

capabilities of the land. If expectations exceed the inherent

capabilities of Wyoming big sagebrush communities, management decisions will be poorly founded, and habitat improvement or restoration projects will fail to achieve desired

results.

Wyoming big sagebrush is commonly 40 to 70 cm tall and

sometimes taller. The crowns are often dense and spreading

with the lower part of the crown in close proximity to the

ground, but not so close as to obstruct sage-grouse nesting.

These features indicate high values for nesting and other

habitat needs of sage-grouse. Potential for crown cover of

many Wyoming big sagebrush communities is within the

range indicated by Connelly and others (2000) to meet the

needs of sage-grouse. Crown cover of stands of Wyoming big

sagebrush often ranges from about 5 to 25 percent. However,

the low percent crown cover (5.2 percent) of sagebrush found

in a relic site by Kindschy (1994) indicates potential for

sagebrush cover can be well below the range recommended

by Connelly and others (2000). Other reports of inherently

low potential for cover are listed above. Also there are many

areas where browsing by wild ungulates, including pronghorn antelope (Antilocapra americana), keeps crown cover

of Wyoming big sagebrush well below the levels most preferred by sage-grouse.

Due to its growth form, height, widespread distribution,

and expansive areas covered, Wyoming big sagebrush is

perhaps the most important plant in the ecology of sagegrouse. Perhaps no other factor is more important to declining populations of sage-grouse than are the changes that

have taken place in the Wyoming big sagebrush system.

Substantial areas have been converted to agriculture. Others have fallen to urban expansion. However, more extensive loss of sagebrush cover is associated with the introduction of annual grasses. In the past few decades, millions of

hectares of Wyoming big sagebrush have been converted to

cheatgrass and other annuals (Billings 1994; Monsen 1994).

Monsen (1994) reported over 1.3 million ha burned between 1979 and 1993 in southern Idaho with much of this in

an ecological setting where cheatgrass is highly competitive.

A table of acres burned from 1979 to 1993 in southern Idaho

(Monsen 1994) shows area burned varying widely from year

to year without an upward trend. In 1999 and 2000, over

121,410 hectares (300,000 acres) burned on lands managed

by the Bureau of Land Management in Idaho. This indicates

large acreages of annual burning will continue. In 1999,

about 68,800 hectares (1.7 million acres) burned in Nevada.

Much of this burning was in sagebrush types that are highly

vulnerable to cheatgrass dominance. The 1999 fire season in

Nevada represents about one-half of the burned acreage

reported by Monsen (1994) for a 14-year period in southern

Idaho. Perhaps the 1999 season in Nevada will prove to be

a rare exception. However, it could also be an indication of

occasional to common fire seasons of the future. Many of the

acres included in annual reports also burned in previous

fires of recent years. The influence of fire is not only size, it

is also frequency. High fire frequency sustains cheatgrass

communities and prevents the return of sagebrush.

Cheatgrass with its associated fire regime becomes the

driver of the system (Billings 1994; Peters and Bunting

1994). This has reduced diversity and eliminated structure

and forage required by sage-grouse.

The displacement of Wyoming big sagebrush by cheatgrass

and its associated fire system over millions of hectares is

likely the most important factor in the future of sage-grouse

in many parts of the West. Reclamation of lands under a

system driven by cheatgrass and fire will be important to the

future of sage-grouse and other sagebrush associated species.

Due to low precipitation inherent in much of the Wyoming

big sagebrush system, reclamation will be difficult with

limited plant materials likely to succeed in reclamation

projects. Hull (1974) evaluated performance of 90 plant

taxa, including many natives, in seedings on rangelands of

southern Idaho. Where annual precipitation was less than

25 cm, only 17 of the 90 taxa rated over 1 on a relative scale

of 1 to 10. Of these 17, only 6 were natives, and none of the
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Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

natives rated over 2.1. Many natives failed to show any

establishment. Although the work of Hull (1974) is over 25

years old, it clearly indicates the difficulty in reclamation of

Wyoming big sagebrush communities. The difficulty inherent

to the low precipitation of many of these communities is

compounded by the highly competitive capability of cheatgrass.

Difficulty of reclamation with a diversity of native plants was

acutely demonstrated by the work of Hull (1974). Later

review of restoration of areas similar to those studied by Hull

(1974) supports Hulls’ work. Monsen (1994) indicated low

potential for successful seedings in areas with less than 250

mm of annual rainfall. To be successful in breaking the high

fire frequency associated with cheatgrass, plant materials

used in restoration projects must be able to compete with

and reduce cheatgrass, and they will be most useful if they

have comparatively low propensity to carry fire (Monsen

1994).

“Hands-off” or passive management is sometimes recommended with an apparent concept that left alone, cheatgrassdominated areas will revert to native sagebrush/grass steppe.

However, this concept is not supported by studies in relic or

natural areas. Kindschy (1994) reported the presence and

increase of cheatgrass in southeastern Oregon’s Jordan

Crater Research Natural Area that has been protected from

human activities including livestock grazing. In Red Canyon

of the Green River, cheatgrass has been found as the most

frequent species where livestock use and other disturbance

have been minimal (Goodrich and Gale 1999). Young and

Tipton (1990) cited two works from southeastern Washington that documented observations of cheatgrass successfully

inserting itself into climax perennial grass/shrub communities that had been protected from fire and grazing for as long

as 50 years. They proposed that the idea of cheatgrass

spreading in a biological vacuum created by excessive grazing may be somewhat misleading or overstated.

Young and Allen (1997) have emphasized that site degradation is not necessary for cheatgrass invasion. In western

Utah, Harper and others (1996) found cheatgrass able to

establish in ungrazed areas in desert shrub communities

where, although native perennials were able to greatly

suppress the size of cheatgrass plants, cheatgrass was able

to maintain a presence by which it could expand upon

disturbance, including gopher mounds.

Austin and others (1986) found cheatgrass present in Red

Butte Canyon of the Wasatch Mountains where livestock

grazing was discontinued in 1905, likely before cheatgrass

reached that area. They reported higher cover values for

cheatgrass in 1983 than for 1935 in Red Butte Canyon. The

Red Butte Canyon study demonstrates increase of cheatgrass

in absence of livestock grazing.

Knight (1994) reported that the cheatgrass problem is not

restricted to land managed for livestock, and he gave an

example of an increase of cheatgrass following fire in Little

Bighorn Battlefield National Monument in southern Montana. He suggested that managing vegetation of a National

Monument to reflect presettlement conditions is a goal that

may be impossible to attain once certain introduced species

become established.

A hands-off approach does not seem to address the capability of cheatgrass. As suggested by Peters and Bunting

(1994), the introduction of annuals, including cheatgrass,

possibly has been the most important event in the natural

history of the Snake River Plain of Idaho since the last glacial

period. The system has changed. A hands-off approach that

anticipates the return of presettlement ecosystems does not

seem to include the reality of the change. Restoration of

sagebrush-grass communities replaced or displaced by

cheatgrass and other introduced plants is difficult. No satisfactory method for controlling weed competition on

large-scale sagebrush restoration projects has been developed (Meyer 1994). Although current technology of dealing

with cheatgrass probably has a long way to go, the future of

much of the Wyoming big sagebrush system will likely be

determined by development and application of restoration

technology. This technology is a focus of these proceedings.

Restoration will include expectations of production, diversity, and potential cover of shrubs and other species. These

expectations can be expected to be associated with costly

disappointments and poorly based management decisions if

inherent as well as induced features of specific sites are not

considered.

Artemisia tridentata ssp. xericensis

Foothills Big Sagebrush, Xeric Big

Sagebrush _____________________

Foothills big sagebrush is known from west central Idaho

where it grows on mesic temperature and xeric moisture

rated soils derived from Columbia River basalt at elevations

between 752 and 1,524 m (Rosentreter and Kelsey 1991).

Morphology and ecology of foothills big sagebrush suggest a

hybrid origin involving basin big sagebrush and Vasey big

sagebrush (Rosentreter and Kelsey 1991).

Classification

Artemisia tridentata ssp. xericensis/Elymus spicatus

(Hironaka and others 1983)

Artemisia tridentata ssp. xericensis/Festuca idahoensis

(Hironaka and others 1983)

Habitat and Capabilities

The habitat and growth form, as indicated above, are

intermediate between basin big sagebrush and Vasey big

sagebrush. Moderate to high values for sage-grouse for

nesting, breeding, and wintering are indicated. Due to drying of herbaceous plants, values for brood rearing in mid and

late summer are likely lower than in higher montane stands

of Vasey big sagebrush. However, brood rearing is common

in big sagebrush areas equally as dry or drier than found in

the range of this taxon.

Artemisia tripartita ssp. rupicola

Wyoming Threetip Sagebrush _____

This shrub is known from east of the Continental Divide

in Wyoming on the Owl Creek Range, Wind River Range,

and the Laramie Range where its elevational range of

dominance is between 1,768 and 2,560 m (Fisser 1962) and

south-central Montana (Wambolt and Frisina 2002).
34 USDA Forest Service Proceedings RMRS-P-38. 2005

Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Classification

Artemisia tripartita ssp. rupicola/Festuca idahoensis

(Johnston [1987] listed references under Artemisia

tripartitafor the Medicine Bow National Forest, Roosevelt

National Forest, and Shoshone National Forest. These

National Forests are within the range of A. t. rupicola.)

Artemisia tripartita ssp. rupicola/Poa secunda (Johnston

[1987] listed two references for this type for the Shoshone

National Forest under Artemisia tripartita. This National

Forest is within the range of A. t. rupicola.)

Artemisia tripartita ssp. rupicola/Stipa comata (Johnston

[1987] listed two references for the Medicine Bow National

Forest under A. tripartita. This National Forest is within

the range of A. t. rupicola.)

Thatcher (1959) found slimstem muhly (Muhlenbergia

filiculmis) associated with A. tripartita ssp. rupicola, but not

with any other sagebrush taxa in his study area in Albany

County, Wyoming.

Habitat and Capabilities

This shrub is found on windswept, shallow, gravelly soils

of foothills and mountains and flats with estimated average

annual precipitation of between 38 and 48 cm (Thatcher

1959). It is associated with a minimum of winter snow

accumulation and is generally not present on sites where

snow tends to drift (Fisser 1962). In addition to the associated grasses listed above under classification, this shrub is

found with blue gramma and fringed sagebrush (Knight

1994).

Height of this somewhat sprawling shrub, excluding the

flowering stems, is seldom as much as 15 cm (Dorn 1992) and

not over 18 cm (Thatcher 1959). Percent crown cover is

generally between 4 and 12, and other shrubs are commonly

of minor cover (Fisser 1962). Fisser (1962) found Hood’s

phlox and three other forbs provided most of the forb cover

in Wyoming threetip sagebrush communities.

Height and crown cover common to this plant indicate

little value for sage-grouse at any time of the year except

when stands of this shrub are adjacent to taller taxa of

sagebrush. Shultz (1986) indicated this shrub is usually on

barren knolls surrounded by well-developed grasslands.

Proximity to big sagebrush is comparatively uncommon.

However, Fisser (1962) found some stands adjacent to big

sagebrush stands. Where adjacent to stands of big sagebrush, stands of Wyoming threetip sagebrush might be

selected for strutting grounds, and the windswept habitat

might indicate some forage value in winter. However, since

the leaves are deciduous, winter value of this shrub is very

low. The sprouting feature indicates high value for reclamation projects.

Artemisia tripartita ssp. tripartita

Idaho Threetip Sagebrush, Tall

Threetip Sagebrush _____________

This deciduous, sprouting shrub ranges from Washington

and adjacent British Columbia south to Elko County, Nevada,

Box Elder and Cache Counties, Utah, and to Teton County,

Wyoming, and the southwestern portion of Montana

(Wambolt and Frisina 2002). Centers of distribution are in

western Washington and east-central Idaho. Much of the

area once covered by threetip sagebrush has been converted

to agricultural land (Schultz 1986).

Classification

Artemisia tripartita ssp. tripartita/Elymus spicatus

(Hironaka and others 1983)

Artemisia tripartita ssp. tripartita/Festuca idahoensis

(Hironaka and others 1983; Mueggler and Stewart 1980;

Johnston [1987] listed six other references that are from

within the range of A. t. tripartita.)

Artemisia tripartita ssp. tripartita/Stipa comata (Hironaka

and others 1983)

Habitat and Capabilities

Idaho threetip sagebrush occurs primarily on fertile volcanic soils (Shultz 1986) that class in a variety of Mollisols or

intergrade with Mollisols (Hironaka and others 1983) on dry

plains and hills at 1,100 to 2,300 m (Cronquist 1994).

Height of Idaho threetip sagebrush is 20 to 100 cm, and the

plant has high capacity to form closed stands with high

percent crown cover. The herbaceous understory can be

fairly productive, but it is sometimes inversely related to

crown cover of the shrubs (Hironaka and others 1983).

The features of Idaho threetip sagebrush communities

indicate high-value habitat for sage-grouse nesting, breeding, and brood rearing. However, due to the deciduous

feature of the shrub, it can be expected to provide little, if

any, forage in winter.

Hironaka and others (1983) noted cheatgrass is generally

not a problem in this type even though the vegetation may

be severely disturbed. The sprouting feature of the shrub

indicates high value for reclamation projects where ability to

recover from fire is important.

References _____________________

Anderson, J. E.; Holte, K. E. 1981. Vegetation development over 25

years without grazing on sagebrush-dominated rangeland in

southeastern Idaho. Journal of Range Management. 34: 25–29.

Ashley National Forest (ASNF). (Unpublished data from permanent study sites on the Ashley National Forest. These studies are

numbered according to the 7.5 minute grid that corresponds to

maps published by U.S. Geological Survey. These studies include

repeat measurements of various vegetation and ground cover

parameters. On file at: Ashley National Forest Supervisors Office, Vernal, Utah).

Austin, D. D.; Urness, P. J.; Riggs, R. 1986. Vegetal change in the

absence of livestock grazing, mountain brush zone, Utah. Journal

of Range Management. 39: 514–517.

Bambrough, D. J. 2002. Greater sage-grouse winter and male and

female summer habitat selection in Strawberry Valley, Utah.

Provo, UT: Brigham Young University. 45 p. Thesis.

Baker, W. L.; Kennedy, S. C. 1985. Presettlement vegetation of part

of northwestern Moffat County, Colorado, described from remnants. Great Basin Naturalist. 45: 747–777.

Barker, J. R.; McKell, C. M. 1986. Differences in big sagebrush

(Artemisia tridentata) plant stature along soil gradients: genetic

components. Journal of Range Management. 39: 147–151.
USDA Forest Service Proceedings RMRS-P-38. 2005 35

Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

Beetle, A. A. 1960. A study of sagebrush, the section Tridentatae of

Artemisia. Bull. 368. Laramie: University of Wyoming, Agricultural

Experiment Station. 83 p.

Beetle, A. A.; Young, A. 1965. A third subspecies in the Artemisia

tridentata complex. Rhodora. 67: 405–406.

Billings, W. D. 1994. Ecological impacts of cheatgrass and resultant

fire on ecosystems in the western Great Basin. In: Monsen, S. B.;

Kitchen, S. G., comps. Proceedings—ecology and management of

annual rangelands; 1992 May 18–21; Boise, ID. Gen. Tech. Rep.

INT-313. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station: 22–30.

Bramble-Brodahl, M. K. 1978. Classification of Artemisia vegetation in the Gros Ventre area, Wyoming. Moscow: University of

Idaho. 126 p. Thesis.

Bunting, S. C. 1985. Fire in sagebrush-grass ecosystems: successional changes. In: Sanders, K.; Durham, J., eds. Rangeland fire

effects—a symposium: 1984 November 27–29; Boise, ID: U.S.

Department of the Interior, Bureau of Land Management, Idaho

State Office: 7–11.

Clary, W. P. 1986. Black sagebrush response to grazing in the eastcentral Great Basin. In: McArthur, E. D.; Welch, B. L., comps.

Proceedings—symposium on the biology of Artemisia and

Chrysothamnus; 1984 July 9–13; Provo, UT. Gen. Tech. Rep. INT200. Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 181–185.

Connelly, J. W.; Schroeder, A.; Sands, A. R.; Braun, C. E. 2000.

Guidelines to manage sage grouse populations and their habitats. Wildlife Society Bulletin. 28: 967–985.

Cronquist, A. 1994. Intermountain flora. Vol. 5. Asteraceae. NY:

New York Botanical Garden. 496 p.

Dorn, R. D. 1992. Vascular plants of Wyoming. Cheyenne, WY:

Mountain West Publishing. 340 p.

Everett, R. L.; Tueller, P. T.; Davis, J. B.; Brunner, A. D. 1980. Plant

phenology in galleta-shadscale and galleta-sagebrush associations. Journal of Range Management. 33: 446–450.

Fisser, H. G. 1962. An ecological study of the Artemisia tripartita

subsp. rupicola and related shrub communities in Wyoming.

Laramie: University of Wyoming. 166 p. Thesis.

Fisser, H. G. 1986. Biology and ecology of sagebrush in Wyoming: II.

Grazing, sagebrush control and forage yield. In: McArthur, E. D.:

Welch, B. L., comps. Proceedings—symposium on the biology of

Artemisia and Chrysothamnus; 1984 July 9–13; Provo, UT. Gen.

Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 303–313.

Fosberg, M. A. 1963. Genesis of some soils associated with low and

big sagebrush complexes in the Brown Chestnut, and Cherozem

Prairie Zones in southcentral and southwestern Idaho. Madison:

University of Wisconsin. 332 p. Dissertation.

Francis, R. E. 1983. Sagebrush-steppe habitat types in northern

Colorado: a first approximation. In: Moir, W. H.; Hendzel, L.,

coords. Proceedings of the workshop on southwestern habitat

types; 1983 April 6–8; Albuquerque, NM. U.S. Department of

Agriculture, Forest Service, Southwestern Region: 67–71.

Freeman, C. D.; Miglia, K. J.; McArthur, E. D.; Graham, J. H.;

Wang, H. 1999. Narrow hybrid zone between two subspecies of big

sagebrush (Artemisia tridentata: Asteraceae): X. Performance in

reciprocal transplant gardens. In: McArthur, E. D.; Ostler, K. W.;

Wambolt, C. L., comps. Proceedings: shrubland ecotones; 1998

August 12–14; Ephraim, UT. Proc. RMRS-P-11. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 15–24.

Goodrich, S. 1981. A floristic study of central Nevada. Provo, UT:

Brigham Young University. 399 p. Thesis.

Goodrich, S. 1986. Vascular plants of the Desert Experimental

Range, Millard County, Utah. Gen. Tech. Rep. INT-209. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 72 p.

Goodrich, S.; Gale, N. 1999. Cheatgrass frequency at two relic sites

within the pinyon-juniper belt of Red Canyon. In: Monsen, S. B.;

Stevens, R., comps. Proceedings: ecology and management of

pinyon-juniper communities within the Interior West; 1997 September 15–16; Provo, UT. Proc. RMRS-P-5. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 69–71.

Goodrich, S.; Huber, A. 2001. Mountain big sagebrush communities

on the Bishop Conglomerate in the Eastern Uinta Mountains. In:

McArthur, E. D.; Ostler, K. W.; Wambolt, C. L., comps. Proceedings: shrubland ecotones; 1998 August 12–14; Ephraim, UT.

Proc. RMRS-P-11. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station: 336–343.

Goodrich, S.; Huber, A. 2004. Studies inventory Ashley National

Forest (Studies 447-3B and 529-10). Unpublished data on file at:

U.S. Department of Agriculture, Forest Service, Ashley National

Forest, Vernal, UT.

Goodrich, S.; Huber, A.; Monroe, B. 2004. Crown cover of sagebrush

and ground cover in post-fire mountain big sagebrush (Artemisia

tridentata var. pauciflora) communities of the Uinta Mountains

and Tavaputs Plateau. Unpublished paper on file at: U.S. Department of Agriculture, Forest Service, Ashley National Forest,

Vernal, UT.

Goodrich, S.; McArthur, E. D.; Winward, A. H. 1985. A new combination and a new variety in Artemisia tridentata. Great Basin

Naturalist. 45: 99–104.

Goodrich, S.; McArthur, E. D.; Winward, A. H. 1999a. Sagebrush

ecotones and average annual precipitation. In: McArthur, E. D.;

Ostler, K. W.; Wambolt, C. L., comps. Proceedings: shrubland

ecotones; 1998 August 12–14; Ephraim, UT. Proc. RMRS-P-11.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Rocky Mountain Research Station: 88–94.

Goodrich, S.; Nelson, D.; Gale, N. 1999b. Some features of Wyoming

big sagebrush communities on gravel pediments of the Green

River in Daggett County, Utah. In: McArthur, E. D.; Ostler, K. W.;

Wambolt, C. L., comps. Proceedings: shrubland ecotones; 1998

August 12–14; Ephraim, UT. Proc. RMRS-P-11. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 159–167.

Graham, J. H.; Freeman, D. C.; Wang, H.; McArthur, E. D. 1999.

Ecological analysis of the big sagebrush hybrid zone. In: McArthur,

E. D.; Ostler, K. W.; Wambolt, C. L., comps. Proceedings: shrubland

ecotones; 1998 August 12–14; Ephraim, UT. Proc. RMRS-P-11.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Rocky Mountain Research Station: 11–14.

Gullion, G. W. 1964. Wildlife uses of Nevada plants. Contributions

toward a flora of Nevada 49. Beltsville, MD: U.S. Department of

Agriculture, Agricultural Research Service, Crops Research Division, National Arboretum. 173 p.

Hall, F. C. 1973. Plant communities of the Blue Mountains in eastern

Oregon and southeastern Washington. R6 Area Guide 3-1.

Portland, OR: U.S. Department of Agriculture, Forest Service,

Pacific Northwest Region. 62 p.

Harniss, R. O.; Murray, R. B. 1973. 30 years of vegetal change

following burning of sagebrush-grass range. Journal of Range

Management. 26: 322–325.

Harper, Kimball T.; Van Buren, Renée; Kitchen, S. G. 1996. Invasion of alien annuals and ecological consequences in salt desert

shrublands of western Utah. In: Barrow, Jerry R.; McArthur, E.

Durant; Sosebee, Ronald E.; Tausch, Robin J., comps. Proceedings: shrubland ecosystem dynamics in a changing environment;

1995 May 23–25; Las Cruces, NM. Gen. Tech. Rep. INT-GTR-338.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 58–65.

Harrington, H. D. 1954. Manual of the plants of Colorado. Chicago,

IL: Sage Books. 666 p.

Hironaka, M. 1963. Plant-environment relations of major species in

sagebrush-grass vegetation of southern Idaho. Madison: University of Wisconsin. 138 p. Dissertation.

Hironaka, M.; Fosberg, M. A.; Winward, A. H. 1983. Sagebrush-grass

habitat types of southern Idaho. Bull 35. Moscow: University of

Idaho, Forest, Wildlife and Range Experiment Station. 44 p.

Hodgkinson, H. S. 1989. Big sagebrush subspecies and management implications. Rangelands. 11: 20–22.

Holmgren, R. C.; Hutchings, S. S. 1972. Salt desert shrub response

to grazing use. In: McKell, C. M.; Blaisdell, J. P.; Goodin, J. R.,

eds. Wildland shrubs—their biology and utilization. Gen. Tech.

Rep. INT-1. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Forest and Range Experiment Station:

153–164.

Houston, D. B. 1973. Wildfires in northern Yellowstone National

Park. Ecology. 54: 1111–1117.
36 USDA Forest Service Proceedings RMRS-P-38. 2005

Goodrich Classification and Capabilities of Woody Sagebrush Communities of Western ...

Hull, A. C., Jr. 1974. Species for seeding arid rangeland in southern

Idaho. Journal of Range Management. 27: 216–218.

Hutchings, S. S.; Stewart, G. 1953. Increasing forage yields and

sheep production on Intermountain winter ranges. Circular 925.

Washington, DC: U.S. Department of Agriculture, Forest Service. 63 p.

Jensen, M. E. 1989. Soil climate and plant community relationships

on some rangelands of northeastern Nevada. Journal of Range

Management. 42: 275–280.

Jensen, M. E. 1990. Interpretation of environmental gradients

which influence sagebrush community distribution in northeastern Nevada. Journal of Range Management. 43: 161–166.

Johnson, K. 1987. Sagebrush types as ecological indicators to

integrated pest management (IMP) in the sagebrush ecosystem

of western North America. In: Onsager, J. A., ed. Integrated pest

management: state-of-the-art in the sagebrush ecosystem. ARS50. Springfield, VA: U.S. Department of Agriculture, Agricultural Research Service: 1–10.

Johnston, B. C. 1987. Plant associations of Region Two. 2nd ed. R2-

Ecol-87-2. Lakewood, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Region. 429 p.

Kindschy, R. R. 1994. Pristine vegetation of the Jordan Crater

Kipukas: 1978–91. In: Monsen, S. B.; Kitchen, S. G., comps.

Proceedings—ecology and management of annual rangelands;

1992 May 18–21; Boise, ID. Gen. Tech. Rep. INT-313. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 85–88.

Knight, D. H. 1994. Mountains and plains: the ecology of Wyoming

landscapes. New Haven, CT: Yale University Press. 338 p.

Marquiss, R.; Lang, R. 1959. Vegetation composition and ground

cover of two natural relic areas and their associated grazed areas

in the Red Desert of Wyoming. Journal of Range Management.

12: 104–109.

McArthur, E. D.; Goodrich, S. G. 1984. Artemisia tridentata ssp.

spiciformis: distribution and taxonomic placement. In: McArthur,

E. D.: Welch, B. L., comps. Proceedings—symposium on the

biology of Artemisia and Chrysothamnus; 1984 July 9–13; Provo,

UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Research Station:

55–57.

McArthur, E. D.; Pope, C. L.; Freeman, D. C. 1981. Chromosomal

studies of subgenus Tridentatae of Artemisia: evidence of autopolyploidy. American Journal of Botany. 68: 589–605.

McArthur, E. D.; Sanderson, S. C. 1999a. Cytogeography and

chromosome evolution of subgensus Tridentatae of Artemisia

(Asteraceae). American Journal of Botany. 86: 1754–1775.

McArthur, E. D.; Sanderson, S. C. 1999b. Ecotones: introduction, scale,

and big sagebrush example. In: McArthur, E. D.; Ostler, K. W.;

Wambolt, C. L. comps. Proceedings: shrubland ecotones; 1998

August 12–14; Ephraim, UT. Proc. RMRS-P-11. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 3–8.

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; 1992 May 18–21; Boise, ID. Gen.

Tech. Rep. INT-313. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 244–251.

Meyer, S. E.; Monsen, S. B. 1992. Big sagebrush germination

patterns; subspecies and population differences. Journal of Range

Management. 45: 87–93.

Miller, R.; Svejcar, T.; Rose, J. 1999. Conversion of shrub steppe to

juniper woodland. In: Monsen, S. B.; Stevens, R., comps. Proceedings: ecology and management of pinyon-juniper communities

within the Interior West; 1997 September 15–16; Provo, UT.

Proc. RMRS-P-5. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station: 385–390.

Miller, R. F.; Rose, J. A. 1999. Fire history and western juniper

encroachment in sagebrush steppe. Journal of Range Management. 52: 550–559.

Monsen, S. B. 1994. Selection of plants for fire suppression on

semiarid sites. In: Monsen, S. B.; Kitchen, S. G., comps. Proceedings—ecology and management of annual rangelands; 1992 May

18–21; Boise, ID. Gen. Tech. Rep. INT-313. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 363–373.

Mooney, M. J. 1985. A preliminary classification of high-elevation

sagebrush-grass vegetation in northern and central Nevada.

Reno: University of Nevada. 123 p. Thesis.

Mozingo, H. N. 1987. Shrubs of the Great Basin. Reno: University

of Nevada Press. 342 p.

Mueggler, W. F.; Stewart, W. L. 1980. Grassland and shrubland

habitat types of western Montana. Gen. Tech. Rep. INT-66.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Forest and Range Experiment Station. 154 p.

Munz, P. A. 1959. A California flora. Berkeley: University of California Press. 1681 p.

Nelson, L. P.; Jensen, M. E. 1987. Sagebrush-grass community

types of the Humboldt National Forest. Elko, NV: U.S. Department of Agriculture, Forest Service, Humboldt National Forest.

80 p.

Padgett, W. G.; Youngblood, A. P.; Winward, A. H. 1989. Riparian

community type classification of Utah and southeastern Idaho.

R4-Ecol-89-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 191 p.

Peters, E. F.; Bunting, S. C. 1994 Fire conditions pre- and postoccurrence of annual grasses on the Snake River Plain. In:

Monsen, S. B.; Kitchen, S. G., comps. Proceedings—ecology and

management of annual rangelands; 1992 May 18–21; Boise, ID.

Gen. Tech. Rep. INT-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 31–36.

Rittenhouse, L. R.; Sneva, F. A. 1976. Expressing the competitive

relationship between Wyoming big sagebrush and crested wheatgrass. Journal of Range Management. 24: 326–327.

Robertson, D. R.; Nielsen, J. R.; Bare, N. H. 1966. Vegetation and

soils of alkali sagebrush and adjacent big sagebrush ranges in

North Park, Colorado. Journal of Range Management. 19: 17–20.

Rosentreter, R.; Kelsey, R. G. 1991. Xeric big sagebrush, a new

subspecies in the Artemisia tridentata complex. Journal of Range

Management. 44: 330–335.

Rowlands, P. G.; Brian, N. J. 2001. Fishtail Mesa; a vegetation

resurvey of a relict area in Grand Canyon National Park, Arizona.

Western North American Naturalist. 61: 159–181.

Sabinske, D. W.; Knight, D. H. 1978. Variation within the sagebrush

vegetation of Grand Teton National Park, Wyoming. Northwest

Science. 52: 195–204.

Shultz, L. M. 1986. Taxonomic and geographic limits of Artemisia

subgenus Tridentatae (Beetle) McArthur. In: McArthur, E. D.;

Welch, B. L., comps. Proceedings—symposium on the biology of

Artemisia and Chrysothamnus; 1984 July 9–13; Provo, UT. Gen.

Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 20–28.

Sturges, D. L. 1983. Long-term effects of big sagebrush control on

vegetation and soil water. Journal of Range Management. 36:

720–765.

Sturges, D. L. 1996. Responses of vegetation and ground cover to

spraying a high elevation, big sagebrush watershed with 2,4-D.

Journal of Range Management. 39: 141–146.

Sturges, D. L.; Nelson, D. L. 1986. Snow depth and incidence of a

snowmold disease on mountain big sagebrush. In: McArthur, E. D.;

Welch, B. L., comps. Proceedings—symposium on the biology of

Artemisia and Chrysothamnus; 1984 July 9–13; Provo, UT. Gen.

Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 215–221.

Summerfield, H. B.; Peterson, F. F. 1971. Soil properties related to

occurrence of big and low sagebrush in northwestern Nevada. In:

Abstracts of papers, 24th annual meeting, Society for Range

Management, Reno, NV. [Place of publication unknown]: Society

for Range Management. Abstract.

Swanson, S. R.; Buckhouse, J. C. 1984. Soil and nitrogen loss from

Oregon lands occupied by three subspecies of big sagebrush.

Journal of Range Management. 37: 298–302.

Tanaka, J. A.; Workman, J. P. 1988. Economic optimum big sagebrush control for increasing crested wheatgrass production. Journal of Range Management. 41: 172–178.

Tart, D. L. 1996. Big sagebrush plant associations of the Pinedale

Ranger District. Pinedale, WY: Bridger-Teton National Forest.

97 p.
USDA Forest Service Proceedings RMRS-P-38. 2005 37

Classification and Capabilities of Woody Sagebrush Communities of Western ... Goodrich

Tew, R. K. 1988. Ecosystem stratification on the Fishlake National

Forest. Richfield, UT: U.S. Department of Agriculture, Forest

Service, Fish Lake National Forest. 177 p.

Thatcher, A. P. 1959. Distribution of sagebrush as related to site

differences in Albany County, Wyoming. Journal of Range Management. 12: 55–61.

Tisdale, E. W. 1986. Canyon grassland and associated shrublands

of west-central Idaho and adjacent areas. Bull. 40. Moscow:

University of Idaho, College of Forestry, Wildlife, and Range

Science. 42 p.

Tueller, P. T.; Blackburn, W. H. 1974. Condition and trend of the big

sagebrush/needle and thread habitat type in Nevada. Journal of

Range Management. 27: 36–40.

Tueller, P. T.; Eckert, R. E., Jr. 1987. Big sagebrush (Artemisia

tridentata vaseyana) and longleaf snowberry (Symphoricarpos

oreophilus) plant associations in northeastern Nevada. Great

Basin Naturalist. 47: 117–131.

U.S. Department of Agriculture. 2001. The plants database, version

3.1 (http://plants.usda.gov). Baton Rouge, LA: U.S. Department

of Agriculture, Natural Resources Conservation Service, National Plant Data Center.

Volland, L. A. 1976. Plant communities of the central Oregon

pumice zone. R-6 Area Guide 4-2. Portland, OR: U.S. Department

of Agriculture, Forest Service, Pacific Northwest Region. 110 p.

Walton, T. P.; White, R. S.; Wambolt, C. L. 1986. Artemisia reproductive strategies: a review with emphasis on plains silver

sagebrush. In: McArthur, E. D.; Welch, B. L., comps. Proceedings—symposium on the biology of Artemisiaand Chrysothamnus;

1984 July 9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 67–74.

Wambolt, C. L.; Frisina, M. R. 2002. Montana sagebrush guide.

Bozeman: Montana Department of Fish, Wildlife, and Parks. 82 p.

Wambolt, C. L.; Payne, G. F. 1986. An 18-year comparison of control

methods for Wyoming big sagebrush in southwestern Montana.

Journal of Range Management. 39: 314–319.

Ward, G. H. 1953. Artemisia section Seriphidium in North America:

a cytotaxonomic study. Contributions from the Dudley Herbarium. 4: 155–205.

Welsh, S. L.; Atwood, D. N.; Goodrich, S.; Higgins, L. C. 1993. A Utah

Flora. 2nd ed. Provo, UT: Brigham Young University Print

Services. 986 p.

Welsh, S. L.; Goodrich, S. 1995. Plant novelties in Lepidium

(Cruciferae) and Artemisia (Compositae) from the Uinta Basin,

Utah. Great Basin Naturalist. 55: 359–362.

West, N. E.; Hassan, M. A. 1985. Recovery of sagebrush-grass

vegetation following wildfire. Journal of Range Management. 38:

131–134.

Winward, A. H. 1980. Taxonomy and ecology of sagebrush in

Oregon. Bull. 642. Corvallis, OR: Oregon State University, Oregon Agricultural Experiment Station. 15 p.

Winward, A. H. 1983. Using sagebrush ecology in wildland management. In: Johnson, K. L., ed. Proceedings of the first Utah shrub

ecology workshop; 1981 September 9–10; Ephraim, UT. Logan:

Utah State University, College of Natural Resources: 15–19.

Winward, A. H. 1991. A renewed commitment to management of

sagebrush grasslands. In: Miller, R. F., ed. Management of

sagebrush steppe. Special Rep. 880. Corvallis: Oregon State

University Agricultural Experiment Station: 2–7.

Winward, A. H.; McArthur, E. D. 1995. Lahontan sagebrush (Artemisia arbuscula ssp. longicaulis): a new taxon. Great Basin

Naturalist. 55: 151–157.

Young, J. A.; Allen, F. L. 1997. Cheatgrass and range science: 1930–

1950. Journal of Range Management. 50: 530–535.

Young, J. A.; Tipton, F. 1990. Invasion of cheatgrass into arid

environments of the Lahontan Basin. In: McArthur, E. D.; Romney, E. M.; Smith, S. D.; Tueller, P. T., comps. Proceedings—

symposium on cheatgrass invasion, shrub die-off, and other

aspects of shrub biology and management; 1989 April 5–7; Las

Vegas, NV. Gen. Tech. Rep. INT-276. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research

Station: 37–40.

Youngblood, A. P.; Padgett, W. G.; Winward, A. H. 1985. Riparian

community type classification of northern Utah and adjacent

Idaho. R4-Ecol-85-1. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Region. 78 p.

Zamora, B.; Tueller, P. T. 1973. Artemisia arbuscula, A. longiloba,

and A. nova habitat types in northern Nevada. Great Basin

Naturalist. 33: 225–24.

Tapertip Hawksbeard
38 USDA Forest Service Proceedings RMRS-P-38. 2005

Clait E. Braun is retired from the Colorado Division of Wildlife and

operates Grouse, Inc., 5572 North Ventana Vista Road, Tucson, AZ 85750

U.S.A., FAX: (520) 529-0365; e-mail: sg-wtp@juno.com. John W. Connelly is

Research Biologist, Idaho Department of Fish and Game, 1345 Barton

Road, Pocatello, ID 83204 U.S.A.; e-mail: JCsagegrouse@aol.com. Michael

A. Schroeder is Upland Bird Research Biologist, Washington Department

of Fish and Wildlife, P.O. Box 1077, Bridgeport, WA 98813 U.S.A.; e-mail:

schromas@dfw.wa.gov

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Seasonal Habitat Requirements

for Sage-Grouse: Spring,

Summer, Fall, and Winter

Clait E. Braun

John W. Connelly

Michael A. Schroeder

Abstract—Sage-grouse (Centrocercus minimus, C. urophasianus)

are dependent upon live sagebrush (Artemisia spp.) for all life

processes across their entire range. This paper describes habitats

used by sage-grouse as documented in the scientific literature. The

leaves of sagebrush are eaten by sage-grouse throughout the entire

year and comprise 99 percent of their winter diets. Spring (late

March through May) habitats are those with intermixed areas of

taller (40 to 80 cm) sagebrush with canopy cover of 15 to 25 percent

and taller (>18 cm) grass/forb cover of at least 15 percent. Sites used

for display have shorter vegetation, frequently few or only short

sagebrush plants, but with taller, more robust sagebrush within 100

to 200 m that is used for escape cover. Nesting cover mimics that

used overall during spring but with clumps of tall (>50 cm), dense

(about 25 percent) live sagebrush and abundant forbs (>10 to 12

percent cover). Early brood rearing areas are those within 200 m

(initial 3 to 7 days posthatch) to 1 km (up to 3 to 4 weeks posthatch)

of nest sites. Forbs and taller (>18 cm) grasses are important for

broods; forbs provide succulent foods, grasses provide hiding cover,

and the grass/forb mixture supports insects used by chicks. Summer

use areas are those with abundant succulent forbs with live, taller

(>40 cm), and robust (10 to 25 percent canopy cover) sagebrush

useful for cover. These areas continue to be used into fall when sagegrouse move to higher benches/ridges where they forage on remaining succulent forbs such as buckwheat (Eriogonum spp.) and switch

to more use of sagebrush leaves. Winter (early December to midMarch) use areas are often on windswept ridges, and south to

southwest aspect slopes as well as draws with tall, robust live

sagebrush. Height (25 to 35 cm) of sagebrush above the surface of

the snow in areas used in winter is important, as is canopy cover (10

to 30 percent). Management of habitats used by sage-grouse should

initially focus on maintaining all present use areas. Practices to

enhance sagebrush habitats to benefit sage-grouse are reviewed, as

is the need to annually monitor sage-grouse numbers along with

systematic monitoring of the health of sagebrush ecosystems.

Introduction ____________________

Sage-grouse (Centrocercus minimus, C. urophasianus)

historically occurred in at least 16 States and three Canadian Provinces (Aldrich 1963; American Ornithologists’ Union

1957; Johnsgard 1973). They have been extirpated in five

States and one Canadian Province (Braun 1998; Connelly

and Braun 1997) and their overall distribution has become

discontinuous (fig. 1). The changes in sage-grouse distribution have been attributed to loss, fragmentation, and degradation of habitats (Braun 1995, 1998; Connelly and Braun

1997), and it is probable that at least one-half of the original

occupied area can no longer support sage-grouse (Braun

1998). Because of the reduced amount of available habitat,

sage-grouse abundance has also markedly decreased with

reported declines of 10 to 51 percent (Connelly and Braun

1997) and as much as 45 to 82 percent since 1980 (Braun

1998). The known decreases in distribution and abundance

have led to concern about stability of sage-grouse populations and the health of sagebrush ecosystems upon which

they depend. Petitions to list sage-grouse under the Federal

Endangered Species Act have been filed for northern sagegrouse (C. urophasianus) and for Gunnison sage-grouse

(C. minimus).

Sage-grouse are dependent upon ecosystems with vast

and relatively continuous expanses of live, robust, taller

sagebrushes (Artemisia spp.) with a strong grass and forb

component. This dependency upon sagebrush, especially the

subspecies of big sagebrush (A. tridentata vaseyana, A. t.

wyomingensis, A. t. tridentata), low sagebrush (A. arbuscula),

black sagebrush (A. nova), silver sagebrush (A. cana), and

three-tip sagebrush (A.tripartita), as well as a variety of less

apparent and abundant species, has been well documented

(Patterson 1952; reviews by Braun and others 1977 and

Connelly and others 2000a). Since the early 1960s, the sagegrouse/sagebrush relationship has focused attention by

Western States and Provinces on the need to maintain

healthy sagebrush-steppe communities over large expanses.

Guidelines for maintenance of sage-grouse habitats were

developed from the scientific literature (Braun and others

1977, completely revised by Connelly and others 2000a) and

promoted by the Western States Sage-Grouse Technical

Committee. The purpose of this paper is to present an overview of the habitat needs of sage-grouse based on the scientific literature, identify the issues that affect maintainance

of useful habitats for sage-grouse, and discuss management strategies to maintain, enhance, and restore habitats
USDA Forest Service Proceedings RMRS-P-38. 2005 39

Seasonal Habitat Requirements for Sage-Grouse: Spring, Summer, Fall, and Winter Braun, Connelly, and Schroeder

for sage-grouse. This paper draws extensively on the published Guidelines to Manage Sage Grouse Populations and

Their Habitats (Connelly and others 2000a).

Habitat Overview ________________

Spring

Timing of spring breeding activities of sage-grouse is

dependent on elevation and amount of persistent snow

cover. Attendance at leks may start in early to mid-March or,

at higher elevations, in early April. Males may attend and

display at leks until late May but most display and mating

activities are greatly reduced by mid-May. Amount and

depth of snow cover greatly influence sage-grouse breeding

activities; thus, snow-free areas are important components

of spring habitat. Habitats used by sage-grouse during the

breeding period are those associated with foraging, leks,

escape, and nesting. Depending upon moisture regimes,

height of sagebrush in used habitats varies from 30 to 80 cm

with canopy cover from 15 to 25 percent (Connelly and others

2000a). Lek sites typically have low amounts of sagebrush

and appear relatively bare, but they may have extensive

cover of low grasses and forbs. Taller, robust live sagebrush

used as escape cover is normally within 100 to 200 m of active

leks. The average distance from a nest to the nearest lek

varies from 1.1 to 6.2 km, and the actual size of the breeding

habitat appears largely dependent on the migratory characteristics of the sage-grouse population as well as distribution

of sagebrush cover with respect to lek location (Connelly and

others 2000a). Habitats selected for nesting are those with

abundant (15 to 30 percent canopy cover) live, taller (30 to 80

cm) sagebrush plants within a community with >15 percent

ground cover of taller (40 to 80 cm) grasses and forbs

(Connelly and others 2000a). Early brood-rearing habitats

(fig. 2) are normally those within 100 m to 1 km of nesting

sites, especially areas with high plant species richness,

moisture, and taller grasses and forbs (Connelly and others

2000a). Adult sage-grouse, while still foraging extensively

on leaves of live sagebrush, eat leaves and flower parts of

forbs during spring, as do chicks (Apa 1998; Drut and others

1994; Dunn and Braun 1986; Klott and Lindzey 1990).

Summer

Habitats used by sage-grouse in summer (early to midJune to mid to late September) are those that provide

Historic

Range

Current

Range

California

Nevada

Arizona

Oregon

Washington

British Columbia Alberta Saskatchewan

North

Dakota

South

Dakota

Nebraska

Wyoming

Kansas

Colorado

Oklahoma

New Mexico Texas

Utah

Montana

Idaho

Figure 1—Historic and current distribution of sage-grouse (map prepared by M. A. Schroeder).
40 USDA Forest Service Proceedings RMRS-P-38. 2005

Braun, Connelly, and Schroeder Seasonal Habitat Requirements for Sage-Grouse: Spring, Summer, Fall, and Winter

adequate forage, especially succulent forbs, and cover useful

for escape. These habitats may include those used for agriculture, especially for native and cultivated hay production,

edges of bean and potato fields, as well as more typical

sagebrush uplands and moist drainages. Taller (>40 cm) and

robust (10 to 25 percent canopy cover) sagebrush is needed

for loafing and escape cover as well as a source of food. Grass

and forb ground cover can exceed 60 percent (hayfields).

Provided moisture is available through water catchments or

from succulent foliage, sage-grouse may be widely dispersed

over a variety of habitats during this period (Connelly and

others 2000a). As late summer approaches, there is movement from lower sites to benches and ridges (fig. 3) where

sage-grouse forage extensively on leaves of sagebrush.

Fall

Fall (late September into early December) is a time of

change for sage-grouse from being in groups of hens with

chicks or males and unsuccessful brood hens to separation

into larger flocks frequently segregated by gender. Some

birds may continue to use lower riparian or hayfield habitats, but there is movement onto higher, frequently northaspect slopes where succulent native forbs, such as buckwheats,

provide green forage. Use of sagebrush leaves for food becomes more common as does use of extensive stands (>20

percent canopy cover) of taller (>25 cm), live sagebrush

(Connelly and others 2000a). Movements can be slow but

there is a general shift toward traditional winter use areas

(Connelly and others 1988).

Winter

Flocks of sage-grouse are somewhat nomadic in early

winter but may remain within chosen areas for periods of

several weeks or more depending upon extent of snow cover

and depth (Beck 1977; Hupp and Braun 1989b). Sagebrush

height (>20 cm, but usually >30 cm, above the surface of the

snow) is important as is the robust (>10 to 30 percent canopy

cover) structure of live sagebrush (Connelly and others

2000a). Sage-grouse use a variety of sites in winter including

windswept ridges with open (10 to 20 percent canopy cover)

(fig. 4) stands of sagebrush to draws with dense (>25 percent

canopy cover) stands. Quality of the snow can be important

because sage-grouse are known to use snow roosts and

burrows (Back and others 1987). Aspect is also important

with south and southwest slopes most used in hilly terrain

(Hupp and Braun 1989b). Leaves of live, vigorous sagebrush

plants provide >99 percent of the foods eaten during the

winter period (early December until early to mid-March)

(Patterson 1952; Remington and Braun 1985; Wallestad and

others 1975). Generally, winter is a time of body mass gain

(Beck and Braun 1978), although severe winter conditions

over prolonged intervals can reduce the amount of area

available for foraging and cover (Beck 1977) and thus affect

body condition (Hupp and Braun 1989a). Overall movement

during winter may be extensive and home ranges can be

large (Connelly and others 2000a). As winter wanes, flocks

of sage-grouse move toward breeding areas that may be

immediately adjacent to or far distant from winter use areas

(Connelly and others 2000a).

Figure 3—Radio-tracking sage-grouse in high-elevation

summer range with a stand of mountain big sagebrush

in the background (photograph by J. W. Connelly).

Figure 2—Sage-grouse brood hen in good quality

Wyoming big sagebrush habitat, North Park,

Colorado (photograph by C. E. Braun).

Figure 4—Sage-grouse winter range in Wyoming big

sagebrush habitat in North Park, Colorado (photograph

by C. E. Braun).
USDA Forest Service Proceedings RMRS-P-38. 2005 41

Seasonal Habitat Requirements for Sage-Grouse: Spring, Summer, Fall, and Winter Braun, Connelly, and Schroeder

Issues _________________________

Decreases in distribution and abundance of sage-grouse

have been ascribed to a complexity of factors (Braun 1987,

1998; Connelly and Braun 1997). The three major causes,

(1) habitat loss (mostly permanent), (2) fragmentation (frequently permanent but reversible at times), and (3) degradation (usually can be corrected), are generally accepted but

the latter two are poorly recognized and understood. Examples of permanent habitat loss include conversion of

sagebrush rangelands to agricultural crops, town and subdivision developments, placement of power plants or surface

mines, and reservoir construction. Fragmentation of habitats occurs with power lines, paved and other high-speed

road development (including maintenance and improvement of farm roads), habitat-type conversion projects, fire,

or any permanent development that reduces the size of

existing habitat patches. Less understood are the impacts of

fences, seasonal use trails, oil and gas wells with surface

pipelines, noise, and so on. Some of these impacts can be

resolved and sage-grouse will reoccupy some formerly disturbed areas (Braun 1987).

Distribution of habitat types useful to sage-grouse is also

important, as these species are habitat specialists using a

variety of areas within a larger landscape mosaic. Thus, not

only is the quantity of sagebrush habitats important, but

also the juxtaposition and quality of those habitats. All

sagebrush habitats are not equal in their acceptability to

sage-grouse, and location of areas used may affect sagegrouse distribution. Size of habitat patches is important and

larger (>30 km2

) is better than smaller, although the spatial

relationships of habitats for sage-grouse are not well understood. Sage-grouse use a mosaic of habitats that is normally

present in sagebrush-steppe because of differences in soils,

moisture, topography, aspect, insect defoliation, wildfires,

and other factors. Sagebrush naturally regenerates as

overmature plants die and seedlings become established.

Use of the term “decadent” for sagebrush is generally inappropriate because it implies that sagebrush communities

are not dynamic with a variety of age classes from seedlings

to overmature. Since most sagebrush communities are resilient and represent a continuum of age classes within a

mosaic of habitats, creation of “edge” to benefit sage-grouse

is rarely needed. Because of human activities, the presence

of too much edge (especially in straight lines) is more

common than too little edge and results in degradation of

sage-grouse habitats.

Sagebrush ecosystems have been managed through a

variety of treatments from domestic livestock grazing, mechanical and chemical clearing or thinning, to use of prescribed fire (Braun 1998). Fire was a natural event in more

mesic sagebrush communities but was infrequent as demonstrated by the lack of resprouting of big sagebrush, black

sagebrush, and low sagebrush. Fire was more common in

areas with three-tip sagebrush and silver sagebrush because both species resprout. Recent research suggests there

is little gain in forage production of grasses and forbs after

fire, because it can take longer than 30 years to return to

preburn conditions (Wambolt and others 2001).

Treatments of sagebrush communities have primarily

been conducted to benefit another treatment (livestock grazing). Use of some treatments has led to plantings of exotic

grasses, invasion of areas by exotic plants, conifer invasion

of sagebrush habitats, and increased fire frequency. Many,

if not most, of these treatments have been applied to improve

rangelands for domestic livestock but have had negative

impacts on sagebrush communities and animals dependent

on them (Braun and others 1976). Further, successive treatments have been applied to landscapes with little understanding of the cumulative effects that may impact both

sagebrush-dependent animals, such as sage-grouse, and the

overall health of the plant community. The impacts of natural

events such as periodic drought are further exacerbated by

human treatments of sagebrush communities. All of these

issues emphasize the need for active protection of habitats

presently used by sage-grouse as well as restoration of

habitats that formerly supported sage-grouse populations.

Sage-Grouse Habitat Management

Strategies ______________________

The objectives of habitat management to benefit sagegrouse, in order of importance, should be (1) to protect and

maintain existing occupied habitats, (2) enhance existing

occupied habitats, (3) restore degraded habitats that still

receive some sage-grouse use, and (4) rehabilitate significantly altered habitats that no longer support sage-grouse.

Strategies to accomplish these objectives should include:

• Vigorous suppression of wildfire.

• Reconsideration of any use of prescribed fire.

• Proper livestock management (including reconsideration of time of grazing, stocking rates, season of use, and

frequency of use).

• Use of nitrogen fertilizer, except in areas infested by

annual weeds.

• Mechanical chopping of sagebrush.

• Fence type and placement.

• Water management.

• Rehabilitation and restoration techniques discussed in

these proceedings.

At times, manipulation of some occupied sage-grouse

habitat may be necessary to enhance the overall quality of a

seasonal range. An example would be removing or reducing

some sagebrush canopy cover in known breeding habitat to

enhance a depleted understory. Removal of 57 percent of

sagebrush cover resulted in a significant decline in a sagegrouse breeding population (Connelly and others 2000b) and

degradation of early brood-rearing habitat (Fischer and

others 1996). More recently, a wildfire that removed about

30 percent of the sagebrush cover in a breeding habitat

resulted in a 60 percent decline in sage-grouse nest success

(Connelly, unpublished data, 1998). Because of this information and the fact that wildfires, drought, and insect

infestations cannot be predicted, any sagebrush removal

efforts should affect a relatively small portion of the occupied

habitat. Connelly and others (2000a) suggested that >80

percent of breeding and winter habitat with vegetative

characteristics necessary for productive sage-grouse habitat

should remain intact to adequately provide for the needs of

sage-grouse. However, an even greater percentage should

be protected if sage-grouse populations are declining or

the population status is unknown. All proposed habitat
42 USDA Forest Service Proceedings RMRS-P-38. 2005

Braun, Connelly, and Schroeder Seasonal Habitat Requirements for Sage-Grouse: Spring, Summer, Fall, and Winter

manipulations should carefully consider the current condition of habitat, status of the sage-grouse population, and

likely outcome of the vegetation treatment, including recovery time necessary for the area to again provide adequate

habitat for sage-grouse nesting and early brood rearing.

Acknowledgments ______________

We thank S. B. Monsen for inviting our participation in

the symposium that led to these proceedings. We further

thank all managers and researchers who have contributed

to the scientific literature and our understanding of sagegrouse and their use of habitats. Much of our knowledge was

gained through research supported through Colorado (CEB),

Idaho (JWC), and Washington (MAS) Federal Aid to Wildlife

Restoration Projects. This is a contribution from the Western States/Provinces Sage and Columbian Sharp-Tailed

Grouse Technical Committee.

References _____________________

Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae.

Journal of Wildlife Management. 27: 529–545.

American Ornithologists’ Union. 1957. Check-list of North American birds. 5th ed. Baltimore, MD: Lord Baltimore Press. 691 p.

Apa, A. D. 1998. Habitat use and movements of sympatric sage and

Columbian sharp-tailed grouse in southeastern Idaho. Moscow,

ID: University of Idaho. 199 p. Dissertation.

Back, G. N.; Barrington, M. R.; McAdoo, J. K. 1987. Sage grouse use

of snowburrows in northeastern Nevada. Wilson Bulletin. 99:

488–490.

Beck, T. D. I. 1977. Sage grouse flock characteristics and habitat

selection in winter. Journal of Wildlife Management. 41: 18–26.

Beck, T. D. I.; Braun, C. E. 1978. Weights of Colorado sage grouse.

Condor. 80: 241–243.

Braun, C. E. 1987. Current issues in sage grouse management.

Proceedings of the Western Association of State Fish and Wildlife

Agencies. 67: 134–144.

Braun, C. E. 1995. Distribution and status of sage grouse in

Colorado. Prairie Naturalist. 27: 1–9.

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.

Fernleaf biscuitroot

Braun, C. E.; Baker, M. F.; Eng, R. L.; Gashwiler, J. S.; Schroeder,

M. H. 1976. Conservation Committee report on the effects of

alteration of sagebrush communities on the associated avifauna.

Wilson Bulletin. 88: 165–171.

Braun, C. E.; Britt, T.; Wallestad, R. O. 1977. Guidelines for

maintenance of sage grouse habitats. Wildlife Society Bulletin. 5:

99–106.

Connelly, J. W. 1998. Unpublished data on file at: Idaho Department of Fish and Game, Pocatello, ID.

Connelly, J. W.; Braun, C. E. 1997. Long-term changes in sage

grouse Centrocercus urophasianus populations in western North

America. Wildlife Biology. 3: 229–234.

Connelly, J. W.; Browers, H. W.; Gates, R. J. 1988. Seasonal

movements of sage grouse in southeastern Idaho. Journal of

Wildlife Management. 52: 116–122.

Connelly, J. W.; Reese, K. P.; Fischer, R. A.; Wakkinen, W. L. 2000b.

Response of a sage grouse breeding population to fire in southeastern Idaho. Wildlife Society Bulletin. 28: 90–96.

Connelly, J. W.; Schroeder, M. A.; Sands, A. R.; Braun C. E. 2000a.

Guidelines to manage sage grouse populations and their habitats. Wildlife Society Bulletin. 28: 967–985.

Drut, M. S.; Crawford, J. A.; Gregg, M. A. 1994. Brood habitat use

by sage grouse in Oregon. Great Basin Naturalist. 54: 170–176.

Dunn, P. O.; Braun, C. E. 1986. Summer habitat use by adult female

and juvenile sage grouse. Journal of Wildlife Management. 50:

228–235.

Fischer, R. A.; Reese, K. P.; Connelly, J. W. 1996. An investigation

on fire effects within xeric sage grouse brood habitat. Journal of

Range Management. 49: 194–198.

Hupp, J. W.; Braun, C. E. 1989a. Endogenous reserves of adult male

sage grouse during courtship. Condor. 91: 266–271.

Hupp, J. W.; Braun, C. E. 1989b. Topographic distribution of sage

grouse foraging in winter. Journal of Wildlife Management. 53:

823–829.

Johnsgard, P. A. 1973. Grouse and quails of North America. Lincoln,

NE: University of Nebraska Press. 553 p.

Klott, J. H.; Lindzey, F. G. 1990. Brood habitats of sympatric sage

and sharp-tailed grouse in Wyoming. Journal of Wildlife Management. 54: 84–88.

Patterson, R. L. 1952. The sage grouse in Wyoming. Denver, CO:

Sage Books, Inc. 341 p.

Remington, T. E.; Braun, C. E. 1985. Sage grouse food selection in

winter, North Park, Colorado. Journal of Wildlife Management.

49: 1055–1061.

Wallestad, R. O.; Peterson, J. G.; Eng, R. L. 1975. Foods of adult sage

grouse in central Montana. Journal of Wildlife Management. 39:

628–630.

Wambolt, C. L.; Walhof, K. S.; Frisina, M. R. 2001. Recovery of big

sagebrush communities after burning in south-western Montana. Journal of Environmental Management. 61: 243–252.
USDA Forest Service Proceedings RMRS-P-38. 2005 43

Bruce A. Roundy is Professor of Range Science, Department of Integrative

Biology, 401 WIDB, Brigham Young University, Provo, UT 84602.

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Plant Succession and

Approaches to Community

Restoration

Bruce A. Roundy

Abstract—The processes of vegetation change over time, or plant

succession, are also the processes involved in plant community

restoration. Restoration efforts attempt to use designed disturbance, seedbed preparation and sowing methods, and selection of

adapted and compatible native plant materials to enhance ecological function. The large scale of wildfires and weed invasion requires

large-scale approaches to restoration. Practices and equipment

from traditional rangeland revegetation are being adapted to establish diverse, native communities. The challenge is to meet the

establishment requirements of different species and to create weedresistant plant communities.

Introduction ___________________

In the past, range scientists developed range improvement techniques directed mainly at controlling unpalatable

woody species and establishing forage grasses for livestock

and erosion control, but also to establish plants critical for

big game habitat (Roundy 1996). Our goals now are to

restore functional, diverse native plant communities. The

processes of restoration are the same processes that operate

in plant succession. Understanding these processes can help

us develop realistic techniques and goals for large-scale

restoration. I will briefly review concepts and processes of

plant succession and discuss associated aspects of community restoration.

Models of Succession __________

Plant succession is the change in vegetation that occurs

over time after fire, heavy grazing, flooding, or other natural

or human-related disturbances. Secondary succession occurs when the land retains some residual soil and biological

components from the plant and animal communities that

existed before the disturbance (Barbour and others 1998).

Primary succession occurs on new substrates, such as on a

newly formed volcanic island. Two major views of this

process were taught by Clements (1916) and Gleason (1926).

In Clement’s model, vegetation changes from pioneer species through a series of predictable communities or seres,

which replace each other in order until a final stable or

climax community dominates the site. This model is said to

be linear (always follows the same order) and deterministic

(is predictable). On the other hand, Gleason suggested that

vegetation change after disturbance was a function of the

kinds of plants involved and their characteristics relative to

the disturbance. More recently, ecologists have recognized

that features of both models may describe what actually

happens. State and transition models that allow for multiple

steady states of vegetation, with different probabilities of

transition or change from one state to another (Westoby and

others 1989), have been proposed. These models work better

with the current recognition that some disturbances, such as

fire, have a natural frequency and play a major role in shaping

many upland plant communities. Similarly, seasonal flooding

shapes riparian communities (Middleton 1999).

Clements (1928) identified the processes of succession as

nudation (disturbance), migration (movement of new seeds

or other plant propagules to the site), ecesis (plant establishment), interaction (sorting out of species that establish),

reaction (the effects of the successful species on the environment), and stability. These processes correspond to revegetation and restoration principles and practices (table 1).

Although all of these successional processes may be active in

most systems, some are more controlling for some systems

Table 1—Plant successional processes that correspond to restoration

and revegetation principles and practices.

Process Principle or practice

Disturbance Site potential after disturbance,

designed disturbance to control

undesirable plants

Dispersal, migration, Sowing sufficient germinable seed,

 residuals preempting resources from or

controlling residual propagules of

undesirable species, renovation to

maintain or stimulate residual

desirable species

Establishment Seedbed preparation and sowing to

maximize germination and seedling

establishment; selecting adapted

plant materials

Interaction/reaction Selecting ecologically functional,

compatible plant materials for mixed

communities that are weed resistant

Stabilization Restoring disturbance regimes and

management strategies that favor

ecological function
44 USDA Forest Service Proceedings RMRS-P-38. 20055

Roundy Plant Succession and Approaches to Community Restoration

and biomes, while others are more controlling for other

biomes (Chambers and others 1992). For example, historically, fire is a major controlling disturbance in grasslands,

but not arid deserts, where drought is most operative.

Interaction (especially competition) and reaction are particularly operative in forest systems, but not as operative in

tundra or desert systems where the harsh environment may

result in fewer highly adapted species.

The state and transition model has been used to describe

successional processes in the sagebrush system (Westoby

and others 1989). Continued heavy spring grazing moves a

sagebrush/bunchgrass system to one dominated mainly by

big sagebrush (Artemisia tridentata). Introduction of annual

weeds like cheatgrass (Bromus tectorum), and the attendant

increased fire frequency holds this system in cheatgrass

dominance unless major inputs in weed control and revegetation move it to another state. The cheatgrass-dominated

system has little likelihood of transitioning to another state

on its own because the rapidly maturing cheatgrass provides

a fine and well-distributed fuel over a long fire season. It is

adapted to establishment after fire, while other species

cannot survive the 3- to 5-year frequency that can occur; so

the system is stuck in a stable, but not highly desirable state.

Reasons that systems can be held in a stable steady state

include (1) frequent or severe disturbances, such as fire, or

heavy, continuous grazing; (2) establishment inertia, or lack

of establishment associated with harsh environmental conditions, such as in arid deserts where establishment occurs

only in unusually wet years; or (3) competitive exclusion

where shrubs or trees that are highly competitive, such as

pinyon and juniper (Pinus edulis, P. monophylla and

Juniperus osteosperma, J. occidentalis), eventually dominate in the absence of disturbance such as fire. Another

example of the latter reason is the replacement of aspen

(Populus tremuloides) by conifers in the absence of fire.

Changes in the disturbance regime interact with the

environmental context of a plant community to result in

transitions or the lack of transitions to other states. For

example, Harper (1985) provides evidence that could be used

to suggest that lack of fire on acidic soils, such as those in the

Uinta Mountains, may result in conifer replacement of

aspen much sooner than on the calcareous limestone soils of

the Wasatch Mountains, Utah. Damming of rivers and

streams controls spring flooding, a process that is necessary for dispersal and establishment of cottonwood (Populus)

and other riparian species (Middleton 1999; Stromberg and

others 1991). This flooding is also essential for the erosion,

deposition, and sediment transport functions of the stream

that result in the natural geomorphologic features that are

necessary to support riparian plant communities.

Lack of fire in communities once dominated by sagebrush

and bunchgrass can result in invasion and dominance by

pinyon and juniper (Tausch 1999). On deeper alluvial or

more fertile soils, tree canopies expand until they touch

while the understory vegetation and seed bank die out.

These communities are then susceptible to catastrophic

crown wildfires and subsequent weed invasion. On shallower soils invaded by pinyon and juniper, resources are too

limited for tree canopies to touch, but loss of understory

vegetation and subsequent wind and water erosion of

interspaces may still degrade the site (Roundy and Vernon

1999).

Transitions to states of much reduced biotic and physical

function are said to have passed a biotic or physical threshold, after which a transition back to the previous state is

highly unlikely without major intervention (Whisenant 1999).

Such thresholds are called irreversible because natural

processes alone are insufficient to move them back to the

prethreshold state. Susceptibility to such thresholds depends on the environmental context and past management

of the site and plant community. For example, sagebrush

communities that lack a good understory of perennial grasses

pass a biotic threshold into cheatgrass dominance after fire

more easily than those with a good perennial grass understory that survives the fire. Invasion and dominance of

pinyon and juniper on a site of high erosion potential (higher

slopes, finer textured soils, and more frequent occurrence of

intense summer thundershowers) may result in major erosion and passing of a physical threshold, while such invasion

on sites of low erosion potential may not (Davenport and

others 1998).

Management to avoid passing biotic and physical thresholds is much preferred and less costly than attempting

restoration after passing these thresholds. For example, use

of fire or mechanical treatments to control pinyon and

juniper before the understory vegetation or soil is lost, or

grazing management to maintain a good perennial understory in sagebrush communities is less costly and risky than

attempts to restore these communities after they have

passed degrading thresholds. Nevertheless, many of our

landscapes have already passed such thresholds and now

require major intervention. Restoration after crossing a

biotic threshold requires control of the dominating vegetation and revegetation with more ecologically functional and

desirable species. Restoration to some historic, natural plant

community after passing a physical threshold may not be

possible at all, requiring that we set our goals as restoration

of ecological function, rather than historic composition. For

example, our goals on an eroded site might be to establish a

persistent perennial plant cover to hold the remaining soil in

place, rather than risk additional soil erosion by attempting

to establish a diverse, native community that may no longer

be adapted to the degraded conditions of the site. On riparian areas, biotic restoration may be very difficult without

restoring the physical disturbance regime or seasonal flooding that drives the biotic responses.

Environmental Context of the

Sagebrush Systems ____________

The environmental context of both succession and restoration efforts has an overriding influence on the outcome.

Every restoration project requires characterization of the

site in order to determine potential for success, species

selection, and seeding methods. Two major sagebrush systems are recognized across the Western United States (West

1983a,b). Sagebrush steppe is north of the drier Great Basin

and Colorado Plateau sagebrush systems and has more

potential for sagebrush renovation and revegetation success

than those drier systems to the south. Mountain big sagebrush (Artemisia tridentata ssp. vaseyana) communities

with higher precipitation have more potential for revegetation
USDA Forest Service Proceedings RMRS-P-38. 2005 45

Plant Succession and Approaches to Community Restoration Roundy

than the lower Wyoming big sagebrush (Artemisia tridentata

ssp. wyomingensis) communities. Black sagebrush (Artemisia nova) communities on shallow soils are more difficult to

revegetate than many Wyoming big sagebrush sites. Higher

elevation pinyon-juniper communities have higher precipitation than lower elevation Wyoming big sagebrush communities, and are therefore not only easier to successfully

revegetate with desirable species, but may also be more

susceptible to invasion of more mesic weeds such as the

knapweeds (Centaurea spp.). Salt-desert shrub communities are especially hard to revegetate due to low precipitation

and fine-textured, saline-alkaline soils that can flow and

crust after disturbance and wetting. In general, directseeding revegetation is risky with annual precipitation <250

mm, and much less risky at >400 mm. Between these ranges,

soils, species selection, seedbed preparation and sowing

methods, but especially precipitation during and following

the year of seeding greatly affect success.

Designed Disturbance:

Residuals _____________________

Just as natural disturbances, such as fire and flooding,

free up resources for new colonization, revegetation or restoration requires a designed disturbance to reduce undesirable plant populations prior to planting (Sheley and others

1996). Methods of undesirable plant control include biological, mechanical, chemical, and fire. Biological control works

best when used in a program of integrated weed control that

employs other methods to greatly reduce weed populations.

Concurrent or subsequent biological control can then work

well to contain weeds. Herbicides have the potential for

greatest control of specific undesirables. Mechanical methods have the versatility of configuring the control across the

landscape in patterns to maximize wildlife benefits by providing cover, edge, and vegetation linkages where desired.

Mechanical methods have less risk of treating nontarget

areas than do fire, which can get away, or herbicides, which

may drift or move with soil in wind or water erosion. Smooth,

Ely (railroad rails attached perpendicular to the chain), or

Sagar (rails attached parallel to the chain) anchor chains

that produce less to more soil disturbance can be chosen to

control large areas of nonsprouting pinyon and juniper trees.

Chaining after fire and broadcast seeding helps cover seeds.

Broadcast seeding without chaining can result in weed

dominance (Ott and others 2003).

The goal of designed disturbance may be to retain components of the original community. For example, sagebrush

communities may be treated to rejuvenate older shrubs and

control enough of them to establish a more diverse and

productive herbaceous understory. Chaining and one-way

harrowing may kill about half of the sagebrush in a stand

treated for renovation, while discing or two-way harrowing

will kill 70 to 90 percent of the sagebrush (Summers and

others 2003). This designed disturbance for renovation is

equivalent to the successional importance of residuals after

a disturbance. Designing plant control to remove undesirables and leave some or most of the desirable plants requires

knowledge of plant characteristics such as regeneration

potential in relation to the kind of disturbance and location

of growing points or seed survival.

Dispersal, Migration, Establishment:

Sowing Sufficient Pure Live Seed in

Seedbeds Prepared to Maximize

Establishment _________________

The restoration equivalent of the successional process of

dispersal is direct seeding or transplanting. Plant communities not dominated by desirable plants will require sowing of

desirable plants after designed disturbance or wildfire to

restore vegetation, ecological function, and value. The largescales of our weed-dominated landscapes and burned areas

require direct seeding for restoration or fire rehabilitation.

Since weed seeds are often in the seed bank or are adapted

to wind or mechanical dispersal to a site, sowing of desirable

plants to reduce weed invasion or reestablishment is necessary. Methods of sowing also employ methods of seedbed

preparation to place seeds in the seedbed where their requirements for germination and seedling establishment will

be met. This can be a challenge when seeding species of

different seed sizes and shapes into the highly variable

seedbeds and soils of wildlands. However, if seeds are not

placed where their establishment requirements are met,

plants will not establish.

Seedbed Preparation and Sowing

Methods

Common methods of large-scale sowing of rangelands

include drilling and broadcasting seeds. With both methods,

the goal is to bury seeds, but to place them at the best depth

for their size. Species with smaller seeds like sagebrush or

kochia (Kochia prostrata) may establish best when broadcast, then firmed into the surface by a rubber-tired

cultipacker. The challenge for these seeds is to firm them in,

but to not bury them deeper than a few millimeters. Sagebrush seeds can also be seeded through a rangeland drill

using a trashy seed box with a pick-wheel inside to force the

seed into the seed tubes. To avoid competition with sown

grasses, sagebrush is commonly seeded separately in its own

rows, and the seed tubes are pulled to let seed fall on top of

the ground and avoid excessive burial. A concern for seeding

sagebrush this way is that seeds may not be anchored to the

surface and may blow away.

Grasses generally establish best when drilled in the fall.

Larger seeds such as those of Indian rice grass (Achnatherum

hymenoides) can be sown deep (2 to 5 cm) in sandy soils,

while most grass species should be sown 1 to 2 cm deep.

Depth bands on the disks of a rangeland drill are used to

prevent excessive seed burial on sandy soils. Newer drills

are now available that provide better control of seed placement than the standard rangeland drill. These drills should

be tested for success with a range of species on different sites

and soils.

Where topography or surface debris makes it impossible to

pull a drill across the landscape, or where small seeds can be

firmed into the soil, broadcasting is used. The best example

of large-scale broadcasting is for fire rehabilitation in

burned pinyon-juniper woodlands. Typically seeds are

broadcast from a whirlybird seeder suspended from a

helicopter or broadcast from a venturi-type seeder on a fixed
46 USDA Forest Service Proceedings RMRS-P-38. 20055

Roundy Plant Succession and Approaches to Community Restoration

wing aircraft. Sites are oneway chained to cover seed after

broadcasting. Broadcasting seeds without chaining or some

other form of seed coverage results in lack of revegetation

success and weed invasion (Ott and others 2003). Larger

seeds, such as those of four-wing saltbush (Atriplex canescens)

or bitterbrush (Purshia tridentata), can be sown while chaining. These seeds are sown from a dribbler box attached above

the tracks of the two caterpillar tractors pulling the chain.

The seeds fall out of the box onto the top of the track and are

buried in the track imprints. Smaller seeds of sagebrush or

rabbitbrush (Chrysothamnus spp.) should not be seeded this

way because they will be buried too deep.

It is a challenge to broadcast seed mixtures of grass, forb,

and shrub seeds with different seed shapes and sizes. Continuous mechanical stirring of seeds mixed with trashy

seeds such as those of sagebrush is necessary to provide

adequate seed flow from broadcast seeders. The different

seed burial requirements of seed mixtures make it difficult

to maximize establishment for any one species. Chaining or

other post-broadcast seed coverage techniques, such as

railing, cabling harrowing, or imprinting, will probably best

help establish diverse mixtures when done soon after broadcasting on soils where a wide range of micro topographically

diverse safe sites will be created. Determining the advantages of different methods requires experimental comparisons for different sites, species, and methods.

Various methods of improving the seedbed environment

have been developed over the years. These include furrowing, imprinting, aerating, or otherwise configuring the seedbed to create safe sites or locations for seeds that favor their

germination and establishment. The idea is to bury seeds at

the proper depth for emergence, but to increase the time of

available water, reduce salinity, moderate temperature, or

otherwise maximize favorable environmental conditions for

establishment. The success of these methods depends on the

soil, seeded species, and precipitation after seeding. Some

methods such as drilling and imprinting can result in excessive seed burial on sandy soils, or lack of sufficient burial on

heavy-textured or compacted soils. Various methods of seedbed enhancement and sowing should be compared experimentally across a range of sites and with a variety of species

in order to make best recommendations for specific sites.

Seedbed enhancement may increase seedling establishment

on average to moderately wet years, but does not ensure

establishment on dry years (Winkel and Roundy 1991).

Seeding Rates

Seeding sufficient germinable seed of adapted species

requires an understanding of germination characteristics as

well as adaptability of candidate species. Traditional rangeland revegetation guidelines recommend sowing 5.4 to 8.9

lbs/acre (6 to 10 kg/ha) of pure live seed of grass species

known to have a fairly wide range of adaptability. These

recommendations have proven successful for introduced

grasses, but additional considerations are needed to successfully sow native species. Pure live seed is the amount of

viable seed in a bag of seeds. It can be expressed as a

percentage of the total weight of viable seeds, plus other

matter such as seed parts, weed seeds, and nonviable seeds.

Viability, or whether the seed is dead or alive, can be

determined by a tetrazolium chloride solution or TZ test,

where the active dehydrogenase enzyme in live seeds results

in a red staining. This test does not determine germinability.

Dormant seeds are viable but not germinable until dormancy is broken by artificial means or by specific environmental conditions. State seed testing laboratories determine germination percentages at temperature, light, and

other incubation specifications generally known to maximize germination for a particular species. Some species may

also be subjected to pretreatments such as seed coat scarification or chilling prior to incubation to maximize germination. When both seed viability and germination are tested,

seed tags may bear germination and hard seed (viable but

dormant) percentages.

Bulk seeding rates are calculated by dividing the recommended pure live seeding rate by the pure live seed percentage. For large-scale fire rehabilitation projects, the Bureau

of Land Management (BLM) typically contracts for lots of

seed specifying at least a 80 percent germination for grasses,

or lesser percentages for some species, such as sagebrush,

that are hard to clean. The BLM sends samples of their seed

purchases to a State seed testing laboratory to verify the

specifications. Because fire rehabilitation seeding is rushed

in the late summer and early fall, seed labs may only have

time to do a TZ test. If seed lots are found to have lower pure

live seed percentages than was specified in the contract, the

BLM may return the seed or adjust their price downward.

The BLM often seeds using bulk rates for introduced grasses

and legumes known to have high germination percentages

(>80 percent). These rates typically run from 1.8 to 3.6 lbs/

acre (2 to 4 kg/ha) of each species in a mixture of three or

more species. For fire rehabilitation in the past, mainly

introduced grasses have been seeded with some native

grasses and a few introduced legumes such as sainfoin

(Onobrychis vicifolia), small burnet (Sanguisorba minor), or

alfalfa (Medicago sativa) (Richards and others 1998).

Successful establishment of native grasses, forbs, and shrubs

may require higher seeding rates than those for simple

introduced species mixtures. Thompson (2002) found successful large-scale establishment of native seed mixtures at

17.8 lbs/acre (20 kg/ha) bulk total seed drilled on burned

sagebrush sites and 16 to 26.8 lbs/acre (18 to 30 kg/ha) total

seed broadcast and chained on burned pinyon-juniper sites.

Bulk rates required to get similar establishment from standard BLM seed mixes were generally lower and cost much

less, but did not result in comparable establishment of

native plants. Pyke and others (2003) found native plants in

BLM fire rehabilitation projects, but they were unable to

determine if those plants were residual to the sites or

established by seeding. Native mixtures may require higher

seeding rates than introduced species, and more careful

species selection for specific sites. In Thompson’s (2002)

study, seeding predominately Indian ricegrass, known for

its ability to emerge from deeper sandy soils, could have

saved the extra expense and failure of other native grasses

that were probably drilled too deep on the sandy sagebrush

site tested.
USDA Forest Service Proceedings RMRS-P-38. 2005 47

Plant Succession and Approaches to Community Restoration Roundy

Interaction/Reaction: Establishing

and Facilitating Diverse, Native

Communities __________________

Plant ecologists have identified numerous combinations

of plant-plant interactions (Barbour and others 1998).

Clements (1928) stressed the importance of competition as

a driving force in succession and what eventually dominates

a site. This makes sense for classic forest succession where

the dominant climax tree species are the ones that eventually develop large enough root and canopy structures to

compete best for resources. Disturbance plays a vital role in

opening up resources for a more diverse suite of species.

Although competition evidently is a major driving force for

the plants that eventually dominate a site after disturbance,

other interactions may be more important in providing for

long-term compatibility and diversity in a community. Plants

may partition resources among themselves in time by growing during different seasons, or in space by accessing different soil depths. Rabbitbrush is evidently less competitive

with grasses than Wyoming big sagebrush because its taproot uses deeper soils and avoids major competition with

shallower grass roots (Frischknecht 1963). On the other

hand, the two-layered surface and taproot system of Wyoming big sagebrush makes it a strong competitor with

perennial bunch grasses. Scientists in Turkmenistan developed range improvement practices to improve forage quality

and quantity for livestock. They selected woody species to

use deeper soil moisture than the extant herbaceous communities (Nechaeva 1985). Agroforestry and intercropping practices are dependent on finding crops and trees that yield

more when grown together than when grown separately.

The best known example is growing nitrogen-fixing legumes

with grasses. However, legume enhancement of grass growth

requires long periods of available soil moisture to work best.

Although shrubs are generally considered competitive

with herbaceous species, they also offer a suite of services to

a diverse community, such as

• Enhance soil fertility

• Catch seeds, spores, soil, and snow

• Moderate the temperature environment

• Improve soil aggregate stability and infiltration rates

• Harbor beneficial insects

(Call and Roundy 1991; West 1989). Because there are many

ecological and management benefits to mixed communities,

we would like to restore them or establish them in fire

rehabilitation seedings. Such a goal is much more ambitious

than the single species or simple introduced species mixes of

past rangeland revegetation. Use of native species in this

effort requires understanding about which ecotypes are best

adapted to specific regions or sites. The large-scale requirements of fire rehabilitation suggest that we should use

native plant materials with a wide range of adaptation if

possible.

Plant Materials Selection and

Improvement

Plant adaptation and plant materials trials in the past

have taken an agronomic approach. Numerous collections

are planted in separate rows or blocks in common gardens

and evaluated over many years. When a particular collection

appears to be more vigorous than the others, it is selected for

release. This approach takes a very long time to release a

given plant material, and fails to address some important

ecological aspects of mixed community restoration. It limits

genetic diversity for out-crossing species by keeping the

collections separate. Mass selection and other crossing techniques could be used to maximize genetic diversity. Very few

examples of such approaches have been tested. An emerging

approach is to certify seeds as “source-identified” (Young

1995). These collections are certified as originally collected

from a particular site, representing a specific environment.

Managers could choose “source-identified” plant materials

from sites with similar regional environmental conditions as

the sites they need to restore or rehabilitate. Once a large

native seed industry is developed, managers could even

choose physical mixes of a number of source-identified plant

materials to best cover their estimated environmental conditions. Such an industry will need establishment of large

seed warehouses and seed storage guidelines to allow stockpiling and a consistent market for these plant materials.

Commitment of government to large restoration efforts such

as the Great Basin Restoration Initiative will also support a

more consistent demand for specific native plant materials.

In the past, the demand for native seeds has been highly

variable and subject to the severity and extent of the current

fire season.

Establishing Diverse, Weed-Resistant

Communities

Another aspect of plant materials evaluation, not generally tested much in the past, is that of how well plant

materials work together rather than separately. The larger

concern, of course, is that of establishing “stable, diverse”

plant communities that are resistant to weed invasion. That

is a major challenge. Not only are we not really sure how

mixtures of plants will persist together, but it is a major

challenge to seed diverse mixtures and have all the seeded

species establish. When we have seeded aggressive, more

weed-resistant introduced grasses in mixtures with native

species, the introduced species eventually dominate (Pyke

1996). Approaches to more successfully establishing diverse

communities include (1) seeding more aggressive species at

a much lower rate than less aggressive ones, (2) seeding

certain species or mixes in separate rows, strips, or patches,

or (3) interseeding slower growing species such as some

shrubs after scalping out established grasses. Because sowing equipment and environmental conditions favor establishment of many grass species over that of shrub and forb

species, you cannot expect that just mixing species will

produce a community in the same proportion as the seed mix

(Newman and Redente 2001).

Weed-control strategies include designed disturbance to

reduce weed populations and controlled establishment of

desirable plants to preempt resources from weeds and prevent their invasion in the future (Sheley and others 1996). In

that regard, we really do not know enough about what

constitutes a community resistant to specific weeds on different sites. Our goal is to establish a suite of desirable plants
48 USDA Forest Service Proceedings RMRS-P-38. 20055

Roundy Plant Succession and Approaches to Community Restoration

that allows for their own coexistence, but excludes weeds.

Goldberg (1990) has suggested that plant-plant interactions

are often indirect through intermediate resources. To guide

seeding mixture recommendations of the future, we must

look at resource needs and use in time and space by desirable

and weedy species. This type of research can require many

years to develop recommendations, given the great range of

species, weeds, and environmental conditions. In the meantime, it is very important that diverse seedings be monitored

for response of both desirable and weedy species. Every fire

rehabilitation or restoration project is an experiment from

which something can be learned to guide future efforts.

It may be unrealistic to expect weed control and successful

revegetation of native plants on sites where precipitation is

low and the proximity of weed populations threaten

reinvasion. On such sites, use of bridging species such as

crested wheatgrass (Agropyron desertorum), which are more

easily established and resistant to weed invasion, may be

necessary. The bridging species could later be controlled and

may be much easier to replace with native species than

weedy species (Cox and Anderson 2004). Restoration will

require innovative approaches to meet the requirements of

native plants.

References ____________________

Barbour, M. G.; Burk, J. H.; Pitts, W. D.; Gilliam, F. S.; Schwartz,

M. W. 1998. Terrestrial plant ecology. Menlo Park, CA: Benjamin/

Cummings. 649 p.

Call, C. A.; Roundy, B. A. 1991. Perspectives and processes in

revegetation of arid and semiarid rangelands. Journal of Range

Management. 44: 543–549.

Chambers, J. C.; MacMahon, J. C.; Wade, G. L. 1992. Differences in

successional processes among biomes: importance for obtaining

and evaluating reclamation success. In: Chambers, J. C.; Wade,

G. L., eds. Evaluating reclamation success: the ecological consideration—proceedings of a symposium; 1990 April 23–26; Charleston, WV. Gen. Tech. Rep. GTR-NE-164. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest

Experiment Station: 59–72.

Clements, F. E. 1928. Plant succession and indicators. New York:

H. W. Wilson Company. 453 p.

Cox, R. D.; Anderson, V. J. 2004. Increasing native diversity of

cheatgrass-dominated rangeland through assisted succession.

Journal of Range Management. 57: 203–210.

Davenport, D. W.; Breshears, D. D.; Wilcox, B. P.; Allen, C. D. 1998.

Viewpoint: sustainability of piñon-juniper ecosystems—a unifying perspective of soil erosion thresholds. Journal of Range

Management. 51: 231–240.

Frischknecht, N. E. 1963. Contrasting effects of big sagebrush and

rubber rabbitbrush on production of crested wheatgrass. Journal

of Range Management. 16: 70–74.

Gleason, H. A. 1926. The individualistic concept of the plant association. Bulletin of the Torrey Botanical Club. 44: 463–481.

Goldberg, D. E. 1990. The components of resource competition in

plant communities. In: Grace, J. B.; Tilman, D., eds. Perspectives on plant competition. San Diego, CA: Academic Press,

Inc.: 27–49.

Harper, K. 1985. Predicting successional rates in Utah aspen

forests. In: Foresters’ future: leaders or followers? Proceedings

Society of American Foresters national convention; 1985 July 28–

31; Fort Collins, CO. [Place of publication unknown]: Society of

American Foresters: 96–100.

Middleton, B. 1999. Wetland restoration, flood pulsing, and disturbance dynamics. New York: John Wiley and Sons. 388 p.

Nechaeva, N. T., ed. 1985. Improvement of desert ranges in Soviet

Central Asia. New York: Harwood Academic Publishers. 327 p.

Newman, G. J.; Redente, E. F. 2001. Long-term plant community

development as influenced by revegetation techniques. Journal

of Range Management. 54: 717–724.

Ott, J. E.; McArthur, E. D.; Roundy, B. A. 2003. Vegetation of

chained and non-chained seedings after wildfire in Utah. Journal

of Range Management. 56: 81–91.

Pyke, D. A. 1996. Rangeland seedings and plantings: exotics or

natives? In: Edge, W. D.; Olson-Edge, S. L., eds. Sustaining

rangeland ecosystems symposium: proceedings; 1994 August 29-

31; La Grande, OR: Eastern Oregon State College SR 953.

Corvallis, OR: Oregon State University: 32–44.

Pyke, D. A.; McArthur, T. O.; Harrison, K. S.; Pellant, M. 2003.

Coordinated Intermountain Project—fire, decomposition, and

restoration. In: Palmer, A. R.; Scogings, P. F., eds. Proceedings of

the VIIth International Rangeland Congress; 2003 July 26–

August 1; Durban, South Africa. African Journal of Range &

Forage Science. 20: 1116–1124.

Richards, R. T.; Chambers, J. C.; Ross, C. 1998. Use of native plants

on Federal lands: policy and practice. Journal of Range Management. 51: 625–632.

Roundy, B. A. 1996. Revegetation of rangelands for wildlife. In:

Krausman, P. R., ed. Rangeland wildlife. Denver, CO: Society for

Range Management: 355–368.

Roundy, B. A.; Vernon, J. L. 1999. Watershed values and conditions

associated with pinyon-juniper communities. In: Monsen, S. B.;

Stevens, R., comps. Proceedings: ecology and management of

pinyon-juniper communities within the Interior West; 1997 September 15–18; Provo, UT. Proc. RMRS-P-9. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 172–187.

Sheley, R. L.; Svejcar, T.; Maxwell, B. D.; Jacobs, J. S. 1996.

Successional rangeland weed management. Rangelands. 18:

155–159.

Stromberg, J. C.; Patten, D. T.; Richter, B. D. 1991. Flood flows and

dynamics of Sonoran riparian forests. Rivers. 2: 221–235.

Summers, D. D.; Roundy, B. A.; Walker, S. C.; Davis, J. N. 2003.

Vegetation response of a Wyoming big sagebrush (Artemisia

tridentata ssp. wyomingensis) community to 6 mechanical treatments in Rich County, Utah. In: Abstracts: diversity through

time. Society for Range Management annual meeting; 2003

February 2–6; Casper, WY. Abstract 282.

Tausch, R. 1999. Historic pinyon and juniper woodland development. In: Monsen, S. B.; Stevens, R., comps. Proceedings: ecology

and management of pinyon-juniper communities within the Interior West; 1997 September 15–18; Provo, UT. Proc. RMRS-P-9.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Rocky Mountain Experiment Station: 12–19.

Thompson, T. W. 2002. Fire rehabilitation in Tintic Valley, Utah,

using native and exotic species. Provo, UT: Brigham Young

University. 67 p. Thesis.

West, N. E. 1983a. Great Basin—Colorado Plateau sagebrush semidesert. In: West, N. E., ed. Temperate deserts and semi-deserts.

New York: Elsevier Scientific Publishing Company: 331–349.

West, N. E. 1983b. Western Intermountain sagebrush steppe. In:

West, N. E., ed. Temperate deserts and semi-deserts. New York:

Elsevier Scientific Publishing Company: 351–374.

West, N. E. 1989. Spatial pattern-functional interactions in shrubdominated plant communities. In: McKell, C. M., ed. The biology

and utilization of shrubs. New York: Academic Press: 283–305.

Westoby, M.; Walker, B.; Noy-Meir, I. 1989. Opportunistic management of rangelands not at equilibrium. Journal of Range Management. 42: 266–274.

Whisenant, S. G. 1999. Repairing damaged wildlands. Cambridge,

UK: Cambridge University Press. 312 p.

Winkel, V.; Roundy, B. A. 1991. Effects of cattle trampling and

mechanical seedbed preparation on grass seedling emergence.

Journal of Range Management. 44: 176–180.

Young, S. A. 1995. Verification of germplasm origin and genetic

status by seed certification agencies. In: Roundy, B. A.; McArthur,

E. D.; Haley, J. S.; Mann, D. K., comps. Wildland shrub and

aridland restoration symposium: proceedings; 1993 October 19–

21; Las Vegas, NV. Gen. Tech. Report INT-GTR-315. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 293–295.
USDA Forest Service Proceedings RMRS-P-38. 2005 49

Thomas A. Jones and Steven R. Larson are Research Geneticists at the

USDA-Agricultural Research Service Forage and Range Research Laboratory,

Logan, UT 84322-6300 U.S.A., FAX: (435) 797-3075, e-mail: tomjones@cc.usu.edu,

stlarson@cc.usu.edu

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Status and Use of Important

Native Grasses Adapted to

Sagebrush Communities

Thomas A. Jones

Steven R. Larson

Abstract—Due to the emphasis on restoration, native cool-season

grass species are increasing in importance in the commercial seed

trade in the Western U.S. Cultivated seed production of these native

grasses has often been hampered by seed dormancy, seed shattering, and pernicious awns that are advantageous outside of cultivation. Relatively low seed yields and poor seedling establishment

have also restricted their cultivation. Most are members of the

Triticeae tribe. Bunchgrasses include Snake River wheatgrass

(Elymus wawawaiensis), bluebunch wheatgrass (Pseudoroegneria

spicata), basin wildrye (Leymus cinereus), and squirreltail (E. elymoides

and E. multisetus). Rhizomatous grasses include western wheatgrass

(Pascopyrum smithii), thickspike wheatgrass, and streambank wheatgrass (both E. lanceolatus), and beardless wildrye (L. triticoides).

Important non-Triticeae native bunchgrasses include native bluegrasses (Poa spp.) and Indian ricegrass (Achnatherum hymenoides).

These grasses may be either self-pollinated, cross-pollinated, or

apomictic. These mating systems are reflected in the patterns of

genetic variation characteristic of these species. At the USDA

Agricultural Research Service (ARS) Forage and Range Research

Laboratory at Utah State University, our goals are to understand

distribution-wide patterns of genetic variation and to develop native cool-season grasses that are adapted to rangeland environments, are reflective of natural patterns of genetic variation, and

are amenable to commercial seed production. To best accomplish

these goals, we are attempting to develop a better understanding of

the correlation between genetic variation and ecological adaptation

at a variety of levels ranging from the whole plant to the DNA

molecule. Besides ourselves, plant materials of these species have

been released by USDA Natural Resources Conservation Service

(NRCS) Plant Materials Centers (Bridger, MT; Aberdeen, ID; Pullman, WA; Lockeford, CA; Los Lunas, NM), Forest Service Shrub

Sciences Laboratory (FS SSL) (Provo, UT), and Upper Colorado

Environmental Plant Center (UCEPC) (Meeker, CO).

Squirreltail (Elymus elymoides

[Raf.] Swezey and E. multisetus

[J.G. Smith] M.E. Jones) __________

Squirreltail is a short-lived perennial grass that is a

prominent understory species in the sagebrush steppe community (Jones 1998). Squirreltail is a complex of five taxa,

all of which are found in southwestern Idaho (Wilson 1963).

Each taxon can be easily identified by spike morphology

using a dichotomous key. E. elymoides ssp. elymoides is the

most common and widespread taxon and is probably most

closely related to E. elymoides ssp. californicus, which is

prominent on the eastern slope of the Sierra Nevadas. E.

elymoides ssp. brevifolius is especially common in the central and southern Rockies where plants and seeds are

exceptionally large; however, these populations are conspicuously different from E. elymoides ssp. brevifoliusplants

originating in the Northwest. E. elymoides ssp. hordeoides is

the most diminutive of the group, and probably the least

common overall.E. multisetus, commonly called big squirreltail, is often considered a distinct species from the others, a

position supported by our molecular data (Larson and others

2003). It is most common in the Northwest.

Squirreltail is a self-pollinated tetraploid (2n = 28) that

genetically consists of the St and H genomes, which include

14 chromosomes (7 pairs) each. These two genomes are

characteristic of Elymus worldwide and are indicative of the

evolutionary history of this polyploid genus. The St genome

originated from the bluebunch wheatgrasses (Pseudoroegneria spp.) and the H genome from the barleys (Hordeum spp.). Most Elymus species are predominately selfpollinating, including blue wildrye (E. glaucus), slender

wheatgrass (E. trachycaulus), and Canada wildrye (E.

canadensis). Prominent cross-pollinated exceptions include

thickspike wheatgrass, Snake River wheatgrass, and the

introduced species, quackgrass (E. repens) (Jensen and Asay

1996; Jensen and others 1990).

Squirreltail germplasms released to date are Sand Hollow

(E. multisetus; Emmett, ID) (Jones and others 1998), Toe

Jam Creek (E. elymoides ssp. californicus; Tuscarora, NV),

Fish Creek (E. elymoides ssp. elymoides; Carey, ID), and

Tusas (E. elymoides ssp. brevifolius; multiple locations in

New Mexico). In addition, several seed growers are producing local proprietary seed sources.
50 USDA Forest Service Proceedings RMRS-P-38. 2005

Jones and Larson Status and Use of Important Native Grasses Adapted to Sagebrush Communities

Squirreltail exhibits a high degree of racial differentiation, as described for other species by Clausen and others

(1947). We evaluated squirreltail accessions (27 in data set 1,

47 in data set 2) for a battery of ecological traits, which could

be used to characterize the ecological relationships between

and within taxa (Jones and others 2003). For data set 1, 13

traits were measured, including days to seedling emergence,

length of the seedling’s first leaf, total plant dry matter, rootto-shoot ratio, leaf area, specific leaf area, root length,

specific root length, heading date, seed mass, emergence

index (from 20 mm), emergence index (from 60 mm), activity

of the nitrate reductase enzyme, plant height, and heading

date. For data set 2, these same traits were measured except

leaf area, specific leaf area, the emergence indices, and

nitrate reductase activity. Seed mass was also measured for

data set 2.

In data set 1, E. multisetus accessions showed greatest

seedling vigor and root development. E. elymoides ssp. elymoides accessions had lowest seed mass and earliest phenological development. E. elymoides ssp. brevifolius had thickest leaves and slowest germination. Taken together, the 13

traits clearly demarcated the three groups of accessions. In

data set 2, E. elymoides ssp. elymoides, E. elymoides ssp.

brevifolius, and E. multisetus accessions again separated

discretely, but E. elymoides ssp. brevifolius accessions separated into three subgroups. Early (subgroup B) and latematuring (subgroup A) accessions from the Rocky Mountains separated apart from each other and also apart from

intermediate-maturing accessions from southwestern Idaho

(subgroup C).

Native Bluegrasses

(Poa secunda Presl.) _____________

Native bluegrasses also serve as important understory

components of the sagebrush steppe vegetation. A large

number of scientific names (P. secunda, P. ampla, P. canbyi,

P. gracillima, P. incurva, P. juncifolia, P. nevadensis, P. scabrella, and P. curtifolia) have been given to various of the

native bluegrasses, but Kellogg (1985, 1990) combined them

all into P. secunda, except P. curtifolia, an endemic from

central Washington. Kellogg argued that, because morphological variation among these entities is continuous,

Table 1—Seed production acreages of all classes of certified seed in the United States from

1996 to 2000 for native cool-season grasses commonly seeded on rangelands in the

Intermountain Regiona.

Species/Cultivar 1996 1997 1998 1999 2000 Mean

Western wheatgrass 934 959 1,626 3,029 3,371 1,984

Rosana 366 355 992 1,593 1,662 994

Arriba 209 255 286 919 1,055 545

Barton 273 272 272 432 548 359

Rodan 46 37 36 45 66 46

Flintlock 40 40 40 40 40 40

Thickspike wheatgrass 931 1,196 1,135 2,183 3,666 1,822

Critana 459 430 501 1,086 1,840 863

Sodar 345 505 299 596 905 530

Bannock 127 229 312 453 505 325

Schwendimar 0 0 23 48 416 97

Elbee 0 32 0 0 0 6

Native bluegrasses 469 560 956 1,267 1,721 995

Sherman 394 494 836 1,239 1,623 917

Canbar 75 66 120 28 98 77

Snake River wheatgrass/Secar 313 292 600 949 2,054 842

Basin wildrye 512 416 572 851 941 658

Magnar 371 369 445 526 448 432

Trailhead 141 47 127 325 493 227

Bluebunch wheatgrass 126 537 603 586 1,260 622

Goldar 49 422 473 401 965 462

Whitmar 77 115 130 185 295 160

Indian ricegrass 190 126 196 307 944 353

Nezpar 129 107 115 133 499 197

Rimrock 26 3 65 55 251 80

Paloma 35 16 16 119 194 76

Beardless wildrye 59 74 93 0 65 58

Shoshoneb 59 74 91 0 65 58

Rio 0 0 2 0 0 0

Squirreltail/Sand Hollow 0 0 0 8 8 3

Proprietary (all species) 30 419 0 289 2 148

Total 3,564 4,579 5,781 9,469 14,032 7,485

a

Figures compiled from AOSCA (1996–2000).

bActually the introduced Leymus multicaulis.
USDA Forest Service Proceedings RMRS-P-38. 2005 51

Status and Use of Important Native Grasses Adapted to Sagebrush Communities Jones and Larson

any attempt to subdivide this group would be arbitrary.

Nevertheless, she tolerated the retention of common names

within the group.

Releases are ‘Canbar’ (WA), commonly known as “canby

bluegrass,” ‘Sherman’ (OR), commonly known as “big bluegrass” because of its larger stature and longer leaves, and

Reliable germplasm (Yakima, WA), commonly known as

“Sandberg bluegrass” (fig. 1). Certified seed production

acreage of Sherman has greatly increased in recent years,

while acreage of Canbar remains low (table 1). Sandberg

bluegrass, a diminutive plant, mimics the phenology of

cheatgrass. It flowers at least as early as cheatgrass and

senesces upon seed ripening in late spring or early summer.

Taxonomic confusion in Poa is enhanced by facultative

apomixis, an asexual form of reproduction by seed that may

preempt sexual reproduction (Kellogg 1987). This means

that new genetic variation generated by sexuality can be

replicated in large quantities at or close to 84 chromosomes,

making them dodecaploids (12x), but big bluegrass plants

usually have about 63 chromosomes (9x) (Hartung 1946).

Obviously, this odd number is something that could only be

fixed asexually. The frequency with which apomixis occurs

probably varies with genotype, but few data have been

collected to determine the mean and range among genotypes. Do 63-chromosome plants arise from hybridization of

84-chromosome plants and 42-chromosome (6x) plants? Do

63-chromosome plants have a consistently higher level of

apomixis than 84-chromosome plants or do they reproduce

sexually, spinning off more odd chromosome-numbered

plants? We believe a better understanding of these issues

would provide a more complete understanding of the taxonomy of the native Poa complex.

We used amplified fragment length polymorphism (AFLP)

analysis to characterize genetic diversity within Canbar,

Sherman, Mountain Home (ID), and Reliable and genetic

divergence between them (Larson and others 2001). AFLP

methodology involves using sets of DNA primers to copy

DNA fragments of different lengths that are then multiplied

via the polymerase chain reaction. The resultant array

serves as a “fingerprint” of the plant’s genotype. Genetic

diversities of the wildland collections, Mountain Home and

Reliable, are similar to one another, but much greater than

Canbar (fig. 2). Sherman shows no genetic diversity whatsoever, which is excellent evidence for apomixis and hints at

Figure 1—Points of origin of plant materials of squirreltail, native bluegrasses, Indian ricegrass, bluebunch wheatgrass, Snake

River wheatgrass, thickspike wheatgrass, basin wildrye, and western wheatgrass.

Secar

Rosana

Arriba

Bluebunch

wheatgrass

Snake River

wheatgrass

Squirreltail Indian 

ricegrass

Thickspike

wheatgrass

Native 

bluegrasses

Basin

wildrye

Western 

wheatgrass
52 USDA Forest Service Proceedings RMRS-P-38. 2005

Jones and Larson Status and Use of Important Native Grasses Adapted to Sagebrush Communities

Figure 2—Genetic divergence (plane) and genetic

diversity (vertical) of ‘Sherman’ big bluegrass,

‘Canbar’ canby bluegrass, and Reliable and

Mountain Home Sandberg bluegrass germplasms.

the second question posed in the previous paragraph. Genetic divergence (separation on the plane) between Mountain Home and Reliable is small. These Sandberg bluegrasses are intermediate to Canbar (another 84-chromosome

genotype) and Sherman (2n = 63), but more similar to the

former.

Indian Ricegrass (Achnatherum

hymenoides [Roem. & Schult.]

Barkworth) _____________________

Indian ricegrass is a short-lived perennial bunchgrass

that favors light-textured soils, especially fine sandy loams,

sandy loams, loamy sands, and sands (Jones 1990). It is

found in especially arid environments in the Intermountain

Region as well as the Great Plains. Though it is a cool-season

C3 grass, it grows longer into the summer heat than native

wheatgrasses and wildryes. This species exhibits great genetic diversity between and within populations. Seed polymorphism is common in Indian ricegrass (Jones and Nielson

1996). Larger seeds typically have greater seed dormancy

than the smaller seeds (Young and Evans 1984). These two

seed morphs are produced on different plants, which usually

differ in other respects as well. Like squirreltail, Indian

ricegrass is highly self-pollinating.

Releases are ‘Paloma’ (Pueblo, CO), ‘Nezpar’ (White Bird,

ID), ‘Rimrock’ (Billings, MT), and Star Lake (McKinley

County, NM), and White River (Rio Blanco County, CO)

germplasms (fig. 1). Certified seed production acreage has

increased in recent years (table 1).

Bluebunch Wheatgrass

(Pseudoroegneria spicata

[Pursh] A. Löve)_________________

Bluebunch wheatgrass is widespread in the Intermountain Region, foothills, and open slopes of the Rocky Mountains and the northern Great Plains. It is probably most

frequent in the region where Washington, Oregon, and

Idaho converge. Many populations have been reduced or

extirpated because this grass is both palatable and highly

susceptible to overgrazing in the spring. Populations may be

awned or awnless or are often mixed. Cultivars include the

awnless Whitmar (Colton, WA) and the awned Goldar
USDA Forest Service Proceedings RMRS-P-38. 2005 53

Status and Use of Important Native Grasses Adapted to Sagebrush Communities Jones and Larson

(Umatilla National Forest, Asotin County, WA) (fig. 1).

Certified seed production acreage of Goldar has greatly

exceeded Whitmar in recent years. ‘Secar’ (Lewiston, ID)

was released as a bluebunch wheatgrass, but was later

discovered to be the newly recognized Snake River wheatgrass (see below). P-7, a multiple-origin polycross of 25

accessions, and Anatone (WA) are recent germplasm releases. Our work at ARS has emphasized selection for

grazing tolerance and improved seed production.

Bluebunch wheatgrass may be either diploid (2n = 14) or

tetraploid (2n = 28), with the diploid being far more common.

Only the St genome is found in bluebunch wheatgrass; it is

present in single or double dose depending on ploidy.

Tetraploids appear to be most frequent in mesic regions such

as southern interior British Columbia and parts of southeastern Washington. Accessions may be mixed for ploidy,

but this is not visually obvious because the two ploidys are

morphologically indistinguishable. In the past, chromosome

number has been measured directly using root-tip cells, but

today we typically measure it indirectly using flow cytometry,

a much more rapid and convenient technique.

While bluebunch wheatgrass is the only New World species of Pseudoroegneria, other taxa are known in the Old

World (Jensen and others 1995). These Old World taxa can

be morphologically indistinguishable from bluebunch wheatgrass and can be successfully hybridized with it. Both the

pollen and the seed of these hybrids are virtually sterile.

AFLP analysis was used to measure genetic diversity and

divergence (see Sandberg bluegrass, above) in and among

Goldar, Whitmar, and P-7 (Larson and others 2000). As

expected, P-7, the multiple-origin polycross, had greater

diversity than either Goldar or Whitmar, both of which

originated from single sites in southeastern Washington

(fig. 3). Divergence was greatest between Goldar and

Whitmar, with P-7 being more similar to Goldar than to

Whitmar.

Snake River Wheatgrass (Elymus

wawawaiensis J. Carlson &

Barkworth) _____________________

Relative to bluebunch wheatgrass, Snake River wheatgrass has a limited distribution within eastern Oregon and

Washington and northern and central Idaho. Its frequency

has likely been severely reduced because of the high degree

of cultivation in that region. Snake River wheatgrass is a

bunchgrass with excellent seed production and drought

tolerance. It has performed very well in seedings in the

Snake River Plain, despite the fact that it is not native to

southern Idaho. The most southerly material of which we

have knowledge originates from near Cambridge, ID. Snake

River wheatgrass is less susceptible to overgrazing than

bluebunch wheatgrass (Jones and Nielson 1997). At ARS we

are selecting for clipping tolerance to improve its tolerance

to grazing.

Until 1985, Snake River wheatgrass was confused with

bluebunch wheatgrass (Carlson and Barkworth 1997). In

1980 ‘Secar’ (fig. 1) was released as a bluebunch wheatgrass.

Snake River wheatgrass is an StH species, thus its hybrids

with bluebunch wheatgrass, an St species, are cytologically

irregular (Carlson 1986). However, hybrids of Snake River

wheatgrass with thickspike wheatgrass, another StH species, are cytologically regular (Jones and others 1995). It was

these observations that initially justified the recognition of

Snake River wheatgrass as a separate species apart from

bluebunch wheatgrass and its placement in the genus Elymus

rather than Pseudoroegneria. Traditionally, analysis of chromosome pairing in hybrids has been used to determine

genomic composition, but we now utilize the FISH (fluorescent in situ hybridization) technique. FISH utilizes DNAbinding dyes that are specific to particular genomes. When

used in combination, these dyes generate red or yellow

fluorescence, depending on the genome. Intermediate genomes fluoresce orange.

While Snake River and bluebunch wheatgrass share a

superficial resemblance, there are many features that distinguish the two from each other (Jones and others 1991).

The spike of Snake River wheatgrass is more compact than

bluebunch wheatgrass, primarily because the rachis internodes of Snake River wheatgrass are shorter. Snake River

wheatgrass is always awned, whereas bluebunch wheatgrass may be awned or awnless. Snake River wheatgrass is

always tetraploid (2n = 28), while bluebunch wheatgrass is

primarily diploid and occasionally tetraploid (2n = 14, 28).

Figure 3—Genetic divergence (plane) and genetic

diversity (vertical) of ‘Goldar’

and ‘Whitmar’ bluebunch

wheatgrass and the P-7

multiple-origin polycross

bluebunch wheatgrass

germplasm.
54 USDA Forest Service Proceedings RMRS-P-38. 2005

Jones and Larson Status and Use of Important Native Grasses Adapted to Sagebrush Communities

Snake River wheatgrass seed is considerably smaller than

bluebunch wheatgrass seed. Snake River wheatgrass glumes

are more lanceolate in shape than the nontapered bluebunch

wheatgrass glumes, but size of the glumes is not diagnostic.

Certified seed production acreage of Snake River wheatgrass has greatly increased in recent years and now greatly

exceeds that of bluebunch wheatgrass (table 1).

Thickspike Wheatgrass (Elymus

lanceolatus [Scribn. & J.G. Smith]

Gould)_________________________

Thickspike wheatgrass is found throughout the Intermountain Region and the northwestern Great Plains, but is

most common in Wyoming. Thickspike wheatgrass is a crosspollinating rhizomatous grass closely related to Snake River

wheatgrass. It is generally found on loamy to sandy soils. Like

Snake River wheatgrass, thickspike wheatgrass is an StH

tetraploid (2n = 28). Thickspike wheatgrass is much more

tolerant of overgrazing than either Snake River wheatgrass

or bluebunch wheatgrass. Cultivars appropriate for our region include Bannock (OR, ID, WA), Schwendimar (The

Dalles, OR), Elbee (AB, SK), Critana (Havre, MT), and Sodar

(Canyon City, OR) (fig. 1). Thickspike wheatgrass rivals

western wheatgrass as the most important species for certified seed production acreage (table 1). In recent years Critana

has been the leading cultivar, followed by Sodar and Bannock.

Sodar is known in the trade as “streambank wheatgrass,”

technically a botanical variety of thickspike wheatgrass

(Dewey 1969). In Canada, thickspike wheatgrass is known

as “northern wheatgrass.”

Basin Wildrye (Leymus cinereus

[Scribn. & Merr.] A. Löve) _________

Basin wildrye is a large-statured bunchgrass that is prominent in the Intermountain region. It is valued for winter

grazing and for the shelter it provides during calving. In the

Snake River Plain, basin wildrye is often found in locations

with deep soils and high water-holding capacity. Basin

wildrye has two chromosome races, tetraploid (2n = 28) and

octoploid (2n = 56), that feature the Ns and Xm genomes in

single or double dose, respectively. The Ns genome originated in Psathyrostachys, the Russian wildryes, but the

origin of the Xm genome is uncertain.

The octoploid race, characterized by more robust plants

that exhibit a glaucous blue color under drought conditions,

is found in central and northeastern Oregon, northern Idaho,

eastern Washington, and interior British Columbia. The

tetraploid race, with somewhat smaller plants that remain

green (rather than turning blue) in response to drought, is

found in Central and southeastern Oregon, southern Idaho,

Nevada, Utah, Colorado, Wyoming, Montana, Alberta, and

Saskatchewan. Magnar is an octoploid cultivar (BC) and

Trailhead is a tetraploid cutivar (Roundup, MT) (fig. 1).

Magnar is the older cultivar, but its certified seed production

acreage has remained stagnant in recent years (table 1).

Trailhead’s acreage has greatly increased in recent years

and it now is equal to Magnar’s. ARS is assembling several

accessions into a multiple-origin polycross oriented towards

northern Nevada, similar to the approach used in the development of P-7 bluebunch wheatgrass.

Beardless Wildrye (Leymus

triticoides [Buckley] Pilger) _______

Beardless wildrye is a rhizomatous corollary to the bunchgrass basin wildrye in the genus Leymus, much as thickspike

wheatgrass is to Snake River wheatgrass in the genus

Elymus. Beardless wildrye is most common in northern

Nevada, eastern Oregon, and northeastern California. It is

an important riparian species, but it is also adapted to arid

upland sites. Beardless wildrye and basin wildrye often

occupy the same site and hybrids are common in such locations. Beardless wildrye has very high levels of seed dormancy,

even higher than Indian ricegrass, a feature that is not

prominent in basin wildrye. The only cultivar of beardless

wildrye is Rio (Stratford, CA), which is propagated commercially by rhizomes because of its poor germination. ‘Shoshone’

is prominent in the seed trade where it is known as beardless

wildrye, but it is now known to be Leymus multicaulis, an

introduced species. Leymus triticoides and L. multicaulis

are easily distinguished when the two species are grown

side-by-side. Shoshone certified seed acreage has not increased in recent years (table 1).

Western Wheatgrass (Pascopyrum

smithii [Rydb.] A. Löve) __________

Western wheatgrass is a cross-pollinating rhizomatous

grass like thickspike wheatgrass. These two grasses are

often confused, but they may be distinguished on the basis

of glume shape. The glume of western wheatgrass is asymmetrical, while the thickspike wheatgrass glume is symmetrical. However, determination of ploidy is the most

foolproof method to separate these grasses; thickspike wheatgrass is always tetraploid (2n = 28) and western wheatgrass

is always octoploid (2n = 56) (Dewey 1975).

Western wheatgrass is believed to have originated from

hybridization of thickspike wheatgrass and beardless wildrye

(Dewey 1975). Western wheatgrass combines the St and H

genomes of Elymus with the Ns and Xm genomes of Leymus.

The frequency and distribution of western wheatgrass exceed either of its parents. Western wheatgrass is rhizomatous because both of its parents are rhizomatous. Western

wheatgrass has an intermediate level of seed dormancy

because beardless wildrye has high seed dormancy and

thickspike wheatgrass does not exhibit seed dormancy.

Compared to thickspike wheatgrass, western wheatgrass is

more rhizomatous, better adapted to heavier soils, and less

adapted to arid conditions.

All cultivars of western wheatgrass originate in the Great

Plains. The two most widely used in the Intermountain

Region are Rosana (Forsyth, MT) to the north and Arriba

(Flagler, CO) to the south (fig. 1). Certified seed production

acreage has increased greatly in recent years (table 1).

Rosana is the leading cultivar (table 1). Western wheatgrass and thickspike wheatgrass acreages are similar and

are greater than those of other species discussed herein.
USDA Forest Service Proceedings RMRS-P-38. 2005 55

Status and Use of Important Native Grasses Adapted to Sagebrush Communities Jones and Larson

References _____________________

Association of Official Seed Certifying Agencies (AOSCA). 1996–

2000. Acres applied for certification by seed certification agencies.

Meridian, ID: Association of Official Seed Certifying Agencies.

Carlson, J. R. 1986. A study of morphological variation within

Pseudoroegneria spicata (Pursh) A Löve (Poaceae:Triticeae).

Corvallis, OR: Oregon State University. Thesis.

Carlson, J. R.; Barkworth, M. E. 1997. Elymus wawawaiensis: A

species hitherto confused with Pseudoroegneria spicata(Triticeae,

Poaceae). Phytologia. 83: 312–330.

Clausen, J.; Keck, D. D.; Hiesey, W. M. 1947. Heredity of geographically and ecologically isolated races. American Naturalist. 81:

114–133.

Dewey, D. R. 1969. Synthetic hybrids of Agropyron albicans X A.

dasystachyum, Sitanion hystrix, and Elymus canadensis. American Journal of Botany. 56: 664–670.

Dewey, D. R. 1975. The origin of Agropyron smithii. American

Journal of Botany. 62: 524–530.

Hartung, M. E. 1946. Chromosome numbers in Poa, Agropyron, and

Elymus. American Journal of Botany. 33: 516-531.

Jensen, K. B.; Asay, K. H. 1996. Cytology and morphology of Elymus

hoffmanni (Poaceae: Triticeae): a new species from the Erzurum

Province of Turkey. International Journal of Plant Science. 157:

750–758.

Jensen, K. B.; Curto, M.; Asay, K. H. 1995. Cytogenetics of Eurasian bluebunch wheatgrass and their relationship to North

American bluebunch and thickspike wheatgrasses. Crop Science. 35: 1157–1162.

Jensen, K. B.; Zhang, Y. F.; Dewey, D. R. 1990. Mode of pollination

of perennial species of the Triticeae in relation to genomically

defined genera. Canadian Journal of Plant Science. 70: 215–225.

Jones, T. A. 1990. A viewpoint on Indian ricegrass: its present

status and future prospects. Journal of Range Management. 43:

416–420.

Jones, T. A. 1998. Viewpoint: the present status and future prospects of squirreltail research. Journal of Range Management. 51:

326–331.

Jones, T. A.; Nielson, D. C. 1996. Seed size and shape and lemma

color polymorphism in Oryzopsis hymenoides (Roem. & Schult.)

Ricker. In: West, N. E., ed. Proceedings; V International Rangeland Congress; 1995 July 23–28; Salt Lake City, UT: 272–273.

Jones, T. A.; Nielson, D. C. 1997. Defoliation tolerance of bluebunch

and Snake River wheatgrasses. Agronomy Journal. 89: 270–274.

Jones, T. A.; Nielson, D. C.; Arredondo, J. T.; Redinbaugh, M. G.

2003. Characterization of diversity among 3 squirreltail taxa.

Journal of Range Management. 56: 474–482.

Jones, T. A.; Nielson, D. C.; Carlson, J. R. 1991. Development of a

grazing-tolerant native grass for revegetating bluebunch wheatgrass sites. Rangelands. 13: 147–150.

Jones, T. A.; Nielson, D. C.; Ogle, D. G.; Johnson, D. A.; Young, S. A.

1998. Registration of Sand Hollow squirreltail germplasm. Crop

Science. 38: 286.

Jones, T. A.; Wang, R. R-C.; Li, L-H. 1995. Meiotic stability of

intersubspecific hybrids of Snake River X thickspike wheatgrasses. Crop Science. 35:962-964.

Kellogg, E. A. 1985. Variation and names in the Poa secunda

complex. Journal of Range Management. 38: 516–521.

Kellogg, E. A. 1987. Apomixis in the Poa secunda complex. American Journal of Botany. 74: 1431–1437.

Kellogg, E. A. 1990. Variation and species limits in agamospermous

grasses. Systematic Botany. 15: 112–123.

Larson, S. R.; Jones, T. A.; Hu, Z-M.; Palazzo, A. J.; McCracken, C. L.

2000. Genetic diversity of bluebunch wheatgrass cultivars and a

multiple-origin polycross. Crop Science. 40: 1142–1147.

Larson, S. R.; Jones, T. A.; McCracken, C. L.; Jensen, K. B. 2003.

Amplified fragment length polymorphism in Elymus elymoides,

Elymus multisetus, and other Elymus taxa. Canadian Journal of

Botany. 81: 789–804.

Larson, S. R.; Waldron, B. L.; Monsen, S. B.; St. John, L.; Palazzo,

A. J.; McCracken, C. L.; Harrison, R. D. 2001. AFLP variation in

agamospermous and dioecious bluegrasses of Western North

America. Crop Science. 41: 1300–1305.

Wilson, F. D. 1963. Revision of Sitanion (Triticeae, Gramineae).

Brittonia. 15: 303–323.

Young, J. A.; Evans, R. A. 1984. Germination of seeds of ‘Paloma’

and ‘Nezpar’ Indian ricegrass. Journal of Range Management.

37: 19–21.

Nineleaf biscuitroot
56 USDA Forest Service Proceedings RMRS-P-38. 2005

Scott C. Walker is Wildlife Research Biologist, Utah Division of Wildlife

Resources, Northern Region Office, 515 East 5300 South, Ogden, UT 84401;

e-mail: scottwalker@utah.gov. Nancy L. Shaw is Research Botanist, Rocky

Mountain Research Station, USDA Forest Service, 316 E. Myrtle, Boise, ID

83702; e-mail: nshaw@fs.fed.us

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Current and Potential Use

of Broadleaf Herbs for

Reestablishing Native

Communities

Scott C. Walker

Nancy L. Shaw

Abstract—Use of forbs for revegetation in the Intermountain West

has been problematic due to the large number of species and lack of

research data. Some forbs are found in numerous plant communities and distributed over wide geographic ranges while others are

more narrowly adapted. Seed sources for revegetation use may be

selected from species and ecotypes indigenous to the planting area.

Management of local stands to improve seed production may be

required to insure the availability of adequate quantities of seed.

Alternatively, seed of an increasing number of commonly used

species is being grown in agricultural settings with more reliable

seed supplies resulting. Advances in wildland seeding methodology

and forb seed production, harvesting, and conditioning technology

have resulted from recent research and plant materials development programs.

Introduction ____________________

Shrublands of the Great Basin have undergone drastic

changes since early settlement. Domestic livestock grazing

and other human impacts, the spread of weedy annuals, and

concurrent changes in fire frequency and intensity have

reduced the diversity of sagebrush (Artemisia spp.) communities. Considerable effort has been directed toward revegetating disturbed rangelands. Seedings have generally been

designed to increase forage production for livestock and,

more recently, to re-establish native plant communities and

wildlife habitat. Seed mixes have included various introduced and native grasses, forbs, and shrubs (Stevens 1983).

Over the past 30 to 40 years most seed mixes were dominated

by introduced grass species. These grasses were seeded

because of their ease of establishment, forage production

potential, ability to compete with invasive exotics, and their

soil stabilization characteristics. In addition, the required

quantities of seed were usually available. Plant materials

development efforts over this period have gradually increased the number of native grass and forb species seeded

and the amount of seed marketed. Research has also improved our understanding of secondary successional processes occurring when combinations of species are planted

together (Walker and others 1995).

The use of native forbs in restoration has been limited due

to a number of factors. These include the large number of

forb species present in the Great Basin, our limited knowledge of seed production and seeding requirements for most

species, the difficulty of harvesting forb seed from wildland

stands, the highly unpredictable quality and quantity of

wildland seed collections, and the frequently high cost of

available seed. Overcoming these obstacles requires selection of accessions adapted to proposed planting areas, development of seed and seeding technology for each species, and

establishment of seed production fields. In some situations,

managing selected wildland stands to improve seed production may be a viable alternative when specific ecotypes are

required for use over extensive areas. Both options require

considerable time and effort for the development of reliable

seed supplies.

Importance of Native Forbs

Native forbs are common and important components of

most native plant communities of the Great Basin. Numerous species occur within this region, but most are not as

widespread as the more common native grasses. Salt desert

shrub and Wyoming big sagebrush (Artemisia tridentata

ssp. wyomingensis) communities support fewer native forbs

than more mesic shrubland communities. Forbs occasionally grow in pure stands, but more frequently they are found

as associated species growing with grasses and other forbs in

grasslands, or as understory species in shrub or tree communities. Individual species may be distributed over wide

geographic ranges and occur in a variety of plant communities, or they may be narrowly distributed edaphic endemics.

Some species exhibit considerable variability within and

among populations (Shaw and Monsen 1983).

The addition of native forbs to revegetation projects contributes to the establishment of a more complex community

and enriches the food supply for sagebrush-associated wildlife, such as mule deer (Odocoileus hemionus), pronghorn

(Antilocapra americana), elk (Cervus elaphus), sage-grouse

(Centrocercus urophasianus), sharp-tailed grouse (Tympanuchus
USDA Forest Service Proceedings RMRS-P-38. 2005 57

Current and Potential Use of Broadleaf Herbs for Reestablishing Native Communities Walker and Shaw

phasianellus), blue grouse (Dendragapus obscurus), and

small mammals. Native forbs play a number of roles in

wildland seedings:

1. They increase community diversity, health, and

resilience.

2. Pioneer forbs provide ground cover and soil stabilization on disturbed and unstable sites.

3. Leguminous forbs improve nitrogen availability.

4. Forbs reduce the ability of exotic species to enter the

community.

5. Vegetative plant parts as well as fruits and seeds of

individual forb species are often valuable seasonal food

sources for specific organisms.

6. The fire resistance of seedings may be improved by the

addition of forbs that remain green well into the summer.

7. Forbs improve the aesthetics of seeded disturbances

and low maintenance landscaping projects.

Sage-Grouse Use of Forbs

Forb-rich sagebrush communities are vital to the survival

of sage-grouse and other species that rely on the complex of

vegetation and associated faunal components of the community. The importance of forbs in the survival of upland bird

species is becoming better understood and is proving to be

more important than previously thought. In spring, sagegrouse diets change from sagebrush-dominated to forbdominated. Breeding habitats used by prelaying hens should

provide a diversity of forbs high in calcium, phosphorus, and

protein; the condition of these areas may greatly affect nest

initiation rate, clutch size, and subsequent reproductive

success (Barnett and Crawford 1994; Connelly and others

2000). Juvenile sage-grouse are dependent upon insects and

succulent forbs as critical food sources after hatching and

until brood dispersal in fall. Chicks follow the habitat use of

the hens with broods and broodless hens (Barnett and

Crawford 1994; Beck and Mitchell 1997; Braun and others

1977; Wallstead 1971).

Reestablishment of

Native Forbs ___________________

Selection of Seed Sources

For seedings on rangeland sites, local seed sources of

native forbs should be added to seed mixes of compatible

grasses and shrubs. Where reestablishment of native communities is the goal, land managers may need to contract

collection and increase of local seed to produce adequate

supplies for the project site. Released forb varieties or

prevariety releases are available for a few species (table 1).

Seed supplies of released materials are generally produced

in agricultural seed fields and sufficient quantities for major

seedings may be available or can be produced under contract. The area of adaptation for these has been determined

to varying degrees, and is described in the release documentation. Species and seed sources known to be competitive

with invasive species must be used if weeds are expected to

be a problem. The availability and cost of native forb seed

remains the greatest obstacle to inclusion of these species in

seeding mixes.

Seeding Mixes

Although forb species may make up only a small portion of

the plant composition and cover in sagebrush habitats, they

are extremely important in the diets of sage-grouse broods

and many other vertebrate and invertebrate species. Consequently, it is important to provide forb diversity in seedings,

even when species must be seeded at low densities or spot

seeded in favorable areas. Established seedings should be

managed to allow forb growth to continue through spring

and summer, particularly in sage-grouse breeding habitat

(Paige and Ritter 1999).

Seeding Techniques

Even the commonly seeded forbs represent a number of

plant families and genera and, consequently, an array of fruit

and seed types and sizes is represented. Successful seeding

requires that seedings be conducted at the appropriate season, usually late fall in the Great Basin, in order that moist

prechilling requirements be met. This permits seeds to

germinate in early spring when soil moisture conditions are

most likely to be favorable. Seed must be planted at the

appropriate depth. Another consideration is the condition of

the fruit or seed structure planted. Although some will flow

well and are easily seeded, others tend to mat together and

must be seeded in drill boxes with agitators. Broadcasting

can generally be successfully accomplished if seeds are

diluted with seeds of grasses and covered as necessary

following seeding.

Relative competitive abilities and affecting factors are

largely unknown for native forbs. Nevertheless, substantial

anecdotal information on common species strongly suggests

that forbs should be sown separately or with other species

that are sown in low densities to minimize competition.

Likewise, the ability of seeded forbs to compete with invasive

species that may be present on the site must be considered.

If site preparation practices are not adequate to control

weeds, the use of scarce and valuable seed, or the seeding

itself, should be reconsidered.

Like all seedings, those including native forbs should be

protected from grazing until plants are fully established and

managed carefully thereafter to prevent degradation of the

seeding or decline or loss of individual species. Monitoring

data can be used to evaluate seeding techniques and may

suggest improvements for future seedings.

The Future of Native Forb

Development ___________________

Native forb seed grown in agricultural fields or collected

from wildland stands is currently available for some species

(table 1). Supplies, however, remain limited and erratic. In

the last several years, threats to Great Basin ecosystems

have fostered efforts to develop plant materials and seed and

seeding technology for native forbs considered priority species by land management agencies. Personnel at State and

Federal laboratories are developing selections of individual
58 USDA Forest Service Proceedings RMRS-P-38. 2005

Walker and Shaw Current and Potential Use of Broadleaf Herbs for Reestablishing Native Communities Table 1—Forb characteristics and releasesa

.

Market

Adaptation availaAnnual Seeds Expected seed quality bility

precipi- per at time of purchase of

Species Soil tation pound Viability Purity Releases seedf

inches number g - - - - - percent - - - - 

Achillea millefolium E/M N P Loam to clay loam 8–26 2,124,000– B/D 85–90 20–90+ None 5

Western yarrow 4,124,000

Agoseris glauca M/L N P Moist, well drained >10 560,000 B/D 20–50; 20–60 None 2

Pale agoseris variable

Antennaria rosea M N P Coarse textured, >12 6,000,000 B 20–50 20–60 None 1

Rose pussytoes well drained

Artemisia ludoviciana E/M/L N P Sandy loam to clay loam >10 2,495,000– B/D 75–85 20–90 ‘Summit’ 2

Louisiana sage 2,500,000,

3,800,000

Aster chilensis M/L N P Coarse to clay loam >10 2,600,000 B/D 80–90 20–90 None 2

Pacific aster

Aster glaucodes M N P Loam to clay loam 12–16 540,000 B/D 80–90 20–90 None 2

Blueleaf aster

Astragalus cicer M I P Sandy loam to clay loam >15 114,000– 95 95 ‘Lutana’, 5

Cicer milkvetch 122,000, ‘Monarch’

130,000 ‘Windsor’

Balsamorhiza sagittata L N P Deep, well drained, 8–25 55,000 D 80–90 95–99 None 3

Arrowleaf balsamroot coarse to medium

 textured

Coronilla varia M I P Coarse to medium >26 109,760, D 75 95 ‘Emerald’ 5

Crownvetch textured, well drained, 138,160

 low fertility, low pH

Crepis acuminata M/L N P Sandy to coarse, 8–20 800,000 80–90 90 None 1

Tapertip hawksbeard gravelly soil

Eriogonum ovalifolium M/L N P Gravelly to sandy clay, 10 120,000 D/B 90 95–98 None 1

Oval-leaved eriogonum well drained

Eriogonum umbellatum M/L N P Well drained, medium 8–18 120,000, D 90 95–98 ‘Sierra’ 2

Sulfur eriogonum textured, slightly basic 209,000,

 to neutral pH 300,000

Hedysarum boreale M/L N P Dry, often rocky soil; 12–18 33,585, D 95 90 ‘Timp’ 3

Northern or well drained sandy loam 70,000,

Utah sweetvetch to basic, silty clay loam 86,000–

96,700,

125,000

Helianthella uniflora M N P Coarse, shallow, 10–35 52,560– B/D 60 75 None 1

Oneflower helianthella well drained 53,560,

267,000

(con.)

Ecological

statusb

Originc

Longevityd

Planting

requirementse
USDA Forest Service Proceedings RMRS-P-38. 2005 59

Current and Potential Use of Broadleaf Herbs for Reestablishing Native Communities Walker and Shaw Table 1—(Con.) Market Adaptation availa- Annual Seeds Expected seed quality bility precipi- per at time of purchase of Species Soil tation pound Viability Purity Releases seedf inches number g - - - - - percent - - - - Linum lewisii M/L N P Fine to coarse textured, 10–24 278,000– B/D 90 95–99 ‘Appar’ 5 Lewis flax well drained 422,325 Linum perenne M I P Fine to coarse textured, 10–24 278,280 B/D 90 95–99 None 2 Perennial flax well drained Lomatium dissectum M/L N P Dry, rocky >14 42,225, D 60–85 85 None 1 Fern-leaved lomatium 134,000 Lomatium nudicaule M N P Sandy B/D 60–85 85 None 1 Barestem lomatium Lomatium nuttallii M N P Light to heavy textured, 13–16 42,200– D 60–80 90 None 1 Nuttall lomatium well drained, basic 42,225 Lomatium triternatum M/L N P Well drained or dry, 8–20 42,000, D 70 75 None 2 Nineleaf lomatium rocky, acidic to basic 100,000 Lupinus argenteus E/M N P Fine to medium, coarse >10 18,300, D 95 90 None 2 Silvery lupine 126,000 Onobrychis viciifolia M I P Sandy loam to clay-loam 12–16 22,000– D 90–95 95–98 ‘Eski’, 5 Sainfoin 26,000 ‘Remont’ Penstemon acuminatus E/M N P Dry, open, sandy areas, 12–30 400,000, B/D 84 95 None 1 Sharpleaf penstemon often in sand dunes 592,379 Penstemon deustus E/M N P Basalt scabland, 10–18 400,000 B/D 90 95 None 1 Scabland penstemon volcanic soils Penstemon eatonii M N P Well drained, rocky to 8–16 350,000– B/D 90–95 95 Richfield 4 Eaton Penstemon sandy loam 400,000 Selected Penstemon pachyphyllus E N P Sandy or gravelly 12–16 336,000 B/D 70 95 None 1 Thickleaf beardtongue Penstemon palmeri E N P Well drained gravelly 10–16 609,675– B/D 85–90 95 ‘Cedar’ 4 Palmer penstemon to sandy loam 610,000, 294,000 Penstemon speciosus E/M N P Coarse-textured soils 8–24 400,000, B/D 66–97 95 None 1 Sagebrush penstemon 481,544 Penstemon strictus M N P Silty clay loam to 14–26 262,000, B/D 85–90 95 ‘Bandera’ 3 Rocky Mountain sandy loam 400,000 penstemon Phlox hoodii M/L N P Heavy clay, silty, B 60–80 None 1 Spiny phlox loamy, or coarse deep soils (con.)

Ecological

statusb

Originc

Longevityd

 Planting

requirementsc
60 USDA Forest Service Proceedings RMRS-P-38. 2005

Walker and Shaw Current and Potential Use of Broadleaf Herbs for Reestablishing Native Communities Table 1—(Con.) Market Adaptation availa- Annual Seeds Expected seed quality bility precipi- per at time of purchase of Species Soil tation pound Viability Purity Releases seedf inches number g - - - - - percent - - - - Sanguisorba minor E/M I P Well drained sandy 12-25 34,000– B/D 90 80–95+ ‘Delar’ 5 Small burnet loam to silty clay, 55,115 basic loams Sphaeralcea coccinea M N P Sandy loam to clay 6–35 500,000– D 90 80–90 ARS-2936 3 Scarlet globemallow 500,660, (with 800,000 gravity table) Sphaeralcea E N P Sandy loam to clay 8–14 500,000– D 90 80–90 ARS-2892 3 grossulariaefolia 500,660, (with Gooseberryleaf 850,000 gravity globemallow table) Trifolium repens M I P Moist, deep soils >20 672,000– B/D 80 95 None 5 White clover 880,000 Vicia americana L N P Sandy, clayey, >9 24,500– D 85–95 90–98 None 1 American vetch medium textured soils 41,314 Viguiera multiflora M N P Sandy loam to clay loam 15–20 800,000, B/D 40–60 90–95 None 2 Showy goldeneye 1,054,885– 1,055,000 a References: Dayton (1960); Davis and others (2002); Goodrich and Neese (1986); Hermann (1966); Hitchcock and Cronquist (1973); Johnson (1993); Jorgensen and Stevens (2004); Link (1993); Monsen (2000); Nold (1999); Plants for a Future (2004); Plummer and others (1968); Redente and others (1982); Rumbaugh (1983); Rumbaugh and others (1993); Shaw and Monsen (1983); Stevens and others (1996); Stevens and Monsen (1988); Stevens and others (1985); U.S. Department of Agriculture, Forest Service (1937, 2004); U.S. Department of Agriculture, Natural Resources Conservation Service (2004). b Ecological status: E = Early seral, M = Mid seral, L = Late seral, I = Introduced, no status. c Origin: N = Native, I = Introduced. d Longevity: P = perennial, B = biennial. e Planting: B = broadcast or aerially seed, D = drill seed. f Market availability: 5 = Excellent, large quanties, 4 = Good, generally sufficient, 3 = Fair, some seed available, 2 = Limited, small quantities may be available, 1 = Very limited, seed not usually available except by contract harvesting. g Literature values vary widely, possibly resulting from differences in seed cleaning practices and the structure considered the seed.

Ecological

statusb

Originc

Longevityd

 Planting

requirementsc
USDA Forest Service Proceedings RMRS-P-38. 2005 61

Current and Potential Use of Broadleaf Herbs for Reestablishing Native Communities Walker and Shaw

species. Additional research conducted by scientists at these

agencies and at universities provides seed and seeding

technology and seeding guidelines for selected species. Plant

materials developed through these efforts are made available to private growers for increase and for commercial seed

production. In some cases, USDA Forest Service, USDI

Bureau of Land Management, and other State and Federal

agencies have provided wildland seed collections directly to

growers for contract production of local seed sources for

specific projects. Private growers have also made sourceidentified seed collections from wildland sites and planted

them in seed fields. The challenge now for public and private

agencies is to set priorities for the seed needs by species and

population to direct research efforts and provide some stability for seed producers attempting to contend with the wide

annual fluctuations in postfire seedings as well as the

funding available for revegetation projects.

References _____________________

Barnett, J. F.; Crawford, J. A. 1994. Pre-laying nutrition of sagegrouse hens in Oregon. Journal of Range Management. 47:

114–118.

Beck, J. L.; Mitchell, D. L. 1997. Brief guidelines for maintaining

and enhancing sage-grouse habitat on private lands in Utah.

Utah Tech Notes. UT 190-7-3. Salt Lake City, UT: U.S. Department of Agriculture, Natural Resources Conservation Service. 9 p.

Braun, C. E.; Britt, T.; Wallstead, R. O. 1977. Guidelines for

maintenance of sage-grouse habitats. Wildlife Society Bulletin. 5:

99–106.

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.

Davis, K. M.; Englert, J. M.; Kujawski, J. L.; Scheetz, J. G. 2002.

Improved conservation plant materials released by NRCS and

cooperators through September 2002. Beltsville, MD: U.S. Department of Agriculture, Natural Resources Conservation Service, National Plant Materials Center. 57 p.

Dayton, William A. 1960. Notes on Western range forbs: Equisetaceae

through Fumariaceae. Washington, DC: U.S. Department of

Agriculture, Forest Service. 254 p.

Goodrich, S.; Neese, E. 1986. Uinta Basin flora. Ogden. UT: U.S.

Department of Agriculture, Forest Service, Intermountain

Region. 320 p.

Hermann, F. J. 1966. Notes on Western range forbs: Cruciferae

through Compositae. Agric. Handb. 293. Washington, DC: U.S.

Department of Agriculture, Forest Service. 365 p.

Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific

Northwest. Seattle: University of Washington Press. 730 p.

Johnson, C. G., Jr. 1993. Common plants of the inland Pacific

Northwest. Washington, DC: U.S. Department of Agriculture,

Forest Service, Pacific Northwest Region. 389 p.

Jorgensen, Kent R.; Stevens, Richard. 2004. Seed collection, cleaning, and storage. In: Monsen, Stephen B.; Stevens, Richard;

Shaw, Nancy L., comps. Restoring western ranges and wildlands.

Gen. Tech. Rep. RMRS-GTR-136-vol-3. Fort Collins, CO: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station: 699–716.

Link, E. 1993. Native plant propagation techniques for National

Parks—interim guide. East Lansing, MI: U.S. Department of

Agriculture, Soil Conservation Service and U.S. Department of

the Interior, National Park Service. 240 p.

Monsen, S. B. 2002. [E-mail to N. L. Shaw]. March 23. 1 leaf. On file

at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, ID; RWU 4253 files.

Nold, R. 1999. Penstemons. Portland, OR: Timber Press, Inc. 259 p.

Paige, C.; Ritter, S. A. 1999. Birds in a sagebrush sea: managing

sagebrush habitats for bird communities. Boise, ID: Partners in

Flight Western Working Group. 47 p.

Plants for a Future [Homepage of Plants for a Future], [Online].

(2004). Available: http://www.comp.leeds.ac.uk/pfaf/index.html.

[2004, February 10].

Plummer, A. P.; Christensen, D. R.; Monsen, S. B. 1968. Restoring

big-game range in Utah. Publ. 68-3. Salt Lake City, UT: Utah

Division of Fish and Game. 183 p.

Redente, E. F.; Ogle, P. R.; Hargis, N. E. 1982. Growing Colorado

plants from seed: a state of the art. Vol. III: Forbs. FWS/OBS-82/

30. Washington, DC: U.S. Department of the Interior, Fish and

Wildlife Service, Office of Biological Services, Western Energy

and Land Use Team. 141 p.

Rumbaugh, M. D. 1983. Legumes—their use in wildlife plantings.

In: Monsen, S. B.; Shaw, N., comps. Managing Intermountain

rangelands—improvement of range and wildlife habitats: proceedings; 1981 September 15–17; Twin Falls, ID; 1982 June 22–

24; Elko, NV. Gen. Tech. Rep. INT-157. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and

Range Experiment Station: 115–222.

Rumbaugh, M. D.; Pendery, B. M.; Mayland, H. F.; Shewmaker,

G. E. 1993. Registration of ARS-2936 scarlet globemallow. Crop

Science. 33: 1106–1108.

Shaw, N.; Monsen, S. B. 1983. Nonleguminous forbs for rangeland

sites. In: Monsen, S. B.; Shaw, N., comps. Managing Intermountain rangelands—improvement of range and wildlife habitats:

proceedings; 1981 September 15–17; Twin Falls, ID; 1982 June

22–24; Elko, NV. Gen. Tech. Rep. INT-157. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 123–131.

Stevens, R. 1983. Species adaptation for seeding mountain brush,

big, black, and low sagebrush, and pinyon-juniper communities.

In: Monsen, S. B.; Shaw, N., comps. Managing Intermountain

rangelands—improvement of range and wildlife habitats: proceedings; 1981 September 15–17; Twin Falls, ID; 1982 June 22–

24; Elko, NV. Gen. Tech. Rep. INT-157. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and

Range Experiment Station: 78–82.

Stevens, R.; Jorgensen, K. R.; Young, S. A.; Monsen, S. B. 1996. Forb

and shrub seed production guide for Utah. Logan: Utah State

University Extension. 51 p.

Stevens, R.; Monsen, S. B. 1988. Cedar Palmer penstemon: a selected

penstemon for semiarid ranges. Rangelands. 10: 163–164.

Stevens, R.; Shaw, N.; Howard, C. G. 1985. Important nonleguiminous forbs for Intermountain ranges. In: Carlson, J. R.;

McArthur, E. D., chairmen. Symposium: range plant improvement; 1985 February 11–15; Salt Lake City, UT. Denver, CO:

Society for Range Management: 210–220.

U.S. Department of Agriculture, Forest Service. 1937. Range plant

handbook. Washington, DC: U.S. Department of Agriculture,

Forest Service. Variously paginated.

U.S. Department of Agriculture, Natural Resources Conservation Service. 2004. The PLANTS Database, Version 3.1.

[Homepage of National Plant Data Center], Baton Rouge, LA

70874-4490 USA. [Online]. Available: http://plants.usda.gov

[2004, February 10].

U.S. Department of Agriculture, Forest Service. 2004. Fire Effects

Information System. [Online]. Available: http://www.fs.fed.us/

database/feis/ [2004, February 10].

Walker, S .C.; Stevens, R.; Monsen, S. B.; Jorgensen, K. R. 1995.

Interaction between native and seeded introduced grasses for 23

years following chaining of juniper-pinyon woodlands. In: Roundy,

B. A.; McArthur, E. D.; Haley, J. S.; Mann, D. K., comps. 1995.

Proceedings: wildland shrub and arid land restoration symposium; 1993 October 19–21; Las Vegas, NV. Gen. Tech. Rep. INTGTR-315. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station: 372–380.

Wallstead, R. O. 1971. Summer movements and habitat use by sagegrouse broods in central Montana. Journal of Wildlife Management. 35: 129–136.
62 USDA Forest Service Proceedings RMRS-P-38. 2005

Michael J. Wisdom is Research Wildlife Biologist, and Barbara C. Wales is

Wildlife Biologist, U.S. Department of Agriculture, Forest Service, Pacific

Northwest Research Station, Forestry and Range Sciences Laboratory, 1401

Gekeler Lane, La Grande, OR 97850. Mary M. Rowland is Wildlife Biologist,

USDI Bureau of Land Management, Forestry and Range Sciences Laboratory, 1401 Gekeler Lane, La Grande, OR 97850. Miles A. Hemstrom is

Research Ecologist, U.S. Department of Agriculture, Forest Service, Pacific

Northwest Research Station, Station Headquarters, 333 SW First Ave,

Portland, OR 97204.

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Landscape Restoration for

Greater Sage-Grouse:

Implications for Multiscale

Planning and Monitoring

Michael J. Wisdom

Mary M. Rowland

Miles A. Hemstrom

Barbara C. Wales

Abstract—Habitats and populations of greater sage-grouse

(Centrocercus urophasianus) have declined throughout western

North America in response to a myriad of detrimental land uses.

Successful restoration of this species’ habitat, therefore, is of keen

interest to Federal land agencies who oversee management of most

remaining habitat. To illustrate the challenges and potential for

landscape restoration, we summarized recent findings of restoration modeling for sage-grouse in the Interior Northwest. Changes in

amount and quality of habitat were evaluated under proposed

Federal management and under two restoration scenarios. Under

the two scenarios, the rate of habitat loss was reduced and the

quality of habitat was substantially improved compared to proposed management. These results have direct implications for

restoration planning and monitoring. First, a strategic, multiscale

approach is needed that links the scale of the stand with scales of the

seasonal, year-round, and multipopulation ranges of sage-grouse.

Second, consideration of connectivity across scales is essential.

Third, extensive and sustained use of a holistic suite of passive and

active restoration treatments is needed. And finally, monitoring of

both habitat and population responses across scales is critical. We

offer suggestions on these and related points for effective restoration planning and monitoring of sage-grouse habitat.

Introduction ___________________

Habitats and populations of greater sage-grouse

(Centrocercus urophasianus) have declined substantially

across the species’ range in response to a variety of detrimental land uses (Connelly and Braun 1997; Schroeder and

others 1999). New guidelines were developed recently

(Connelly and others 2000a) to help managers conserve and

restore habitats for the species at the stand scale, but similar

guidelines do not exist for landscape scales that encompass

all or major portions of the species’ range. The cumulative

effects of management at these larger scales can greatly

influence regional extirpation of sage-grouse (Raphael and

others 2001), and recent landscape evaluations (Hemstrom

and others 2002; Raphael and others 2001; Wisdom and

others 2000, 2002a,b) offer new insights for effective restoration planning across the species’ range.

The prospect of continued and widespread habitat declines for sage-grouse and other sagebrush (Artemisia spp.)

obligates (Raphael and others 2001; Wisdom and others

2000, 2002a) points to the urgent need for development of

restoration efforts across large landscapes. Without such

restoration efforts, continued management of Federal lands

under current land use plans will likely result in further loss

and degradation of sagebrush steppe, with an increasingly

high risk of population extirpation for sagebrush-dependent

species (Raphael and others 2001).

In this paper, we summarize results of recent landscape

evaluations to restore habitats for sage-grouse on lands

administered by the U.S. Department of Agriculture, Forest

Service, and U.S. Department of the Interior, Bureau of

Land Management (FS-BLM) in the Interior Columbia Basin and adjacent portions of the Great Basin (Basin) (fig. 1).

The 58 million-ha Basin encompasses a major portion of

current and historical range of greater sage-grouse (fig. 1)

(Wisdom and others 2002a). Proposed management of the

Basin’s sagebrush steppe will therefore substantially affect

sage-grouse and other sagebrush obligates. That as context,

our goals were to summarize the conditions projected for

greater sage-grouse from a previous study within the Basin

(Raphael and others 2001) in relation to two restoration

scenarios recently developed and evaluated by Hemstrom

and others (2002) and Wisdom and others (2002a) and to

place the results in appropriate biological context for management of sage-grouse, particularly in terms of multiscale

land use planning and monitoring.
USDA Forest Service Proceedings RMRS-P-38. 2005 63

Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . . Wisdom, Rowland, Hemstrom, and Wales

Restoration Scenarios __________

Raphael and others (2001) evaluated the effects of proposed management of FS-BLM lands, projected 100 years in

the future, on sage-grouse and other vertebrates that depend on sagebrush steppe in the Basin. These effects were

associated with three management alternatives proposed in

the Supplemental Environmental Impact Statement (SEIS)

(USDA-FS and USDI-BLM 2000) of the Interior Columbia

Basin Ecosystem Management Project (ICBEMP). Under

proposed management, as well as current land management, Raphael and others (2001) found that most species

that depend on sagebrush steppe, including sage-grouse,

had a high probability of local or regional extirpation.

Hemstrom and others (2002) and Wisdom and others

(2002a) evaluated the benefits of dramatically increasing

the extent and intensity of restoration in sagebrush steppe

to gain insight into the potential for improving environmental conditions for sage-grouse and thereby reducing the risk

of extirpation compared to that under proposed management. Evaluations were based on Hemstrom and others’

(2002) modeling of two restoration scenarios that substantially increased the combination of passive and active

restoration of sagebrush steppe within the historical range

of sage-grouse in the Basin.

As the basis for the two scenarios, Hemstrom and others

(2002) defined passive restoration as “the process of modifying or eliminating existing management activities (for example, livestock grazing, roads, or recreation) that contribute to environmental degradation of desired resources.” In

contrast, Hemstrom and others (2002) defined active restoration as “the application of treatments that contribute to

recovery of targeted resources (for example, appropriate

use of wildfire suppression, prescribed fire, or seeding with

native plants).” These definitions are similar to those

summarized for rangeland restoration by McIver and Starr

(2001).

The two scenarios substantially increased the levels of

both passive and active restoration in relation to proposed

management because of managers’ desire to understand the

magnitude by which sagebrush habitats could be improved

relative to what was originally proposed. Scenario 1 assumed a 50 percent reduction in detrimental grazing effects

by livestock as the main form of passive restoration. Detrimental grazing effects were defined as the probability,

associated with grazing, of moving from a desired vegetation

state, which provides habitat for sage-grouse (for example,

Figure 1—The Interior Columbia Basin assessment area in the Western United States,

encompassing eastern Washington (WA), eastern Oregon (OR), most of Idaho (ID),

northwestern Montana (MT), and adjacent areas of northwestern Wyoming (WY),

northwestern Utah (UT), and northern Nevada (NV), and the historical habitats of

greater sage-grouse within the species’ historical range in the Basin (from Wisdom and

others 2000).

WA

MT

ID

OR

WY

NV UT

Historical habitat of 

greater sage-grouse 

within the species’ 

historical range in Basin 

OR
64 USDA Forest Service Proceedings RMRS-P-38. 2005

Wisdom, Rowland, Hemstrom, and Wales Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . .

gray boxes, fig. 2), to an undesired state (for example, white

boxes, or nonhabitat, fig. 2). Accordingly, 50 percent and 100

percent reductions in detrimental grazing effects represented like reductions in the probability of transitioning

from desired to undesired states for sage-grouse in relation

to livestock grazing. Detailed rationale and supporting literature regarding these grazing effects on sagebrush habitats, and on sage-grouse, are described in Hemstrom and

others (2002).

To achieve reductions of 50 percent and 100 percent in

detrimental grazing effects, like reductions in stocking rate

of livestock were assumed in combination with additional,

positive changes in grazing systems (for example, increasing

rest periods in rest-rotation systems) (Hemstrom and others

2002). This form of passive restoration under scenario 1 was

applied to 6.4 million ha of FS-BLM lands in the Basin that

have potential to be sage-grouse habitat or that currently

serve as habitat (referred to as potential sage-grouse habitat). Two points are important here. First, not all grazing

effects were assumed by Hemstrom and others (2002) to be

detrimental to sage-grouse habitat. However, there is growing awareness that the herbaceous component of sagebrush

stands, which can be reduced substantially in occurrence

and percent cover with intensive livestock grazing (Anderson and Inouye 2001), is a primary requirement for successful nesting and brood rearing by sage-grouse (Barnett and

Crawford 1994; Connelly and others 2000a; Crawford 1997).

Consequently, there is a need to mitigate the detrimental

effects of livestock grazing on native grasses and forbs

important to sage-grouse productivity. And second, reduction in stocking rate of livestock can effectively restore

native, herbaceous components in sagebrush steppe (Anderson and Inouye 2001). In defense of this point, Hemstrom

and others (2002) stated

Our assumed reductions in stocking rate needed to achieve

a desired reduction in detrimental grazing were based on

empirical data demonstrating that herbage production on

rangelands is affected mostly by variation in stocking rate,

and less so by changes in grazing system (Holechek and others

1998; Van Poolen and Lacey 1979). On arid rangelands such as

those dominated by sagebrush, a positive response in herbage

production must include a reduction in stocking rate in combination with active restoration treatments (see empirical

synthesis by Holechek and others 1998).

Active restoration under scenario 1 was integrated with

passive restoration on the same 6.4 million ha of potential

sage-grouse habitat (Hemstrom and others 2002). By contrast, active restoration under proposed management targeted approximately 1.1 million ha of potential sage-grouse

habitat. Thus, scenario 1 represented a sixfold increase in

areas treated with active restoration beyond that identified

in proposed management.

Key forms of active restoration included seedings and

plantings of desired vegetation, particularly after fire events;

wildfire suppression in vegetation types where such fires

would facilitate invasion of exotic plants; prescribed fire in

vegetation types where such fires would reduce woodland

encroachment; and use of a variety of other chemical and

mechanical treatments to control invading conifers and

enhance composition of native grasses and forbs (Hemstrom

and others 2002). The specific combination of active restoration treatments was tailored to the unique, desired response

of each sagebrush community to the treatments. For example, use of prescribed fire to suppress juniper (Juniperus

spp.) invasion, and enhance growth of herbaceous vegetation, was applied to many areas dominated by mountain big

sagebrush (A. tridentata ssp. vaseyana), where fire effects

are largely beneficial (Miller and Eddleman 2000). By

contrast, suppression of wildfire, in combination with chemical treatments and native seedings to control spread of

exotic grasses, was applied to many areas dominated by

Wyoming big sagebrush (A. tridentata ssp. wyomingensis),

where cheatgrass (Bromus tectorum) and other annuals

often supplant native vegetation following fire events (Miller

and Eddleman 2000).

Restoration scenario 2 was based on a 100 percent reduction in detrimental grazing effects by livestock, with a like

reduction in stocking rate (Hemstrom and others 2002). This

high level of passive restoration was integrated with the

same level of active restoration assumed for scenario 1, with

the same 6.4 million ha of FS-BLM lands targeted for

treatment. Detailed methods, assumptions, and rationale

associated with the scenarios are described in Hemstrom

and others (2002) and Wisdom and others (2002a).

Conditions under the two restoration scenarios were projected 100 years into the future, as was done for proposed

management. Restoration activities for each scenario were

sustained throughout the 100-year period, with the frequency, intensity, and type of each activity designed to

substantially recover or maintain desired conditions

(Hemstrom and others 2002). Three landscape variables for

sage-grouse were targeted for improvement as part of the

restoration scenarios: (1) habitat amount, and two indices of

habitat quality, (2) HRV departure (an acronym for historical range of variability departure, as defined by Hann and

others 1997), and (3) uncharacteristic grazing (Hemstrom

and others 2002; Wisdom and others 2002a).

Habitat amount is the area of sage-grouse habitat within

the Basin, as defined by Wisdom and others (2000). Sagegrouse habitats in the study area primarily include low- to

medium-height shrublands in basin big sagebrush (Artemisia tridentata ssp. tridentata), Wyoming big sagebrush,

mountain big sagebrush, and low sagebrush (A. arbuscula)

communities, as well as herbaceous wetlands.

HRV departure was used to index the degree to which

exotic plants have invaded and displaced components of

native sagebrush steppe, particularly native grasses and

forbs that are required by sage-grouse for successful nesting

and brood rearing (Connelly and others 2000a; Crawford

1997; Drut and others 1994; Sveum and others 1998). Uncharacteristic grazing, (UG) was used to index changes in

species richness, height, and cover of native understory

grasses and forbs in response to livestock grazing, and to

subsequent effects on quality of nesting and brood-rearing

habitat for sage-grouse (Wisdom and others 2002a).

Restoration activities were designed to enhance habitat

quantity (through increased habitat amount) and quality

(through reductions in HRV departure and uncharacteristic

grazing). Restoration was particularly designed to retard

the cheatgrass (Bromus tectorum)-wildfire cycle (fig. 2), a

pervasive problem in the Wyoming big sagebrush communities that compose >60 percent of sage-grouse habitat in the

Basin (Hemstrom and others 2002).
USDA Forest Service Proceedings RMRS-P-38. 2005 65

Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . . Wisdom, Rowland, Hemstrom, and Wales Succession, 21 y Agropyron/Poa Closed herbland Wyoming big sage Open low shrub Fire, 0.01 Succession, 30 y Poa/Festuca Closed herbland Grazing, 0.025 Fire, 0.01 Grazing, 0.021 Fire, 0.008 Cheatgrass Open herbland Wyoming big sage/Poa Open low shrub Succession, 1000 y Grazing + exotics, 0.011 Succession, 31 y Succession, 31 y Fire, 0.008 Grazing, 0.01 Wyoming big sage/cheatgrass Open low shrub Grazing + exotics, 0.011 Seeded native grass Closed herbland Herbicide and seed native species, 0.01 Exotic perennial grasses Closed herbland Herbicide and seed exotic species, 0.0001 Succession, 1000 y Fire, 0.05 Succession, 81 y Fire, 0.05 Herbicide and seed exotic species, 0.0001Threshold Figure 2—Example state-and-transition model (based on concepts of Westoby and others 1989 and Laycock 1991) for the potential vegetation type of warm, dry Wyoming big sagebrush (defined by Hann and others 1997) under an active restoration prescription in the Interior Columbia Basin (from Hemstrom and others 2002). Boxes are vegetation states and arrows are transitions from one state to another. In this example, transitions are brought about by succession and disturbance agents of fire, grazing, chemical treatments, and seedings. When disturbance is the primary cause for transition, the annual probability of moving from one state to another is shown in association with the disturbance that causes the transition (for example, fire, 0.01). Successional transitions of 1,000 years (y) indicate stable states. The dotted line indicates a transition threshold, below which restoration is difficult and requires a sustained, extensive combination of active and passive treatments.
66 USDA Forest Service Proceedings RMRS-P-38. 2005

Wisdom, Rowland, Hemstrom, and Wales Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . .

Methods used to model these improvements under the

restoration scenarios were deliberately conservative in terms

of the assumed enhancements that such activities could

produce. A conservative modeling approach was adopted

because of the high uncertainty of restoration outcomes in

sagebrush steppe (West 1999). This high uncertainty is

related to incomplete knowledge of appropriate restoration

methods and technologies, and the logistical challenges

posed by sustained and integrated application of restoration

treatments across vast areas of sagebrush steppe, which to

date has not been attempted (Knick 1999).

Results from the restoration modeling were evaluated in

terms of risk of regional extirpation of sage-grouse, as

expressed in five outcome classes (see population outcome

model described by Wisdom and others 2002b). Outcome A

was defined as a very low risk of regional extirpation,

followed by low (outcome B), moderate (outcome C), high

(outcome D), and very high (outcome E) degrees of risk.

These levels of risk corresponded to empirical findings of

Wisdom and others (2002b), showing that areas of the Basin

historically occupied by sage-grouse were associated with

outcome A, whereas areas of current extirpation were associated with outcome E. Moreover, areas of the Basin currently occupied by sage-grouse have undergone an intermediate level of habitat loss and degradation between that

estimated for historically occupied areas versus currently

extirpated areas, resulting in an intermediate outcome of

class C (Wisdom and others 2002b).

Risk of extirpation was assessed for FS-BLM lands and for

all lands. The five outcome classes that indexed risk on FSBLM lands were referred to as environmental outcomes

(Raphael and others 2001).

Restoration Effects _____________

Results from Hemstrom and others (2002) showed that

under proposed management, sage-grouse habitat on FSBLM lands would decline by 27 percent compared with the

current amount (weighted average of percent declines across

the sagebrush communities shown in fig. 3). However, habitat declined more slowly under restoration scenarios 1 and

2 (by about 19 percent and 17 percent, respectively), but

neither scenario halted the long-term downward trend.

Most future habitat loss was associated with sagebrush

transitions to herblands and grasslands dominated by

cheatgrass and other exotic plants in large areas of the

Wyoming big sagebrush communities. Substantially smaller

habitat losses were projected in the future in mountain big

sagebrush communities, with losses due mostly to encroachment by juniper. In mountain big sagebrush communities,

however, some loss to exotic plant invasion was projected at

lower elevation, drier sites, while loss to woodland and forest

encroachment was projected at higher elevation, mesic sites.

Additionally, small declines in habitat amount were projected for other sagebrush communities, such as low sagebrush (Hemstrom and others 2002).

Restoration scenarios 1 and 2 increased habitat amount,

relative to proposed management, by about 0.6 million ha

and 0.8 million ha, respectively. The model projections

indicated that a substantial increase in habitat from passive

and active restoration would be offset by large (>1 million ha)

losses associated mostly with wildfire and the subsequent

invasion of cheatgrass in the Wyoming big sagebrush communities.

In contrast to results for habitat amount, the quality of

habitat improved substantially under the restoration scenarios compared with proposed management, as indexed by

substantial reductions in UG and HRV departure (fig. 4).

Only 22 percent and 12 percent of subwatersheds were

characterized by high UG under scenarios 1 and 2, whereas

highUG occurred in 68 percent and 53 percent of subwatersheds

during the current period and under proposed management,

respectively. Percentage of subwatersheds with high HRV

departure under the restoration scenarios (2 percent) also

was substantially lower than the percentage with high HRV

departure currently (6 percent) and under proposed management (7 percent). The restoration scenarios also were

associated with a higher percentage of subwatersheds in the

low and none classes of UG and HRV departure compared to

current conditions and proposed management (fig. 4).

Risk of sage-grouse extirpation on FS-BLM lands was

reduced to a moderate level under the two restoration

scenarios compared to a high risk under proposed management (fig. 5). The moderate risk of extirpation under the

restoration scenarios was the same as that estimated for the

current period (fig. 5). The difference between a moderate

versus a high risk of extirpation, as evaluated under the

outcome classes for sage-grouse, was found by Wisdom and

others (2002b) to represent a substantial difference in the

probability of regional extirpation for the species.

Three landscape variables contributed to the increased

risk of sage-grouse extirpation under proposed management

(Wisdom and others 2002a): (1) reduced habitat quantity

and quality, as reflected in an overall reduction in habitat

capacity; (2) increased contraction of the species’ range,

-35

-30

-25

-20

-15

-10

-5

0

Percent change

Wyoming big sagebrush Mountain big sagebrush

Current

Proposed 

Management

Restoration

Scenario 1

Restoration

Scenario 2

Figure 3—Percent change in habitat amount for sagegrouse, by major sagebrush communities, in 1,831 FSand BLM-dominated subwatersheds within the historical

range of sage-grouse in the Interior Columbia Basin

(adapted from Hemstrom and others 2002). Results for

proposed management (PM) and the restoration scenarios

are for 100 years in the future. Decline is relative to

amount of habitat estimated for historical conditions (circa

1850–1900) (Hann and others 1997).
USDA Forest Service Proceedings RMRS-P-38. 2005 67

Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . . Wisdom, Rowland, Hemstrom, and Wales

owing to continued habitat loss; and (3) decreased connectivity of habitats that remained within the contracted range.

Wisdom and others (2002a) provide details.

Implications for Multiscale Planning

and Monitoring ________________

Results of restoration modeling by Hemstrom and others

(2002) and Wisdom and others (2002a) have substantial and

direct implications for management of sage-grouse habitats.

We offer the following suggestions for planning and monitoring restoration activities that follow directly from results of

the restoration scenarios:

1. A strategic, multiscale approach is needed that

links the scale of individual sagebrush stands with

scales of the seasonal, year-round, and multipopulation ranges of sage-grouse. Identification of nesting and brood-rearing areas is critical for effective management of sage-grouse summer range. Similarly, identification

of wintering areas is important for maintaining conditions

adequate for winter survival. Information about conditions

of individual sagebrush stands within seasonal ranges, as

well as information about overall conditions on seasonal

ranges, can be synthesized at larger scales to evaluate

composite conditions for the species on a year-round basis.

In turn, this information can be further synthesized to scales

of local or multiple populations of sage-grouse, with patterns

identified and summarized at larger, regional scales, such as

the Snake River Plain or Columbia River Plateau. At each

scale, relevant information is available for identifying management threats, setting restoration priorities, and implementing and monitoring a desired suite of restoration activities. While local areas are the traditional focus for restoration

planning and implementation, the larger, regional scales

are a critical and effective complement to local work. Information at regional scales, for example, can be used to target

large areas that may deserve high priority for restoration

and monitoring. By contrast, information about conditions

on seasonal ranges, or of individual stands within seasonal

ranges, is important for effective implementation of local

restoration priorities.

2. Consideration of connectivity across scales is

essential.Connectivity of summer range with winter range,

and of local populations with multiple populations, is critical

for maintaining viable populations of sage-grouse across the

species’ range. Raphael and others (2001) and Wisdom and

others (2002a,b) used a landscape method to evaluate connectivity of sage-grouse habitats in the Basin as part of their

population outcome model of extirpation risk. The method

assessed the degree to which subwatersheds containing

sage-grouse habitat fell within the median dispersal distance of juvenile grouse. This measure of connectivity was

later validated as an important landscape measure of extirpation risk (Wisdom and others 2002b). Specifically, the

connectivity of subwatersheds in areas currently occupied

by sage-grouse was 61 percent (on a scale of 0 to 100 percent,

where 100 percent represents habitats that are fully connected across the range of the species). By contrast, connectivity in areas where sage-grouse have been extirpated

was only 23 percent (Wisdom and others 2002b). Similar

0

10

20

30

40

50

60

Current PM RS1 RS2

Subwatersheds (%) 

High

Moderate

Low

None

0

10

20

30

40

50

60

70

80

90

Current PM RS1 RS2

Subwatersheds (%)

Uncharacteristic grazing

HRV departure

Class

Figure 4—Changes in habitat quality for sagegrouse across time, as indexed by classes of

uncharacteristic grazing and HRV departure in

1,831 FS- and BLM-dominated subwatersheds

within the historical range of sage-grouse in the

Interior Columbia Basin (from Hemstrom and others

2002). Results for proposed management (PM)

and the two restoration scenarios (RS1 and RS2)

are for 100 years in the future.

Historical Current PM RS1 RS2

Environmental outcome, FS-BLM lands

Alternative/time period

Outcome class

A

B

C

D

E

Very Low

Low

Moderate

High

Very High

Extirpation risk

Figure 5—Risk of regional population extirpation

for sage-grouse in the Interior Columbia Basin

(adapted from Wisdom and others 2002a), as

indexed by classes of environmental outcome (A =

very low risk, B = low, C = moderate, D = high, and

E = very high) projected for historical and current

periods and 100 years in the future under proposed

management (PM) and two restoration scenarios

(RS1 and RS2). Results for environmental outcomes

are for Forest Service (FS) and Bureau of Land

Management (BLM) lands. Historical, current, and

PM results are from Raphael and others (2001).
68 USDA Forest Service Proceedings RMRS-P-38. 2005

Wisdom, Rowland, Hemstrom, and Wales Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . .

measures of connectivity need development and validation

at a variety of scales to allow managers to understand how

well restoration plans might improve the connectivity of

habitat for sage-grouse, and to monitor the population response of sage-grouse to presumed improvements in connectivity. Development and validation of such connectivity

measures will be most successful if conducted as a partnership between land managers and scientists, owing to the

absence of research on this topic and the challenges of

management application at multiple scales.

3.Sustained use of a comprehensive suite of passive

and active restoration treatments over extensive areas is needed. Hemstrom and others (2002) and Wisdom

and others (2002a) found that restoration of sagebrush

habitats will require monumental spatial and temporal

scales of application if downward trends are to be slowed or

reversed. Expansive and sustained habitat restoration can

maintain desired conditions and reduce the future risk of

sage-grouse extirpation on FS-BLM lands. Local restoration

efforts, without coordination and implementation across

large areas as an adaptive management experiment, appear

to have a low probability of reducing extirpation risk for

sage-grouse in the Basin. This is due to the vast areas over

which restoration must occur and the comprehensive, integrated manner in which a suite of restoration treatments

must be implemented (Knick 1999). Knowledge voids about

effective methods of restoration pose a major challenge. For

example, few methods exist for effective restoration of native

forbs in sagebrush habitats, and these forbs are critical for

successful nesting and brood rearing by sage-grouse (Barnett

and Crawford 1994; Drut and others 1994). Continued

spread of exotic plants presents a formidable challenge to

successful restoration, and warrants substantial research

and management attention. In particular, the lower elevation, Wyoming big sagebrush communities are most susceptible to future loss from wildfire and subsequent invasion by

cheatgrass. These areas warrant special attention for restoration activities. Moreover, results from Hemstrom and others

(2002) suggest that suppression of wildfire, combined with

improvements in grazing management, are critical for preventing expansive conversions of sagebrush to cheatgrass in Wyoming big sagebrush communities (fig. 2).

4. Monitoring of both habitat and population responses across scales is critical. Three types of monitoring have been defined and used by Federal land management agencies: (1) implementation, (2) effectiveness, and

(3) validation monitoring. Implementation monitoring is the

assessment of whether restoration and other management

actions are implemented in the manner specified. By contrast, effectiveness monitoring evaluates whether the desired results from implementation were achieved, while

validation monitoring determines the scientific validity of

the concepts, methods, and predictions associated with the

expected benefits of the management actions. For sagegrouse, all three types of monitoring are needed. For example, goals may be set for improving the amount, quality,

and distribution of sage-grouse habitat under a restoration

plan. The primary goal may be to improve habitat attributes,

but invariably, the ultimate goal of such plans is to increase

population growth of sage-grouse and associated species.

Measuring such population responses will require regional

scales of monitoring, and are best accomplished as part of

research. Nonetheless, implementation monitoring is needed

to determine whether the treatments are applied in the

manner specified. Moreover, effectiveness monitoring is

needed for two purposes: (1) to assess whether the desired

habitat improvements were achieved with successful implementation (for example, were the desired improvements in

habitat amount, quality, and distribution actually accomplished?); and (2) to determine whether the associated population of sage-grouse responded positively from the habitat

improvements (for example, did the improvements in composition of understory bunchgrasses and forbs increase nest

success and brood survival?). Finally, validation monitoring

may be needed to understand why certain habitat restoration efforts might have failed, or how such restoration efforts

worked successfully. Unfortunately, nearly all management

and research of sage-grouse has focused on habitats or

populations, but not both. Consequently, few monitoring

efforts have considered effects on both habitats and populations, and considered all three types of monitoring. A notable

exception is Connelly and others’ (2000b) monitoring of sagegrouse response to prescribed burning in southeastern Idaho;

this integrated type of habitat and population monitoring is

needed to guide future restoration work. The model of

Edelmann and others (1998), which evaluates the quality of

sage-grouse habitat and predicts effects on growth rate of

sage-grouse populations, provides a comprehensive framework for conducting all three types of monitoring at stand

and landscape scales.

5.A comprehensive set of species that depend on the

sagebrush ecosystem needs to be targeted for restoration planning and monitoring. A common management

assumption about restoration efforts for sage-grouse is that

such efforts will confer like benefits to a larger set of plants

and animals that depend on sagebrush steppe. This assumption, however, has not been evaluated with empirical research and needs testing at multiple scales (Rich and Altman

2001). Moreover, new approaches that explicitly consider

the needs of a comprehensive set of species need to be

developed for effective restoration planning for all key attributes of the sagebrush ecosystem. Recently, Wisdom and

others (2002c) developed a habitat network for a large set of

vertebrates of conservation concern that depend on sagebrush steppe in the Basin. Watersheds for these species

were characterized as one of three habitat conditions: Condition 1—habitats of high resiliency, abundance, and quality; Condition 2—habitats of high abundance but moderate

resiliency and quality; and Condition 3—habitats that are

highly degraded, fragmented, and isolated, or that have

been extirpated. This type of characterization of a comprehensive, multispecies habitat network could be used to

maintain habitats in a relatively unchanged state from

historical conditions (Condition 1), to improve habitats where

quality and resiliency have declined (Conditions 2 and 3), to

restore habitats in areas of extirpation or low abundance

and quality (Condition 3), and to improve connectivity where

spatial gaps have developed (Condition 3).
USDA Forest Service Proceedings RMRS-P-38. 2005 69

Landscape Restoration for Greater Sage-Grouse: Implications for Multiscale Planning . . . Wisdom, Rowland, Hemstrom, and Wales

Conclusions___________________

Restoration of sage-grouse habitat represents a daunting

task. Without consideration of multiple scales of planning

and monitoring, chances for success may be substantially

reduced. New approaches that integrate a holistic suite of

restoration treatments, including changes in management

of livestock, are essential. Results of recent restoration

modeling for sage-grouse provide a starting point for development of multiscale strategies that could facilitate effective

recovery of key habitats across large areas of sage-grouse

range. In addition to restoration planning and monitoring

for sage-grouse, efforts that consider a comprehensive set of

sagebrush-dependent species are needed. An example is the

development of a habitat network and related multispecies

approaches for restoration planning (Wisdom and others

2002c), which could facilitate a more holistic recovery of the

sagebrush ecosystem. Without such efforts, managers will

be faced with a high likelihood of continued habitat loss and

increasing extirpation risk for species that depend on sagebrush habitats.

Acknowledgments _____________

We thank T. Quigley, T. Mills, and R. Graham for their

longstanding support of this work. We also thank A. Miller,

C. McCarthy, and N. Shankar for mapping and analysis, and

J. Augsburger, S. Sather-Blair, and G. Buckner for comments

on our paper. Our work was funded by the U.S. Department

of Agriculture, Forest Service, and the U.S. Department of

the Interior, Bureau of Land Management.

References ____________________

Anderson, J. E.; Inouye, R. S. 2001. Landscape-scale changes in

plant species abundance and biodiversity of a sagebrush steppe

over 45 years. Ecological Monographs. 71: 531–556.

Barnett, J. K.; Crawford, J. A. 1994. Pre-laying nutrition of sagegrouse hens in Oregon. Journal of Range Management. 47:

114–118.

Connelly, J. W.; Braun, C. E. 1997. Long-term changes in sagegrouse Centrocercus urophasianus populations in western North

America. Wildlife Biology. 3: 229–234.

Connelly, J. W.; Reese, K. P.; Fischer, R. A.; Wakkinen, W. L. 2000b.

Response of a sage-grouse breeding population to fire in southeastern Idaho. Wildlife Society Bulletin. 28: 90–96.

Connelly, J. W.; Schroeder, M. A.; Sands, A. R.; Braun, C. E. 2000a.

Guidelines to manage sage-grouse populations and their habitats. Wildlife Society Bulletin. 28: 967–985.

Crawford, J. A. 1997. Importance of herbaceous vegetation to female

sage-grouse Centrocercus urophasianus during the reproductive

period: a synthesis of research from Oregon, USA. Wildlife Biology. 3: 271.

Drut, M. S.; Crawford, J. A.; Gregg, M. A. 1994. Brood habitat use

by sage-grouse in Oregon. Great Basin Naturalist. 54: 170–176.

Edelmann, F. B.; Ulliman, M. J.; Wisdom, M. J.; Reese, K. P.;

Connelly, J. W. 1998. Assessing habitat quality using population

fitness parameters: a remote sensing based habitat-explicit model

for sage-grouse. Tech. Bull. 25. Moscow: University of Idaho,

Forest, Wildlife, and Range Experiment Station. 33 p.

Hann, W. J.; Jones, J. L.; Karl, M. G.; Hessburg, P. F., Keane, R. E.;

Long, D. G.; Menakis, J. P.; McNicoll, C. H.; Leonard, S. G.;

Gravenmier, R. A.; Smith, B. G. 1997. Landscape dynamics of the

basin. In: Quigley, T. M.; Arbelbide, S. J., tech. eds. An assessment

of ecosystem components in the Interior Columbia Basin and

portions of the Klamath and Great Basins. Gen. Tech. Rep. PNWGTR-405 Vol. II. Portland, OR: U.S. Department of Agriculture,

Forest Service, Pacific Northwest Research Station: 337–1055.

Hemstrom, M. A.; Wisdom, M. J.; Rowland, M. M.; Wales, B.; Hann,

W. J.; Gravenmier, R. A. 2002. Sagebrush-steppe vegetation

dynamics and potential for restoration in the Interior Columbia

Basin, USA. Conservation Biology. 16: 1243–1255.

Holechek, J. L.; Pieper, R. D.; Herbel, C. H. 1998. Range management: principles and practices. 3rd ed. Upper Saddle River, NJ:

Prentice Hall. 542 p.

Knick, S. T. 1999. Requiem for a sagebrush ecosystem? Northwest

Science. 73: 53–57.

Laycock, W. A. 1991. Stable states and thresholds of range condition

on North American rangelands: a viewpoint. Journal of Range

Management. 44: 427–434.

McIver, J. L.; Starr, L. 2001. Restoration of degraded lands in the

Interior Columbia River Basin: passive vs. active approaches.

Forest Ecology and Management. 153: 15–28.

Miller, R. F.; Eddleman, L. L. 2000. Spatial and temporal changes

of sage-grouse habitat in the sagebrush biome. Tech. Bull. 151.

Corvallis: Oregon State University, Agricultural Experiment

Station. 35 p.

Raphael, M. G.; Wisdom, M. J.; Rowland, M. M.; Holthausen, R. S.;

Wales, B. C.; Marcot, B. M.; 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.

Rich, T.; Altman, B. 2001. The greater sage-grouse as an umbrella

species. Grouse Partnership News. 2: 20.

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: life histories for the 21st century. Vol. 11. No.

425. Philadelphia, PA: The Birds of North America, Inc. 28 p.

Sveum, C. M.; Edge, W. D.; Crawford, J. A. 1998. Nesting habitat

selection by sage-grouse in south-central Washington. Journal of

Range Management. 51: 265–269.

U.S. Department of Agriculture, Forest Service, and U.S. Department of the Interior, Bureau of Land Management (USDA-FS

and USDI-BLM). 2000. Interior Columbia Basin supplemental

environmental impact statement, BLM/OR/WA/Pt-00/019+1792.

Portland, OR: U.S. Bureau of Land Management. variously

paginated.

Van Poolen, H. W.; Lacey, J. R. 1979. Herbage response to grazing

systems and stocking intensities. Journal of Range Management.

32: 250–253.

West, N. E. 1999. Managing for biodiversity of rangelands. In:

Collins, W. W.; Qualset, C. O., eds. Biodiversity in agroecosystems. Boca Raton, FL: CRC Press: 101–126.

Westoby, M.; Walker, B.; Noy-Meir, I. 1989. Opportunistic management for rangelands not at equilibrium. Journal of Range Management. 42: 266–276.

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. R. 2000. Source habitats for terrestrial

vertebrates of focus in the Interior Columbia basin: broad-scale

trends and management implications. Gen. Tech. Rep. PNWGTR-485. Portland, OR: U.S. Department of Agriculture, Forest

Service, Pacific Northwest Research Station.

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.

Wisdom, M. J.; Wales, B. C.; Holthausen, R. S.; Hann, W. J.; Hemstrom,

M. A.; Rowland, M. M. 2002c. A habitat network for terrestrial

wildlife in the Interior Columbia Basin. Northwest Science. 76:

1–14 .
70 USDA Forest Service Proceedings RMRS-P-38. 2005

John A. Fairchild is Habitat Conservation Coordinator, Utah Division of

Wildlife Resources, P.O. Box 146301, Salt Lake City, UT 84114-6301 U.S.A.;

e-mail: johnfairchild@utah.gov. James N. Davis is Project Leader, Big Game

Range Trend Study, Utah Division of Wildlife Resources, 735 North 500 East,

Provo, UT 84606-1865 U.S.A.; e-mail: jimdavis@utah.gov. Jack D. Brotherson

is Professor, Department of Integrative Biology, Brigham Young University,

Provo, UT 84602 U.S.A.; e-mail: jack_brotherson@byu.edu

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Big Sagebrush Response to

One-Way and Two-Way Chaining

in Southeastern Utah

John A. Fairchild

James N. Davis

Jack D. Brotherson

Abstract—A decadent, mixed stand of Wyoming big sagebrush,

Artemisia tridentata wyomingensis, and mountain big sagebrush,

Artemisia tridentata vaseyana, located north of Cisco, Utah, was

subjected to one-way and two-way chaining treatments in November 1987. The effect of the treatments on plant community characteristics and shrub vigor was documented over a 3-year period.

Stand density was reduced 60 percent on sites chained two ways

and 43 percent on sites chained over once. Shrubs on one-way

chained sites produced more leader growth in 1989 and 1990 than

those on untreated sites or sites chained two ways. Browse production on one-way chained sites surpassed that of untreated sites and

two-way chained sites by 140 percent and 350 percent, respectively.

Over the short term, a one-way chaining was shown to be an

effective method for improving sagebrush vigor and production on

a critical mule deer winter range.

Introduction ___________________

The importance of big sagebrush as winter forage for mule

deer has received considerable attention as researchers

have attempted to determine the species’ role in mule deer

nutrition. There is widespread agreement that it is frequently a dominant constituent of the winter diet of mule

deer (Kufeld and others 1973; Leach 1956; Pederson and

Welch 1982). Research shows it to be a highly digestible food

during the winter (Urness and others 1977; Wallmo and

others 1977; Welch and Pederson 1981), and its nutritional

value is enhanced when taken with other forages (Nagy and

others 1964; Smith 1959). Although its importance in the

diet increases in late winter (Carpenter and others 1979;

Wallmo and others 1977), it is not a starvation food, and has

been shown to contribute substantially to the diet of mule

deer in the fall and early winter when other forages are

available.

Historically, mule deer carrying capacity has increased on

winter ranges that have experienced an increase in shrub

density (Urness 1979). However, on many sagebrush ranges

in Utah, shrub density has peaked, and plant communities

have become dominated by old, decadent shrubs. Browse

production has decreased dramatically in these areas, especially during recent drought periods (Davis and others

2000).

In the 1960s, Plummer and others (1968) suggested chaining as a viable method for thinning stands that had become

extremely dense and decadent. This method was suggested

with or without seeding on depleted game ranges to retain

native grasses and forbs. Similarly, Cain (1971) reported on

the effectiveness of the Ely chain in restoring depleted

rangelands dominated by pinyon-juniper and big sagebrush.

Mechanical treatment, as opposed to spraying or burning,

allows for controlled or partial treatments where an adjustment in relative shrub density is desired (Urness 1979).

Such treatments retain shrubs in the treated area and avoid

the problems associated with establishing shrubs from seed

on semiarid sites.

This study compared one-way and two-way chaining as

tools for restoring decadent stands of big sagebrush on

important big game winter ranges. Treatment and control

sites were identified during the planning phase of a 134-ha

chaining project. Two sites were used to evaluate each

management alternative (no chaining, one-way chaining,

and two-way chaining). The unmodified anchor chain that

was used for the project measured 76 m in length and

weighed 60 kg/m.

Location _______________________

The study site was located on an important mule deer

winter range at the base of the Book Cliff Range, just east of

Nash Wash in southeastern Utah. The elevation of the study

area ranged from 1,945 to 1,980 m. The deer winter range

was characterized by a narrow band of pinyon-juniper woodland, interspersed with sagebrush parks. The parks were

dominated by Wyoming big sagebrush (Artemisia tridentata

wyomingensis), but mountain big sagebrush (Artemisia

tridentata vaseyana) was also present, a common situation

on sagebrush winter ranges in the region (McArthur, personal communication).
USDA Forest Service Proceedings RMRS-P-38. 2005 71

Big Sagebrush Response to One-Way and Two-Way Chaining in Southeastern Uta Fairchild, Davis, and Brotherson

Methods _______________________

Plant Community Assessment

A 30-m baseline was located at each of the six study sites

following treatment. Three 30-m transects were run perpendicular to the baseline and centered on the 15-m mark. The

transects were randomly located at 3-m intervals along the

line. A 1-m2 quadrat was positioned at 3-m intervals on

alternating sides of the transect to yield 10 sampling quadrats per transect and 30 quadrats per site.

Cover was estimated for all plants, individual species,

bare ground, litter, rock, and cryptogamic crust within each

quadrat using a modified Daubenmire (1959) method described by Davis and others (2000). Shrub density was

estimated along a 0.005-ha strip plot centered on the 30-m

survey tape. Each shrub rooted within the strip was counted

and placed in one of the following age classes: seedling,

young, mature, decadent, or dead (USDI-BLM 1996). Mature shrubs were classified as decadent when 25 percent or

more of the branches were dead. A one-way analysis of

variance test was used to determine if mean plant cover and

shrub density values differed among treatment and control

sites. Sagebrush production was estimated at each site

using a modified method for predicting annual production

based on shrub crown measurements (Dean and others

1981). The modification involved estimating crown denseness as percent cover in two dimensions, rather than percent

volume in three-dimensional space. Regression equations

were developed for each site to predict annual biomass from

crown measurements. Shrubs selected for analysis were

aged according to the method described by Ferguson (1964).

Stem diameter and annual ring count data for each shrub

were subjected to regression analysis to allow prediction of

ages from stem diameter measurements.

Shrub Vigor Assessment

At 2-m intervals along each baseline, the nearest mature

sagebrush plant was selected for the shrub vigor analyses.

Subspecies determinations were made for each shrub according to the method described by Stevens and McArthur

(1974). Baseline shrubs were sampled in fall to obtain

estimates of annual leader growth, number of seed stalks

per shrub, and seed stalk frequency, weight, and length. A

one-way analysis of variance test was used to determine if

mean shrub vigor measurements differed among treatment

and control sites.

One of each of the replicated treatment and control sites

was selected to evaluate browse utilization by mule deer.

The first 10 shrubs selected for the shrub vigor assessment

along the baseline of each of the three sites were included in

the utilization study following the method described by

Smith and Urness (1962). The “use-index” (leader length

times percent use) described by Smith and Urness (1962)

was used to evaluate treatment effects on browse use by

mule deer.

Results ________________________

Precipitation

A rain gauge was located within 1.6 km of the study area

and monitored monthly by the USDI-BLM. A summary of

moisture received from November to May and from June to

October is given for the years 1987 to 1990 in table 1. Winter

precipitation, which greatly influences annual vegetative

growth, decreased steadily from 203 cm (1987 to 1988) to 91

cm (1989 to 1990). The severity of the drought in the Nash

Wash area was verified by observations of pinyon pine

(Pinus edulis) trees at lower elevations that died in 1989 and

1990. The dead trees were scattered across the landscape,

and many exceeded 100 years in age (Davis, personal communication).

Plant Community Responses

Shrub Density and Production—Pretreatment shrub

density averaged 18,500 plants/ha on the six sites. Density

estimates in the final year (1990) showed one-way chained

sites supporting 10,500 plants/ha and two-way chained sites

supporting 7,500 plants/ha, for reductions of 43 percent and

60 percent, respectively. Sagebrush seedling establishment

was negligible on treated and control sites (<1 percent).

Annual production (kg/ha) of shrubs in the one-way chained

treatment was consistently greater than in the two-way

chained treatment or the control from 1988 to 1990 (table 2).

Over the 3 years, browse production on one-way chained

sites surpassed that on untreated sites and two-way chained

sites by 140 percent and 350 percent, respectively.

Table 1—A summary of moisture received (mm) from November to

May and June to October, 1987 to 1990, at a BLM precipitation gauge 1.6 km from the Nash Wash study site.

1987 to 1988 1988 to 1989 1989 to 1990

- - - - - - - - - - - - - - mm - - - - - - - - - - - - - - 

June to October 236 91 99

November to May 203 155 91

Total 439 246 190

Table 2—Browse production on chained one-way, two-way, and control

treatmentsa

.

Year

Treatment 1988 1989 1990 Average

- - - - - - - - - - - - - - - - kg ha-1 - - - - - - - - - - - - - - 

One-way 430 570 480 490a1

Two-way 130 130 160 140b

Control 370 380 300 350c

a

Values not sharing the same letter superscript are significantly different at the

0.05 level.
72 USDA Forest Service Proceedings RMRS-P-38. 2005

Fairchild, Davis, and Brotherson Big Sagebrush Response to One-Way and Two-Way Chaining in Southeastern Utah

Age Structure—Seedling establishment did not contribute significantly to shrub density of the treatment or

control sites at Nash Wash. In the 3 years following the

chaining, seedlings averaged about 1 percent of the stand

density on treated sites and were almost nonexistent on

untreated sites.

The untreated sites supported a shrub population that

had a higher proportion of older shrubs than was found on

either treatment. The regression equation generated to predict age (y) based on stem diameter (x) was y = 0.83x + 12.26,

with a coefficient of simple determination of 0.52. Even

though only 52 percent of the variation in age was accounted

for by measuring stem diameters, some trends were apparent. Shrub age on control sites ranged from 32 to 97 years

and averaged 54 years. Shrubs averaged 43 years of age

(range: 19 to 62 years) on the one-way chained sites and 39

years (range: 19 to 62 years) on the two-way chained sites.

Older shrubs with wide stem diameters lacked flexibility,

offered resistance to the force of the chain, and were more

effectively uprooted than were the younger plants.

Although age-class determinations (seedling, young, mature, and decadent) can vary among observers, their value in

characterizing shrub populations has been recognized by

many researchers and land management agencies. There

was no significant difference in the contribution made by

each age-class on one-way and two-way chained sites. However, a higher proportion of decadent shrubs was consistently found in the untreated sites (table 3). The widest

margin of difference was noted in 1990 following the dry

winter of 1989 to 1990. The proportion of older shrubs

classified as mature rose to a high in 1989 on all sites.

Plant Cover—Cover class estimates for sagebrush did

not reveal cover differences between treatments, but did

show a difference between treatments and controls. The 3-

year average for sagebrush cover on treated sites and control

sites was 6 percent and 14 percent, respectively. Total

herbaceous plant cover was highly variable and limited on

all sites. The 3-year average for herbaceous plant cover on

one-way chained, two-way chained, and control sites was 7

percent, 14 percent, and 5 percent, respectively. Cheatgrass

made up about 50 percent of the herbaceous plant cover on

all sites (table 4). The extremely low cover values for all sites

in 1990 reflected the severity of the drought.

Shrub Vigor Responses

Leader Growth—The most striking impact of the thinning treatments was an increase in the length of vegetative

leaders on remnant shrubs. Average leader length for sagebrush in treated areas was almost double that of control sites

in 1988 (table 5). No significant difference in leader growth

between the two treatments was observed the first year

after chaining, but sagebrush vegetative growth on oneway chained sites surpassed that of two-way chained sites in

1989 and 1990. Essentially no growth was observed on

control sites in 1990 due to the drought. In addition, leader

growth in 1990 on shrubs from treated sites was well below

the average observed on control sites during the previous

two growing seasons.

Seed Production—The reproductive growth response to

thinning was more gradual than the vegetative growth

response. During the first growing season, seed stalk production was similar on treated and untreated sites. No

significant differences were detected in the average number

of seed stalks per shrub or the average length of seed stalks

among treatments and controls (table 6). In 1989, the number of seed stalks per shrub increased slightly (although

not significantly), and seed stalk length on treated sites

Table 3—Proportion of decadent shrubs in the stand on chained one-way, two-way, and control treatmentsa.

Year

 Shrub 1988 1989 1990

 condition 1-way 2-way Control 1-way 2-way Control 1-way 2-way Control

- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - percent - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 

Decadent 86 90 99* 59 58 64 76 74 98*

Not decadent 14 10 1* 41 42 46 24 26 2*

a Proportions followed by asterisks are significantly different (<0.05 level) than those of the other treatments based on categorical analysis using the chi-square statistic.

Table 4—Total herbaceous cover and percent contribution from

cheatgrass (in parentheses) on chained one-way, two-way,

and control treatmentsa.

Year One-way Two-way Control

- - - - - - - - - - - - - percent - - - - - - - - - - - - - 

1988 11a (45) 18b (61) 10a (90)

1989 9a (67) 18b (50) 4a (25)

1990 2a (50) 5a (20) 2a (50)

a

Values not sharing the same letter superscript are significantly different at the

0.05 level.

Table 5—Average sagebrush leader length in chained one-way, twoway, and control treatmentsa

.

Year

Treatment 1988 1989 1990

- - - - - - - - - - - - - - - cm - - - - - - - - - - - - - - 

One-way 7.5a 9.4a 1.4a

Two-way 7.1a 7.3b 1.2b

Control 4.0b 3.9c 1.0c

a

Values not sharing the same letter superscripts in each column are significantly different at the 0.05 level.
USDA Forest Service Proceedings RMRS-P-38. 2005 73

Big Sagebrush Response to One-Way and Two-Way Chaining in Southeastern Uta Fairchild, Davis, and Brotherson

exceeded that on control sites by 170 percent. In addition to

almost a doubling in length, seed stalk weight per cm was

6 times greater on shrubs from treated versus control sites,

averaging 0.097 gm/cm and 0.016 gm/cm, respectively. In

1990, shrubs from treated sites produced far more seed

stalks than shrubs from control sites; however, stalk length

did not vary significantly (table 5).

Browse Use by Mule Deer—Treatment effects on browse

utilization by wintering mule deer are summarized in table 7.

No significant differences were detected in browse use between the two treatments, but there were differences between treatments and controls. Deer utilized more forage

from shrubs in treated areas than untreated areas even

though utilization percentages were comparable. Average

utilization for each treatment and control ranged from 67 to

77 percent during winter 1988 to 1989 and from 51 to 67

percent in 1989 to 1990. Significantly higher use-index

values (leader length times percent use) were associated

with shrubs from treated versus untreated areas during

both winters.

Discussion _____________________

Utah Division of Wildlife Resources (UDWR) and BLM

biologists have recognized the depleted condition of the deer

winter range in the Nash Wash area since the mid-1960s.

The close proximity of this critical winter range to the

adjacent salt desert shrub type at lower elevations, traditionally used by sheep during the winter, has led to excessive

use of sagebrush and preferred herbaceous species. Coles

and Pederson (1967) were concerned that the combined

winter use by deer and sheep on sagebrush in the Nash Wash

area was excessive, and that if allowed to continue, would

result in a widespread sagebrush die-off. In an effort to avoid

such a loss, the Grand Resource Area (USDI BLM) modified

the grazing plan for the Nash Wash area and limited winter

sheep grazing to the salt desert shrub community below the

critical deer winter range.

The increase in forage production per shrub on chained

sites revealed the high degree of intraspecific competition

that existed among sagebrush plants in the study area.

Shrub density on one-way chained sites was sufficient to outproduce untreated control sites the following year and in

each succeeding year of the study. Sites chained two ways

did not respond with sufficient browse production to compensate for the additional shrub removal. The second pass

reduced sagebrush production to 50 percent of that found on

control sites.

The reproductive growth response to sagebrush thinning

was not as striking as the vegetative response. A gradual

increase in reproductive growth response to sagebrush thinning (both treatments combined) was observed over the 3-

year study period. No significant differences were observed

in stalk length, weight, or number the first year following

treatment. A reproductive response was observed the second

year in increased seed stalk lengths and stalk weights. Seed

stalk production (number per plant) did not increase until

the third flowering period following treatment.

Several factors appeared to be working against sagebrush

productivity in the study area. In general, the sagebrush

plants were old and decadent. Shrub age averaged 54 years,

and the oldest shrubs were close to 100 years in age. Seasonal precipitation patterns, typical of the Colorado Plateau,

may have limited the potential for sagebrush production at

Nash Wash. Weather data collected at Thompson, UT (16

km from the study site), over the past 30 years, showed that

only 55 percent of the annual precipitation was received

during the winter (November to May) period. Stations at the

Salt Lake City Airport and Fillmore reflected the trend in

the Great Basin where approximately 70 percent of the

annual precipitation was received during the November to

May period. Since winter precipitation is known to influence

big sagebrush production (Daubenmire 1975; Elderkin and

others 1986), and only 55 percent of the annual precipitation

at Nash Wash is received during the winter, shrub health

may be vulnerable, especially during drought periods.

Drought conditions were exacerbated at Nash Wash, where

competition among sagebrush plants was evident. Although

not evaluated in this study, the prevalence of cheatgrass, a

Table 7—Browse utilization by mule deer on chained one-way, two-way, and control sitesa.

Year

1989 1990

Treatment Leader length Use Use index Leader length Use Use index

cm percent cm percent

One-way 5.1a 67a 3.2ab 4.7a 64a 3.2a

Two-way 5.4a 76a 4.1a 4.4a 67a 3.0a

Control 2.7b 77a 2.1b 2.2b 51b 1.1b

a

Values not sharing the same letter superscripts in each column are significantly different at the 0.05 level.

Table 6—Seed stalk length and number per shrub in chained one-way,

two-way, and control treatmentsa

.

Year

1988 1989 1990

Treatment Length Number Length Number Length Number

cm cm cm

One-way 16.9a 0.5a 17.0a 3.3a 5.1a 14.7a

Two-way 17.2a 0.4a 15.1a 1.3a 4.9a 10.2a

Control 10.1a 1.0a 9.4b 1.3a 2.4a 0.3b

a

Values not sharing the same letter superscripts in each column are significantly different at the 0.05 level.
74 USDA Forest Service Proceedings RMRS-P-38. 2005

Fairchild, Davis, and Brotherson Big Sagebrush Response to One-Way and Two-Way Chaining in Southeastern Utah

winter annual, could have had a significant impact on

sagebrush production and seedling establishment.

Chaining in one direction with a light, unmodified anchor

chain, was shown to be an effective treatment for thinning a

decadent stand of sagebrush without reducing the carrying

capacity for a wintering deer herd. The treatment should be

considered with or without artificial seeding, depending on

the condition of the herbaceous understory, as a practical

method to improve sagebrush health and wildlife habitat on

critically important sagebrush ranges. Different results

would be expected from chaining one-way with heavier

chains, chains modified with rails welded crosswise to the

links, or chains pulled in a more aggressive “J” configuration

as opposed to the “U” shape maintained during the Nash

Wash project.

References _____________________

Cain, Don. 1971. The Ely chain: a practical handbook of principles

and practices of chaining and vegetative manipulation. Ely, NV:

U.S. Department of the Interior, Bureau of Land Management.

32 p.

Carpenter, L. H.; Wallmo, O. C.; Gill, R. B. 1979. Forage diversity

and dietary selection by wintering mule deer. Journal of Range

Management. 32: 226–229.

Coles, F. H.; Pederson, J. C. 1967. Big game inventory 1965–66.

Bull. 67-1. Salt Lake City, UT: Utah State Department of Fish

and Game.

Daubenmire, R. F. 1959. A canopy-coverage method of vegetational

analysis. Northwest Science. 33: 43–64.

Daubenmire, R. F. 1975. Ecology of Artemisia tridentata in the State

of Washington. Northwest Science. 49: 24–35.

Davis, J. N. 1987. Personal communication. Provo, UT: Utah Division of Wildlife Resources.

Davis, J. N.; Green, A.; Farmer, M. 2000. Utah big game range trend

studies. Publ. 00-38. Salt Lake City, UT: Utah Division of Wildlife

Resources: 202–210.

Dean, S.; Burkhardt, J. S.; Meeuwig, R. O. 1981. Estimating twig

and foliage biomass of sagebrush, bitterbrush, and rabbitbrush in

the Great Basin. Journal of Range Management. 34: 224–227.

Elderkin, R. L.; Dietz, D. R.; Welch, B. L.; Ralphs, P. M.; Czarnecki,

J. K. 1986. Irrigation by mine discharge water and fertilization of

big sagebrush: effects on nutritional composition and mule deer

use. In: McArthur, E. D.; Welch, Bruce L., comps. Proceedings—

symposium of the biology of Artemisia and Chrysothamnus; 1984

July 9–13; Provo, UT. Gen. Tech. Rep. GTR-INT-200. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 140–145.

Ferguson, C. W. 1964. Annual rings in big sagebrush, Artemisia

tridentata. Papers of the Laboratory of Tree-Ring Research 1.

Tucson, AZ: University of Arizona Press. 95 p.

Kufeld, R. C.; Wallmo, O. C.; Feddema, C. 1973. Foods of the Rocky

Mountain mule deer. Res. Pap. RMRS-RM-111. Fort Collins, CO:

U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station. 31 p.

Leach, H. R. 1956. Food habits of the Great Basin deer herds of

California. California Fish and Game. 42: 243–308.

McArthur, E. D. 1987. [Personal communication]. Provo, UT: U.S.

Department of Agriculture, Forest Service, Rocky Mountain

Research Station, Shrub Sciences Laboratory.

Nagy, J. G.; Steinhoff, H. W.; Ward, G. M. 1964. Effects of essential

oils of sagebrush on deer rumen microbial function. Journal of

Wildlife Management. 28: 785–790.

Pederson, J. C.; Welch, B. L. 1982. Effects of monoterpenoid exposure on ability of rumen inocula to digest a set of forages. Journal

of Range Management. 35: 500–502.

Plummer, A. P.; Christensen, D. R.; Monsen, S. B. 1968. Restoring

big game ranges in Utah. Publ. 68-03. Salt Lake City, UT: Utah

Division of Fish and Game. 183 p.

Smith, A. D. 1959. Adequacy of some important browse species in

overwintering of mule deer. Journal of Range Management. 12:

8–13.

Smith, A. D.; Urness, P. J. 1962. Analysis of the twig length method

of determining utilization of browse. Publ. 62-9. Salt Lake City,

UT: Utah State Fish and Game.

Stevens, R.; McArthur, E. D. 1974. A simple field technique for

identification of some sagebrush taxa. Journal of Range Management. 27: 325–326.

U.S. Department of the Interior, Bureau of Land Management

(USDI BLM). 1996. Sampling vegetation attributes, Interagency

Tech. Ref. BLM/RS/ST-96/002+1730. Denver, CO: U.S. Department of the Interior.

Urness, P. J. 1979. Wildlife habitat manipulation in sagebrush

ecosystems. In: The sagebrush ecosystem: a symposium; April

1978; Logan, UT: Utah State University: 169–178.

Urness, P. J.; Smith, A. D.; Watkins, R. K. 1977. Comparison of in

vivo and in vitro dry matter digestibility of mule deer forages.

Journal of Range Management. 30: 119–121.

Wallmo, O. C.; Carpenter, L. H.; Regelin, W. L.; Gill, R. B.; Baker,

D. L. 1977. Evaluation of deer habitat on a nutritional basis.

Journal of Range Management. 30:122–127.

Welch, B. L.; Pederson, J. C. 1981. In vitro digestibility among

accessions of big sagebrush by wild mule deer and its relationship

to monoterpenoid content. Journal of Range Management. 34:

497–500.

Hoary arnica
USDA Forest Service Proceedings RMRS-P-38. 2005 75

Scott M. Lambert is Regional Seed Coordinator, Idaho State Office, Bureau

of Land Management, U.S. Department of the Interior, Boise, ID 83709; e-mail:

Scott_Lambert@blm.gov

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Seeding Considerations in

Restoring Big Sagebrush Habitat

Scott M. Lambert

Abstract—This paper describes methods of managing or seeding to

restore big sagebrush communities for wildlife habitat. The focus is

on three big sagebrush subspecies, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), basin big sagebrush (Artemisia

tridentata ssp. tridentata), and mountain big sagebrush (Artemisia

tridentata ssp. vaseyana). Natural colonization of the native plant

community may be the preferred management action on sites where

native seed sources are available to successfully reestablish the

desired wildlife habitat. On highly disturbed or otherwise damaged

sites and where competition from weeds is excessive, seeding will be

utilized to restore big sagebrush for wildlife habitat. Big sagebrush

seed is never seeded alone in site rehabilitation and restoration

projects. The best time to seed or interseed big sagebrush seed

mixes, including grasses and forbs, is in late fall or early winter. The

overall best method to reestablish big sagebrush is to use a rangeland drill at a shallow setting following site preparation, including

tillage and weed control. When big sagebrush is drill seeded with

other seed types, it is recommended that it be seeded through a

separate drill box to permit very shallow seeding and proper seed

placement for plant establishment. Seedings of native plants,

including big sagebrush, should be protected from grazing for at

least 3 to 5 years to allow time for the shrubs and forbs to become

established.

Introduction ____________________

Big sagebrush dominated plant communities occupy over

25 percent (67 million acres) of the landscape in the Great

Basin region of Idaho, Utah, Nevada, Oregon, and California.

A total of about 96 million acres in the Western States has

historically been big sagebrush habitat (Blaisdell 1953). In

some areas, over one-half of the big sagebrush areas have

been severely disturbed, altered, or even removed by wildfire, grazing, prolonged drought, and other natural events or

human activities. This paper will concentrate on seeding considerations for habitat restoration, including big sagebrush

(Artemisia tridentata) subspecies Wyoming big sagebrush

(Artemisia tridentata ssp. wyomingensis), basin big sagebrush (Artemisia tridentata ssp. tridentata), and mountain

big sagebrush (Artemisia tridentata ssp. vaseyana).

Beginning in the 1930s and 1940s, big sagebrush-dominated plant communities were reseeded to introduced forage

grasses and forbs such as crested wheatgrass (Agropyron

cristatum), Siberian wheatgrass (Agropyron fragile ssp.

sibericum), alfalfa (Medicago sativa), and pubescent wheatgrass (Elytrigia intermedia). Intermediate wheatgrass

(Elytrigia intermedia) was one of the preferred introduced

species seeded on mountain big sagebrush sites, which

usually have higher annual precipitation. These plant species were often used to improve forage production and reduce

soil erosion after wildfires and other site disturbances.

Since the 1990s, the emphasis has been to restore wildlife

habitat with a diversity of native plant species on public

lands in the Western States. Seeds of many native and

introduced species are commercially available, including

certified cultivars and source-identified native germplasms.

Basin big sagebrush and Wyoming big sagebrush seeds

are light brown to grayish brown or black and very similar

in size and shape (Parkinson 2004). Mountain big sagebrush

seeds are slightly larger and darker than basin big sagebrush or Wyoming big sagebrush seeds.

One proven method to obtain seed of the desired big

sagebrush subspecies is to procure Certified Source Identified (SI) seed of the big sagebrush subspecies appropriate, or

adapted, to the site. SI seed has been verified as to the

subspecies. The SI seed is collected from mature plants

growing on an identified natural site.

Identification of big sagebrush seed to subspecies may be

difficult to determine just by looking at the seed. In the past,

Bureau of Land Management and others that seeded big

sagebrush seed were not always provided with seed of the

subspecies that was specified in seeding contracts. Wyoming, basin, and mountain big sagebrush establish and

thrive on sites with different environmental conditions.

Using seed of the inappropriate subspecies of big sagebrush

may be a reason for stand failure on some sites (Lysne and

Pellant 2004).

The habitat management option of “do not seed” has been

the action taken on about 50 percent of the potential rehabilitation or restoration sites on public land in the Great

Basin States (Lambert and Hamby 2003). Natural colonization of native or introduced species is often allowed to occur

on sites where the seeds of desired plants exist in the soil

seed bank or on adjacent lands. Natural plant recovery will

only be successful on those sites with adequate soil and

moisture conditions and where competition from weeds is

not a problem.

Some big sagebrush site restoration projects, especially

those in Wyoming big sagebrush habitat, may initially

require seeding for soil stabilization and weed control. These

seedings may include adapted introduced plant species. At

a later time, other desirable native plants can be seeded (or
76 USDA Forest Service Proceedings RMRS-P-38. 2005

Lambert Seeding Considerations in Restoring Big Sagebrush Habitat

interseeded) to provide a greater diversity of native species

for wildlife habitat.

Descriptions of Big Sagebrush

Subspecies ____________________

Basin Big Sagebrush (Artemisia tridentata

ssp. tridentata Nutt.)

Basin big sagebrush is an evergreen shrub commonly 3 to

6 feet, but sometimes as much as 14 feet tall and 5 to 8 feet

wide. It is erect, spreading, and heavily branched, with an

uneven top. This subspecies of big sagebrush is native to the

Interior Western United States, including the Great Basin.

It is found on well-drained, moderately deep to deep loam or

silt loam soils, and at elevations up to 9,500 feet. The

moisture regime is semidry; 8 to 18 inches of mean annual

precipitation is required. Basin big sagebrush is the most

abundant shrub in the Western United States aridlands

(Passey and others 1982). It is sometimes interspersed with

Wyoming big sagebrush, and may also occur in riparian

sites. It is generally fire intolerant.

Basin big sagebrush averages 2,500,000 cleaned seeds per

pound (McArthur and others 1979); seed is harvested in the

late fall. The drilled seeding rate is 0.1 to 0.2 lb pure live seed

(PLS) per acre in a seed mixture. Big sagebrush seed is

planted very shallow (approximately 1⁄16 inch deep). Caution: Basin big sagebrush seed is very short-lived and may be

fragile when improperly handled. The seed must be kept in

cold storage (<38 °F and <25 percent relative humidity

[RH]). It is recommended the seed be used within 1to 2 years

of harvest.

Mountain Big Sagebrush (Artemisia

tridentata ssp. vaseyana [Rydb.] Beetle)

Mountain big sagebrush is an evergreen shrub that is

usually less than 3 feet tall with a spreading and even topped

crown. It is native to the Interior Western Unites States,

including the Great Basin region. It occurs on soils that are

deep and well drained with pH usually about 7.0. It is

naturally found in association with mountain shrub plant

communities at moderate to high elevations, up to 10,000

feet. Its moisture regime is semidry; at least 13 inches of

mean annual precipitation is required to maintain a population of this subspecies. After fire, mountain big sagebrush

often recovers from seed remaining in the soil. On many

restoration sites, mountain big sagebrush sites do not require reseeding unless there is an overwhelming population

of weedy plants. In such situations, weed control would be

required prior to seeding.

Mountain big sagebrush averages 2,250,000 cleaned seeds

per pound (McArthur and others 1979); the seed is harvested

earlier in the fall than the other two subspecies. The drilled

seeding rate for mountain big sagebrush is 0.1 to 0.2 lb PLS

per acre in a seed mixture. The seed should planted about 1⁄16

inch deep in the soil (Jacobson and Welch 1987). ‘Hobble

Creek’ is a native cultivar that originated in Wasatch County,

Utah, and was released by the USDA Forest Service, Shrub

Sciences Laboratory, Provo, Utah (Welch and others 1986).

Caution: mountain big sagebrush seed is very short-lived

and may be fragile when improperly handled. The seed must

be stored in cold storage (<38 ° F) with humidity maintained

at <25 percent R.H. It is recommended the seed be used

within 1 to 2 years of harvest.

Wyoming Big Sagebrush (Artemisia

tridentata ssp. wyomingensis Beetle &

Young)

Wyoming big sagebrush is an evergreen shrub usually up

to 3 feet tall and usually up to 3 feet wide at maturity. It is

basally branched and rounded in form with an uneven top.

It is native to the Interior Western United States, including

the Great Basin region. This subspecies grows on shallow,

gravelly, or sandy to silt clay loams at elevations from 2,000

to 7,000 feet. Its moisture regime is dry; 7 to 14 inches of

mean annual precipitation is required for Wyoming big

sagebrush. It is considered to be fire-intolerant and does not

resprout after wildfire.

Wyoming big sagebrush averages 2,500,000 cleaned seeds

per pound (McArthur and others 1979); the seed is harvested

in late fall. The drilled seeding rate for Wyoming big sagebrush is 0.1 to 0.2 lb PLS per acre in a seed mixture. The seed

is planted shallow in soil (1⁄16 inch in depth). Cultivar:

‘Gordon Creek’ Wyoming big sagebrush originated in Carbon County, Utah, and was released by the USDA Forest

Service, Shrub Sciences Laboratory (Welch and others 1992).

Currently there is no commercial seed production of Gordon

Creek. Caution: Wyoming big sagebrush seed is very shortlived and may be fragile when improperly handled. The seed

must be stored in cold storage (<38 °F) with humidity

maintained at <25 percent RH. It is recommended the seed

be used within 1 to 2 years of harvest.

Seeding Specifications ___________

Seeding Dates for Big Sagebrush

Late fall is the overall best time to seed big sagebrush, and

rangeland drill seeding is the recommended method. Early

winter is the preferred time for aerial seeding. Whichever

seeding method is used, it is best to seed prior to any

significant snowfall.

Seeding Requirements

A firm, packed, weed-free seedbed should be prepared on

restoration sites. The best soils for establishing big sagebrush from seed are silt loam to sandy loams, deep to shallow

and well drained. One of the best methods for successful

establishment of big sagebrush by seed is to broadcast the

seed on the soil surface and then lightly rake or harrow to

barely cover the small seed. Another method of big sagebrush establishment that has been successfully used is to

apply the seed with a rangeland seed drill (Truax, Tye, and

so forth). The seed drill is set to place the seed very shallow,

about 1⁄16 inch deep. The packer wheels of the seed drill

should lightly cover the seed with soil. The best chance for a

successful seeding is if the seed drill has a separate seed box

for the big sagebrush seed.
USDA Forest Service Proceedings RMRS-P-38. 2005 77

Seeding Considerations in Restoring Big Sagebrush Habitat Lambert

Aerial or broadcast seeding has been successful on some

sites in late fall or just prior to snowfall in early winter.

Aerial seeding may not be as good a seeding method as a

properly installed drilled seeding or broadcasting seeding

followed by harrowing to lightly cover seed with soil (Lysne

and Pellant 2004).

Seeding Rate for Big Sagebrush

Subspecies

The typical drill seed rate, pure live seed (PLS), for all

three big sagebrush subspecies is 1⁄10 lb per acre in a seed

mix. Basin big sagebrush has, on average, 2,500,000 cleaned

seeds per pound (1⁄10 lb = 57 seeds per square foot). Mountain big sagebrush has 2,250,000 cleaned seeds per pound

(1⁄10 lb = 51 seeds per square foot). Wyoming big sagebrush

has 2,500,000 cleaned seeds per pound, according to McArthur

(1979) (1⁄10 lb = 57 seed per square foot).

Seed Mixes

Challenges to Seeding Small Seeds—Big sagebrush

seed is much smaller than most other species that are

typically drill seeded. Big sagebrush seedlings are slow

growing and often less competitive than most other species

that occur in the same habitat. The best options to establish

big sagebrush are to use a rangeland drill with a separate

seed box for big sagebrush or broadcast seed and very lightly

cover with soil. An additional option is to grow the seed in a

greenhouse to produce container seedlings and then transplant to the field location, or grow the seed in a field to

produce bare-root stock and transplant to the desired site.

Establishing Big Sagebrush With Understory Species—Big sagebrush subspecies are not seeded alone. Sagebrush seed is included in seed mixtures with other species,

usually grasses and forbs. If big sagebrush is drill seeded, it

is recommended that it be seeded through a separate drill

box to permit very shallow seeding. The drilled seed mix

should be installed at the PLS rate of 60 to 80 seeds per

square foot. To determine a seed mix for your specific

ecological situation, utilize information available through

the PLANTS database (http://plants.usda.gov/). The PLANTS

Web site includes links to VegSpec, a database to help with

seeding prescriptions, and the Ecological Site Information

System (ESIS).

The following seed mixes are examples for each of the

three big sagebrush subspecies and include understory

grasses and native forbs for habitat restoration projects

(tables 1, 2, and 3). Examples of two seed mixes for soil

stabilization and erosion control are also provided (table 4).

Checklist of Potential Causes for Seeding

Problems or Failure

The seed doesn’t grow when initially planted.

1. The seed selected was not appropriate for the given

environment. This problem may be due to inaccurate

seeding specifications or inadequate seed availability. For

many native species, a local seed source may be preferable to

those sources that originate further away from the seeding

Table 1—Example of generic basin big sagebrush steppe seed mix,

drilled rate (PLS).

Native species Pounds per acre

Basin big sagebrush 0.1

Rubber rabbitbrush 0.1

Bluebunch wheatgrass 2.0

Indian ricegrass, on sandy soil 2.0

Thurber’s needlegrass 1.0

Sandberg’s bluegrass 1.0

Bottlebrush squirreltail 2.0

Six-weeks fescue 1.0

Pale agoseris 0.2

Threadstalk milkvetch 0.2

Western yarrow 0.1

Arrowleaf balsamroot 0.2

Tapertip hawksbeard 0.2

Nineleaf biscuitroot 0.2

Fleabane spp. 0.1

Total 8.4

Table 2—Example of generic mountain big sagebrush community

seed mix, drilled rate (PLS).

Native species Pounds per acre

Mountain big sagebrush 0.1

Woods rose/mountain snowberry 0.5

Sulfur-flowered buckwheat 0.2

Idaho fescue 2.0

Bluebunch wheatgrass 2.0

Bottlebrush squirreltail 1.0

Slender wheatgrass 1.0

Pale agoseris 0.2

Woollypod milkvetch 0.2

Arrowleaf balsamroot 0.2

Tapertip hawksbeard 0.2

Sagebrush mariposa-lily 0.2

Total 7.8

Table 3—Example of generic Wyoming big sagebrush steppe seed

mix, drilled rate (PLS).

Native species Pounds per acre

Wyoming big sagebrush 0.1

Rubber rabbitbrush 0.1

Indian ricegrass (on sandy site) 2.0

Thurber’s needlegrass 1.0

Sandberg’s bluegrass 1.0

Bottlebrush squirreltail 2.0

Six-weeks fescue 1.0

Pale agoseris 0.2

Threadstalk milkvetch 0.2

Western yarrow 0.1

Tapertip hawksbeard 0.2

Nineleaf biscuitroot 0.2

Fleabane spp. 0.1

Total 8.25
78 USDA Forest Service Proceedings RMRS-P-38. 2005

Lambert Seeding Considerations in Restoring Big Sagebrush Habitat

Table 4—Samples of soil stabilization seed mixes for sites with less

than 12 inches mean annual precipitation.

Species and variety Pounds per acre

PLS, drilled rate

A. Species for a sandy soil site

Siberian wheatgrass, ‘Vavilov’ or ‘P27’ 2

Indian ricegrass, ‘Nezpar’ 1

Thickspike wheatgrass, ‘Critana’, ‘Bannock’,

 or ‘Schwendimar’ 1

Sand dropseed 1

Alfalfa, ‘Ladak’ 2

Total 7

B. Species for a silt-clay loam soil site

Crested wheatgrass, ‘Nordan’ or ‘Hycrest’ 2

Western wheatgrass, ‘Arriba’ 1

Snake River wheatgrass, ‘Secar’ 2

Pubescent wheatgrass, ‘Luna’ 1

Sainfoin, ‘Eski’ 1

Total 7

location in terms of distance, elevation, or other site factors.

The seed from nonlocal collections or seed production fields

may be less well adapted to the restoration site. Seed not

adapted to the site may have no or a low percentage of seeds

that germinate, establish successfully, and persist over time.

2. The seed may remain dormant in the soil for varying

periods of time. Most seed planted in late fall or early

winter will germinate the following spring. Some seed, such

as the hard-coated seeds of Indian ricegrass, may take

several years under normal conditions before initiation of

germination.

3. There may have beeen poor seed storage conditions

prior to seeding. In general, seed stored for more than 2

weeks must be held in a climate-controlled warehouse with

temperatures not to exceed 80 °F and 30 percent RH.

4. The seed delivered was not viable. This results when

viability has declined following testing or when seed is

damaged during transport or handling. To overcome this

potential problem, arrange for a certified seed sampler to

confirm or deny the original seed tag information by taking

seed samples of the seed lots prior to mixing. Then have the

seed samples sent to a certified seed lab for purity and

germination or TZ tests and noxious weed seed analysis. Do

not accept seed lots with low germination rates or unacceptable PLS. Acceptable germination rates or PLS percent

should be determined by the agency or by the Seed Certification standards for the seed type or species. An example of

the Certified seed standard minimums set for bluebunch

wheatgrass are 85 percent pure seed, 80 percent germination, and 68 percent PLS (AOSCA 2001).

5. The seed was planted at too great a depth. Be sure that

all seed drills and other equipment are set to install the seed

in the soil correctly before you start the seeding operation.

6. The seed was damaged during application. Some damage to seed may occur with rough handling, transportation

of seed to the field site, or during hydroseeding applications.

7. If seed is applied in mulch, the medium may not be

capable of sustaining seed germination on dry sites. Some

temporary mulches, especially those made of recycled newsprint,

can contain inks and metals that are toxic to the newly

germinated seedlings.

8. The time of seeding was past the normal germination

season for that species or population. This problem can vary

among species. Some species will actually germinate under

snow cover during the winter, while others will break seed

dormancy and germinate in the following spring or summer.

The seed has germinated and later dies off.

1. Soil nutrients and moisture content are insufficient to

sustain seedling growth.

2. The soil type, such as silt loam, clay loam, or sandy

loam, is not capable of sustaining seed germination and

seedling development of plants that were seeded.

3. Diseases, such as damping off (a fungal disease), may

kill seedlings soon after germination.

Environmental conditions may cause additional stress

and prevent seedling establishment.

1. Conditions that may increase seedling mortality include drought, excessive heat or cold spells, wind, flood,

early frost, or late frost.

2. Competing vegetation, especially weeds such as

cheatgrass (Bromus tectorum), will cause seedling mortality in less competitive plant species (West and Hassan

1985). Control of weeds may be essential to seedling establishment.

3. After seeding it was found that the seeding mix contained some weed seeds that germinated and took over the

site. To overcome this problem, arrange for all seed lots to be

tested by a certified seed lab for noxious and other weed

seeds prior to mixing. Do not accept seed lots with unacceptable weed seeds on the seed lab analysis.

4. Animals and insects may have eaten the seed/seedlings.

You may need to provide some manner of protection to

emerging seedlings. Seedings of native plants, including big

sagebrush, should be protected from grazing for at least 3 to

5 years to allow time for the shrubs and forbs to become

established.

5. The soil may lack the microorganisms and fungal

mycorrhizae necessary for seedling establishment. Inoculation of seed or soil with microorganisms may be necessary

for plant health.

Monitoring Seedings_____________

Use monitoring protocols from the BLM/NRCS Monitoring Handbook (Elzinga and others 1998) or other monitoring

protocols as determined by the administering agency to

evaluate the success of seedings. On arid sites, a seeding is

often considered to be successful if at least 0.5 plants per

square foot are established.

Planting Big Sagebrush Using Seedlings

On some sites in the Western United States, good establishment of big sagebrush has resulted when seedlings are

planted in early spring on locations with the “best” soils and

aspect (Everett 1980). Arrangements should be made with a

nursery or grower to purchase or produce big sagebrush
USDA Forest Service Proceedings RMRS-P-38. 2005 79

Seeding Considerations in Restoring Big Sagebrush Habitat Lambert

seedlings when restoration plantings are the preferred plant

establishment option. Field-grown bare-root stock or greenhouse-grown containerized plants may be used. The time of

sowing seed in a production field to lifting of conservation

grade bare-root stock ready for planting could be as much as

2 years. Greenhouse grown seedlings may be ready to plant

within a 6 to 8 month time period.

To establish the big sagebrush seedlings it is recommended that they be randomly placed in clumps or blocks on

the best sites for restoring big sagebrush. Seedlings planted

in natural blocks or clumps become fertile islands of big

sagebrush as they mature. These may establish additional

plants throughout the adjacent areas.

Sources for Big Sagebrush Seed

Wildland Collected Seed (Source-Identified)—Big

sagebrush seed is collected from natural big sagebrush

stands. Most seeds of big sagebrush subspecies are sold as

Source-Identified seed from wildland collection sites. Certified Source Identified seed of native plants is recommended

for all rehabilitation and restoration seedings. The Source

Identified Certification tag, usually yellow, verifies the species, County, State, elevation, seed lot number, and sometimes other geographic location information for the collected

seed.

Commercial Seed Field Production—Certified seed,

cultivars or source-identified, of native or introduced plants

is recommended for all rehabilitation and restoration

seedings. The Certified tag on a seed bag (blue tag) verifies

the cultivar, species, location where the seed was grown, no

noxious weed seed for the States specified, and the minimum

percent pure seed and percent germination.

Hobble Creek is the only big sagebrush cultivar currently

available in limited quantities from seed vendors. Native

and introduced cultivars of other plant species are available

in limited to good quantities from seed vendors (table 5).

Summary ______________________

Natural colonization of the native plant community may

be the preferred management action on sites where native

seed sources are available to successfully reestablish the

desired wildlife habitat. On highly disturbed or otherwise

damaged sites or where excessive competition of weeds

occurs, seeding will be used to restore the big sagebrush for

wildlife habitat.

Big sagebrush seed is never seeded alone in site rehabilitation and restoration projects. The best time to seed or

interseed big sagebrush seed mixes, including grasses and

forbs, is in late fall or early winter. The best method to seed

big sagebrush is by using a rangeland seed drill. The seed is

drilled shallowly with the seed drill set for the appropriate

planting depth and PLS seeding rate. Wyoming big sagebrush sites are usually the most arid and shallowest soil

sites where big sagebrush naturally occurs. Mountain big

sagebrush sites are those sagebrush sites, in general, with

the coolest average temperature and annual rainfall

greater than 12 inches. Basin big sagebrush sites are often

transitional between the foothills and mountains where the

Table 5—Partial list of commercial big sagebrush seed vendors (these

seed vendors may have seed of other species also).

Vendor Vendor

Barton Seed Co.

222 E. Union Street

Manti, UT 84642

Phone: (435) 835-9200

Comstock Seed Co.

917 Hwy 88

Gardnerville, NV 89410

Phone: (775) 746-3681

Fremont Trading Co.

450 S. 50 E.

Ephraim, UT 84627

Phone: (435) 283-4701

Geertson Seed Farms

1665 Burroughs Road

Adrian, OR 97901

Phone: (541) 339-3768

Granite Seed Co.

1697 W. 2100 N.

Lehi, UT 84043

Phone: (801) 768-4422

Harvest Moon Seed Co.

PO Box 532

Richfield, UT 84701

Phone: (435) 979-8549

Intermountain Seed Co.

Box 62

370 W. 300 N.

Ephraim, UT 84627

Phone: (435) 283-4703

Landmark Seed Co.

N. 120 Wall St., Suite 400

Spokane, WA 99201

Phone: (509) 835-4967

Maughan Seed Co.

PO Box 72

700 W. 2100 S.

Manti, UT 84642

Phone: (435) 835-0401

NP Seed Co.

206 E. 300 S.

Manti, UT 84642

Phone: (435) 835-8301

Rainier Seed Co.

PO Box 70

Port Orchard, WA 98367

Phone: (800) 828-8873

Native-Seed Co.

7361 Pineridge Drive

Park City, UT 84098

Phone: (435) 640-0557

Plummer Seed Co.

PO Box 70

Ephraim, UT 84627

Phone: (435) 283-4844

Stevenson Intermountain Seed Co.

PO Box 2

Ephraim, UT 84627

Phone: (435) 283-6639

Southern Utah Seed Co.

PO Box 124

192 W. 100 S.

Junction, UT 84740

Phone: (435) 577-2142

Wagstaff Seed

1900 E. Oakhill Lane

Wallsburg, UT 84082

Phone: (435) 654-3439

mountain big sagebrush community is found and the arid

lowlands are dominated by the Wyoming big sagebrush

community. Basin big sagebrush often naturally occurs on

microsites with deeper soils in Wyoming big sagebrush

habitat and may be found along Interior Western riparian

zones. After seeding, the restored sites should be rested, or

protected, from grazing for at least 3 to 5 years to allow time

for the shrubs, forbs, and grasses to become fully established

for wildlife habitat.

References _____________________

Association of Official Seed Certifying Agencies (AOSCA). 2001.

Genetic and crop standards of the AOSCA. Meridian, ID: Association of Official Seed Certifying Agencies. 118 p.

Blaisdell, J. P. 1953. Ecological effects of planned burning of sagebrush-grass range on the Upper Snake River Plain. Tech. Bull.

1075. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 38 p.

Elzinga, C. L.; Salzer, D. W.; Willoughby, J. W. 1998. Measuring and

monitoring plant populations. Tech. Ref. 1730-1. Denver, CO:

U.S. Department of the Interior, Bureau of Land Management.

492 p.
80 USDA Forest Service Proceedings RMRS-P-38. 2005

Lambert Seeding Considerations in Restoring Big Sagebrush Habitat

Everett, R. L. 1980. Use of containerized shrubs for revegetating

arid roadcuts. In: Reclamation Review. Vol. 3. London: Pergamon

Press Ltd.: 33–40.

Jacobson, T. L.; Welch, B. L. 1987. Planting depth of ‘Hobble

Creek’ mountain big sagebrush seed. Great Basin Naturalist.

47: 497–498.

Lambert, S. M.; Hamby, J. L. 2004. Seed use by BLM on public land

in the Great Basin States. In: Abstracts presented at the 57th

annual meeting of the Society for Range Management: Rangelands in transition; 2004 January 24–30; Salt Lake City, UT: 98.

Lysne, C. R.; Pellant, M. 2004. Establishment of aerially seeded big

sagebrush following southern Idaho wildfires. Tech. Bull. 2004-

01. Boise, ID: U.S. Department of the Interior, Bureau of Land

Management. 14 p.

McArthur, E. D.; Blauer, A. C.; Plummer, A. P.; Stevens, R. 1979.

Characteristics and hybridization of important Intermountain

shrubs. III. Sunflower family. Res. Pap. INT-220. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Forest and Range Experiment Station. 82 p.

Parkinson, H. 2004. Germination of Artemisia tridentata ssp.

tridentata, ssp. vaseyana, and ssp. wyomingensis after one and

two years of storage. Unpublished paper on file at: U.S. Department of the Interior, Bureau of Land Management, Idaho State

Office, Boise, ID. 14 p.

Passey, H. B.; Hugie, B. K.; Williams, E. W.; Ball, D. E. 1982.

Relationships between soil, plant community, and climate on

rangelands of the Intermountain West. Tech. Bull. 1669. Washington, DC: U.S. Department of Agriculture, Soil Conservation

Service. 123 p.

USDA, Natural Resources Conservation Service. 2004. The PLANTS

Database, Version 3.5 [Online]. Baton Rouge, LA: U.S. Department of Agriculture, National Resources Conservation Service,

National Plant Data Center. Available: http://plants.usda.gov

Welch, B. L.; McArthur, E. D.; Nelson, D. L.; Pederson, J. C.; Davis,

J. N. 1986. ‘Hobble Creek’—a superior selection of low-elevation

mountain big sagebrush. Res. Pap. INT-370. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station. 10 p.

Welch, B. L.; Nelson, E. D.; Young, S. A.; Sands, A. R.; Wagstaff, F.

J.; Nelson, D. L. 1992. ‘Gordon Creek’—a superior, tested

germplasm of Wyoming big sagebrush. Res. Pap. INT-461. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 7 p.

West, N. E.; Hassan, M. A. 1985. Recovery of sagebrush-grass

vegetation following wildfire. Journal of Range Management. 38:

131–134.

Big sagebrush
USDA Forest Service Proceedings RMRS-P-38. 2005 81

Mike Pellant is Coordinator, Great Basin Restoration Initiative, Nevada

State Office, Bureau of Land Management, Boise ID 83709, U.S.A.; e-mail:

Mike_Pellant@blm.gov. Cindy R. Lysne is Environmental Specialist, Power

Engineers, Inc., Boise, ID 83707; e-mail: clysne@powereng.com

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Strategies to Enhance Plant

Structure and Diversity in

Crested Wheatgrass Seedings

Mike Pellant

Cindy R. Lysne

Abstract—Crested wheatgrass (Agropyron cristatum sensu amplo

[L.] Gaertn.) is an introduced, caespitose grass that has been seeded

on millions of acres of Western rangelands. In some areas, crested

wheatgrass seedings overlap with critical sage-grouse (Centrocercus

urophasianus; C. minimus) habitat, raising the question of how

plant diversity might be restored in these closed plant communities.

A three-step process is described to reduce crested wheatgrass

competition, introduce desired species, and manage to maintain

desired species for use long term. Crested wheatgrass is a strong

competitor with other species and a prolific seed producer, which

hinders treatments to reduce its influence and improve conditions

for establishment of desirable seeded species. Herbicides, burning,

mechanical treatments, livestock grazing, droughts, and combinations of these are effective to varying degrees in reducing crested

wheatgrass competition. Once crested wheatgrass competition is

reduced, either seed or seedlings can be used to increase diversity in

these seedings. Post-establishment management and monitoring

are essential components of the strategy to maintain plant diversity

into the future.

Introduction ____________________

Use of introduced species invokes a range of emotions from

unequivocal support to outright opposition and has even been

linked to fascism and racism (Simberloff 2003). In the Western United States, the planting of introduced perennial

wheatgrasses for rangeland rehabilitation has and continues

to be practiced after disturbances such as wildfires, on cropland taken out of production, and to increase forage production for livestock. Given the focus of this symposium on the

restoration of habitat for sage-grouse, using crested wheatgrass (Agropyron cristatum sensu amplo [L.] Gaertn.) to

meet certain land-use objectives must be evaluated relative

to the millions of acres already planted to these grasses and

their continued use in rangeland rehabilitation projects.

This paper will focus on a review of the characteristics,

use, and control techniques for crested wheatgrass and

other closely related introduced, caespitose bunchgrasses

(Siberian wheatgrass [Agropyron fragile], and Russian

wildrye [Psathyrostachys juncea]), prior to reintroducing

plant diversity. Crested wheatgrass and its close relatives

were introduced from Eurasia and selected for land rehabilitation in the Central and Western United States. The areas

where crested wheatgrass has been extensively used in the

Western United States overlap closely with historic sagegrouse distribution in the low elevation rangelands where

annual precipitation ranges from 8 to 12 inches annually

(USDA NRCS 2004). The first part of this paper will include

a review of the historical use, competitive characteristics,

and concerns regarding the use of crested wheatgrass, especially with regard to sage-grouse habitat. The remainder of

this paper will emphasize potential treatments to reduce

crested wheatgrass competition, where acceptable functional or structural vegetation components required by sagegrouse are not present, prior to increasing the diversity of

desirable herbs and shrubs.

Crested Wheatgrass: Introduction,

Uses, and Issues in Western

Ecosystems ____________________

The first collections of crested wheatgrass were made in

1897 to 1989 and again in 1906 from the dry steppes of

Eastern Russia (Dillman 1946; Rogler and Lorenz 1983).

These collections were classified as crested wheatgrass and

desert wheatgrass (Agropyron desertorum) and were distributed to 15 experiment stations throughout the West.

Minimal use of these introduced grasses occurred until the

1930s when a combination of cheap labor (for example,

Civilian Conservation Corps), the “dust bowl” in the Midwest,

and the need to reestablish perennial vegetation on abandoned farmlands prompted their increased use. Crested wheatgrass, primarily the caespitose bunchgrasses, were used extensively to revegetate abandoned croplands that were subject

to wind erosion in the Northern Great Plains (Holechek

1981; Young and Evans 1986). The first planting of crested

wheatgrass in the Intermountain area occurred in eastern

Idaho in 1932 (Hull and Klomp 1966). As the need for plants

to reclaim abandoned cropland increased, the production of

seed of crested wheatgrass also increased (Sharp 1986).

With greater demands for red meat production from Western rangelands during World War II, Congress allocated

funds to convert unproductive sagebrush rangelands to more

productive introduced grasslands (Young and McKenzie

1982). Researchers for the Forest Service developed a series of

bulletins on rangeland seeding, emphasizing the use of
82 USDA Forest Service Proceedings RMRS-P-38. 2005

Pellant and Lysne Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings

crested wheatgrass and other introduced wheatgrasses in

Idaho (Hull and Pearse 1943), Nevada (Robertson and Pearse

1943), and Utah (Plummer and others 1943).

Following World War II, the pace of rehabilitation accelerated again during the 1950s and early 1960s when millions of acres of Central and Western rangelands were

seeded to crested wheatgrass. The objectives of these seedings

included increasing forage for livestock, weed control, watershed stabilization, and reducing wildfire hazards. The

use of crested wheatgrass to biologically suppress halogeton

(Halogeton glomeratus), a nonnative poisonous forb, was

funded by Congress in 1952 and ultimately paid for a major

portion of crested wheatgrass seedings in Nevada and other

Great Basin States (Young and Evans 1986).

The rehabilitation of degraded rangelands with crested

wheatgrass was also accelerated by the development of

equipment to control competitive plants and distribute seed

effectively across a wide range of edaphic conditions. In

particular, the development of the rangeland drill in the

early 1950s hastened the ability of land managers to seed

large acreages to crested wheatgrass, while the brushland

plow provided managers with a tool for sagebrush removal

prior to seeding (Young and McKenzie 1982). By the early

1970s, referred to by Young and Evans (1986) as the “golden

age” of seeding crested wheatgrass, an estimated 12.4 million acres were seeded to this species (Dewey and Asay

1975). A recent report (USDI 2001) on the condition of public

lands managed by the Bureau of Land Management in the

Western United States indicates that approximately 5

million acres of rangelands have been seeded (USDI 2001),

the majority of which we estimate included crested wheatgrass in the seed mixture. However, since this information

is based in large part on inventory information collected in

the mid-1970s, the acreage of public lands seeded in part

with crested wheatgrass is expected to exceed this figure.

The use of crested wheatgrass has come under increasing

scrutiny since the 1970s. Legislation, such as the Surface

Mining Control and Reclamation Act (PL 95-87, 1977),

required the use of native species for mine reclamation. The

National Environmental Policy Act of 1969 (Pub. L. 91-190,

42 U.S.C. 4321-4347) required the preparation of an impact

analysis on activities funded by the government; this included the use of introduced species in seedings. Federal

agency guidance on this subject has also changed. Prior to

1984, the Bureau of Land Management’s guidance on postwildfire seeding encouraged the use of introduced grass

species given their cost, ease of establishment, and erosion

prevention capability (USDI BLM 1981). More recently,

Presidential Executive Order 13112 on Invasive Species

(Clinton 1999) directs Federal agencies to use native species

when feasible to restore ecosystems where invasive species

are a problem. Finally, the BLM’s Great Basin Restoration

Initiative (GBRI) gives preference to the use of native species in seeding projects, “pending seed availability, cost and

chance for success.” (USDI BLM 2000).

Competitive Characteristics of

Crested Wheatgrass _____________

An understanding of the competitive characteristics of

crested wheatgrass is essential in order to develop strategies

to increase plant diversity in seedings dominated by this

species. The same features that make crested wheatgrass

appealing to land managers (for example, provide soil stability and compete with and control invasive species) can also

result in community dominance of this species, displacement of native species, and reduced plant diversity (Broersma

and others 2000; D’Antonio and Vitousek 1992; Marlette

and Anderson 1986; Roundy and others 1997). Although

some studies reported that crested wheatgrass is not “mobile” and does not deter the reestablishment of native species (Broersma and others 2000; Krzic and others 2000),

several other studies have shown that established stands

have spread beyond the original seeded area (Hull and

Klomp 1966; Marlette and Anderson 1986). Other studies

have shown that crested wheatgrass seedings resulted in

near monospecific stands (Hull and Klomp 1966; Looman

and Heinrichs 1973; Schuman and others 1982).

There are several characteristics of crested wheatgrass

that contribute to its competitiveness with both invasive

species and native vegetation. At the seedling stage, crested

wheatgrass has an advantage over some native plants, due

in part to its ability to efficiently capture nutrients and

water (Bakker and Wilson 2001; Schuman and others 1982).

Established crested wheatgrass plants were more efficient

at securing phosphorus than native bluebunch wheatgrass

(Pseudoroegneria spicatum) when both species were grown

in association with big sagebrush (Artemisia tridentata)

(Caldwell and others 1985). Big sagebrush was also negatively affected by the ability of crested wheatgrass to rapidly

extract soil water during the same period that sagebrush

requires this resource (Cook and Lewis 1963; Eissenstat and

Caldwell 1988; Sturges 1977). Other studies have shown the

competitive advantage of crested wheatgrass during the

initial stages of plant establishment for Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) and antelope bitterbrush (Purshia tridentata) (Blaisdell 1949; Fortier

2000;Hall and others 1999; Schuman and others 1998).

Native grasses may also reduce shrub seedling establishment; however, the effect is less than that exhibited by crested

wheatgrass (Eissenstat and Caldwell 1988; Hubbard 1957).

For example, Frischknecht and Bleak (1957) reported that

seeded stands of bluebunch wheatgrass were more likely to

permit sagebrush seedling recruitment than seeded stands

of crested wheatgrass.

Another attribute that favors crested wheatgrass establishment is its prolific seed production. Marlette and Anderson (1986) germinated seed from a crested wheatgrass

monoculture planting and reported around 500 crested

wheatgrass seedlings per m2. Pyke (1990) compared the

demography of crested wheatgrass and bluebunch wheatgrass and found crested wheatgrass to have a decided

advantage over the native grass in seed production, seed

bank carryover, seed dispersal, and seedling survival.

Seed production for crested wheatgrass ranged from 1,772

seeds per m2 in a wet year to 1,037 seeds per m2 in a dry

year. Bluebunch wheatgrass seed production during this

same period ranged from 26 seeds per m2 in the wet year

to no seed production in the dry year. In a recent study,

Romo (2005) found an average return of 2 percent of crested

wheatgrass seeds sown while Heidinga and Wilson (2002)

reported a 4 percent return. For example, seed production

rates of 1,000 per m2 with a 3 percent rate of return would
USDA Forest Service Proceedings RMRS-P-38. 2005 83

Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings Pellant and Lysne

result in an initial count of 30 crested wheatgrass seedlings

per m2.

Carryover of germinable seed for more than 1 year is

another competitive characteristic of crested wheatgrass.

Under ideal seed storage conditions in a lab, crested wheatgrass seeds remained germinable for over 20 years (Ackigoz

and Knowles 1983). Seed life is much shorter in the natural

environment; however, crested wheatgrass has been observed by the authors to germinate in the second and, in a

few cases, the third growing season after the seed was

planted.

Sage-Grouse and Crested

Wheatgrass ____________________

Crested wheatgrass has been planted or has the potential

for establishment (Rogler and Lorenz 1983) over a large

portion of the historic and current range of sage-grouse

(Connelly and others 2004). The question of the quality of

crested wheatgrass seedings as habitat for sage-grouse is

difficult to answer since it depends on both spatial and

temporal scales that are rarely monitored (Connelly and

others 2004). The size of the seeding, juxtaposition on the

landscape in relation to suitable habitat, species composition of the seeding, and sagebrush cover are other factors

that affect the utility of crested wheatgrass seedings as

sage-grouse habitat. A recent review of sage-grouse habitat

needs and associated habitat threats does not directly identify introduced seedings as a factor in the decline of sagegrouse (Wambolt and others 2002). They indicated that the

extreme reduction in canopy cover of sagebrush, and associated loss in understory plant diversity typical of some

introduced species seedings, can significantly reduce sagegrouse habitat quality throughout the year.

Understory plant diversity is important to sage-grouse in

the spring and summer, whereas sagebrush provides essential cover and forage in the winter and late fall (Connelly and

others 2000; Wambolt and others 2002). Brood-rearing habitats with an array of plant species provide a diversity of

insects important to sage-grouse, especially during brood

rearing (Connelly and others 2000; Drut and others 1994).

The recently published “Guidelines for Sage-Grouse Habitat” (Connelly and others 2000) identified grass height (over

18 cm) and canopy cover (greater than 15 percent for breeding and brood-rearing habitats) as important habitat requirements for nesting sage-grouse. Therefore, crested wheatgrass, in the appropriate proportions, could provide similar

habitat structure compared to native bunchgrasses.

Connelly and others (2000) recommend that nonnative

species that are functionally equivalent to natives be used in

restoration projects if native forbs and grasses are unavailable.

Another situation where the use of crested wheatgrass

may be necessary is the restoration of habitat that is severely degraded and dominated by aggressive invasive species (Pyke 1994). Beginning in the 1930s numerous studies

were conducted that showed the superior competitive ability of crested wheatgrass with cheatgrass in the Intermountain area (Hull 1974; Hull and Holmgren 1964; Hull

and Pehanec 1947; Hull and Stewart 1948). Recently, the

establishment of crested wheatgrass as a “bridge” plant

community to replace cheatgrass-dominated lands for

future restoration to a more diverse plant community has

been proposed as an alternative to seeding a full complement

of native species in one treatment. This strategy, also referred to as “assisted succession” (Cox and Anderson 2004),

will be discussed later in this document.

It is known that sage-grouse will not thrive in large

homogenous stands of a single plant species (Crawford and

others 2004). The modification of composition and structure

of existing crested wheatgrass seedings to increase plant

diversity for sage-grouse should be implemented only after

considering social impacts, economic considerations, and

land-use objectives. This synthesis paper was not produced

to support the replacement of crested wheatgrass with

native species. It is intended to provide a review of existing

science and knowledge that can be used to increase diversity

and structure of crested wheatgrass seedings to benefit

sage-grouse, other wildlife species, and livestock. Local

land-use plans, science, and public input should be incorporated into the decisionmaking process when selecting

crested wheatgrass seedings to implement the strategies

described in this paper.

Steps Required to Enhance

Structure and Diversity in Crested

Wheatgrass Seedings ____________

The competitive characteristics of crested wheatgrass,

discussed in the previous section, illustrate the difficulty in

designing treatments to increase plant diversity in crested

wheatgrass seedings. This grass is extremely resistant to

grazing by herbivores. It has a variable response to mechanical or chemical treatments and a large viable seed reserve in

the soil that must be considered in any control treatment.

Increasing plant diversity in established stands of crested

wheatgrass is presented as a three-step process.

Step 1. Reduce competition of crested wheatgrass to facilitate the establishment and persistence of the

desired species.

Step 2. Introduce the desired plant(s) as seed or seedlings.

Step 3. Implement appropriate management and monitoring to maintain plant diversity of the seeding.

The discussion that follows focuses on step 1, because the

knowledge and literature on seed or plant application (step 2) and

managing restored seedings (step 3) are generally available.

Step 1: Reduce Crested Wheatgrass

Competition

Grazing by domestic livestock, mechanical or chemical

treatments, and fire are potential treatments that can be

implemented singly or in combination to reduce crested

wheatgrass competition prior to introducing desired plants

to a seeding. Periodic droughts also offer opportunities,

again singly or in combination with the above treatments, to

reduce crested wheatgrass competition.

Livestock Grazing—The design of livestock grazing

systems to maintain crested wheatgrass and reduce encroachment of sagebrush into these seedings has been studied for almost as long as crested wheatgrass has been used.
84 USDA Forest Service Proceedings RMRS-P-38. 2005

Pellant and Lysne Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings

Much of this early research focused on grazing systems and

utilization levels to maintain the productivity of seedings

and minimize the return of sagebrush in seeded areas (Cook

and others 1958; Hull and Klomp 1974) Sagebrush was

considered an “invader” in crested wheatgrass seedings

because it reduced the productive capability and economic

returns from seedings in direct proportion to the ratio of

shrub to grass. Rittenhouse and Sneva (1976) determined

that each 1 percent increase in Wyoming big sagebrush

canopy cover was associated with a 3.3 to 5.2 percent decline

in crested wheatgrass production in eastern Oregon. This

early research can now be used “in reverse” to develop

grazing systems to reduce crested wheatgrass competition

in order to increase plant diversity, especially shrubs, in

seedings.

Since crested wheatgrass can withstand heavy grazing

(Caldwell and others 1981; Cook and others 1958; Hull 1974;

Laycock and others 1981), reducing competition using only

livestock may be insufficient to permit establishment of

desirable seeded species. In general, high levels of utilization by livestock during the growing season reduces the

vigor of crested wheatgrass and may lead to mortality of

some, but not all, plants (Cook 1973; Wilson and Partel

2003). Early summer grazing may be detrimental to crested

wheatgrass due to lower carbohydrate (Trlica and Cook

1972) and nitrogen reserves (de Kroon and Bobbink 1997) of

grazed plants. Other studies have demonstrated that heavy

use alone (up to 70 percent) did not significantly affect

stands of crested wheatgrass (Frischknecht and Harris

1968; Lodge and others 1972; Springfield 1963). Heavy

utilization by livestock was also cited by some of these

authors as necessary to reduce development of crested

wheatgrass “wolf plants.” Wolf plants are crested wheatgrass plants that contain a higher proportion of dead stems

than consistently grazed plants and, as a result, are not

preferentially selected by livestock.

Olson and others (1988a,b) found that grazing to reduce

vigor, cause mortality, or reduce establishment of new

crested wheatgrass seedlings is most effective if the treatment is done during or immediately after tiller elongation

(internode elongation) and results in removal of the apical

meristem. These same studies showed that grazing crested

wheatgrass before internode elongation had little effect on

reducing tiller replacement and could increase tiller density

if grazing intensity and timing were not closely monitored.

Olson and others (1988b) indicated that short-duration

grazing at a conventional stocking rate in eastern Oregon

increased tiller density of crested wheatgrass. This study

also demonstrated that most crested wheatgrass tillers are

produced in the fall, overwinter, and flower the following

growing season. If an adjacent crested wheatgrass plant was

removed, the tiller production and resource uptake of its

neighbor was increased. Thus, attempts to reduce crested

wheatgrass by livestock grazing are influenced by growing

season conditions, level of utilization in relation to plant

phenology, degree of use of neighboring plant, and the

dynamics of tiller production.

It is well established that crested wheatgrass is adapted

to withstand heavy livestock use with minimal mortality.

How does livestock grazing affect the recruitment and establishment of crested wheatgrass? Most crested wheatgrass

recruitment occurs between the rows of established plants in

a seeding (Salihi and Norton 1987). The success of recruitment in the interspaces is reduced by the impacts of livestock

trampling since cattle (Bos sp.) generally avoid stepping on

plant tussocks (Balph and Malechek 1985). As crested

wheatgrass plants age, elevated tussocks develop because

of the plant’s caespitose growth form, further increasing

cattle avoidance of stepping on the mature plants. This

results in increased mortality of seedlings that are trampled

(Salihi and Norton 1987), and accelerates soil erosion and

compaction (Balph and others 1985) in the interspace areas.

In another study, Krzic and others (2000) stated that longterm grazing of crested wheatgrass did not result in degradation of soil properties, with one exception: soil compaction was

greater in seedings grazed in spring compared to native

rangeland. Salihi and Norton (1987) measured less than

1 percent crested wheatgrass seedling survival in grazed

plots compared to 12 percent survival in ungrazed plots.

The combination of properly timed livestock use to reduce

vigor and survival of mature crested wheatgrass plants

along with the trampling of new recruits in the interspace

areas should result, over time, in a reduction in both numbers and vigor of mature crested wheatgrass plants and

their recruitment potential. The decision to use these intensive grazing treatments must be weighed against the detrimental effects of heavy grazing on soil properties, weed

entry and/or expansion, and erosion potential as well as the

management objectives for the seeding.

Another benefit of livestock use at the appropriate time

and intensity in crested wheatgrass seedings is to facilitate

the return of sagebrush. As mentioned earlier, control of

“reinvading” sagebrush in crested wheatgrass seedings was

the focus of past research on treatments to physically remove the sagebrush or livestock management systems to

maintain the crested wheatgrass productivity and minimize

the reinvasion of sagebrush. It is well established that

sagebrush encroachment in seedings is less under light to

moderate spring livestock use, but increases under high

crested wheatgrass utilization levels for this same period

(subject to climatic, grazing management system, and initial

treatment variables) (Frischknecht and Harris 1968; Hull

and Klomp 1974; Laycock and Conrad 1981; Robertson and

others 1970). For example, crested wheatgrass utilization

levels of 80 percent on a Utah seeding resulted in loss in vigor

of crested wheatgrass and an increase in sagebrush

(Frischknecht and Harris 1968). By comparison, fall grazing

by cattle resulted in less sagebrush encroachment in seedings

when compared to heavy spring livestock use (Laycock and

Conrad 1981). However, grazing by sheep (Ovis aries) in fall

often resulted in reduced production or mortality of sagebrush (Frischknecht 1978).

Therefore, assuming that there is a sagebrush seed source

in or near the target crested wheatgrass seeding, a grazing

system that promotes heavy spring livestock use over a

period of years could promote an increase of sagebrush in

crested wheatgrass seedings. Angell (1997) found that this

same grazing management system would also promote the

survival of juvenile sagebrush plants due to decreased soil

water depletion by crested wheatgrass. He found that only

the short duration, double stocking rate treatment in spring

resulted in an increase in juvenile sagebrush plants when
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Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings Pellant and Lysne

compared to the continuous grazing and moderate short

duration grazing treatments. In a similar study, Owens and

Norton (1990) found that juvenile sagebrush survival was

greater in a pasture that received high intensity use for

repeated short durations (short duration grazing system)

during the growing season when compared to a traditional

continuous growing season treatment.

Thus, once juvenile sagebrush plants are established in a

seeding, continued heavy livestock use will accelerate sagebrush growth and potentially increase additional sagebrush

recruitment. This strategy is predicated on concentrated

heavy use of crested wheatgrass and may require temporary

fencing to concentrate livestock in a smaller portion of a

larger seeded pasture. The temporary fence could then be

moved to another portion of the seeding to increase sagebrush establishment over a larger area, if desired. Other

considerations in applying this strategy to increase sagebrush in crested wheatgrass seedings are the effects of soil

compaction, potential for weed entry, increased soil erosion,

and effects on residual native grasses and forbs in the heavy

use areas. Introduction of sagebrush seed may be required if

a seed source is not already present in or immediately

adjacent to the treatment area.

Drought and Livestock Grazing—Periodic droughts

provide another window of opportunity to reduce crested

wheatgrass density, especially when combined with properly timed, heavy levels of livestock use. Tiller regrowth of

crested wheatgrass was limited by clipping and drought over

a 2-year period (Busso and Richards 1995). They cautioned

that repeated late spring grazing under droughts lasting 2 or

more years could reduce the persistence of crested wheatgrass in a stand. Conversely, light or moderate grazing

(around 40 percent) of crested wheatgrass in a drought was

found to enhance production and survival because of a

decrease in the leaf area and associated respiration

(Mohammad and others 1982). In this same study, no plant

recovery occurred when water stress was severe and crested

wheatgrass defoliation was 80 percent.

Crested wheatgrass has the potential to recover rapidly

after a drought due to the high accumulation of total

nonstructural carbohydrate reserves accumulated in the

plant organs during times of stress (Busso and others 1990).

Thus, any benefits in reduction in competition of crested

wheatgrass achieved by livestock grazing during droughts

may be lost quickly if treatments to increase diversity are

not implemented in a timely manner. Another concern with

using drought and livestock to reduce competition of crested

wheatgrass is the opportunity for an increase in invasive

species during periods between droughts and the average or

above average precipitation periods following droughts

(Svejcar 2003). Heavy livestock use may also increase the

potential for loss of biological soil crusts and residual native

plants in the seeding (Anderson and others 1982; Kimball

and Schiffman 2003). Even with these concerns, livestock

grazing during multi-year droughts may reduce crested

wheatgrass competition sufficiently to allow successful reintroduction of desired species.

Herbicide Application—The application of an appropriate herbicide at the proper time can reduce perennial

grass density (Nelson and others 1970; Whisenant 1999). A

number of different herbicides are effective in reducing vigor

or causing mortality of crested wheatgrass. Glyphosate (N-

[phosphonomethyl] glycine) is a contact herbicide that stunts

or kills the entire plant upon application. Application of

glyphosate (trade name RoundupTM) reduced crested wheatgrass cover from 12 to 4 percent in 1 year and had no effect

in the second year of a 2-year study in Utah that looked at the

utility of several treatments to reduce competition prior to

seeding native species (Cox and Anderson 2004). This difference in effect between years, probably due to timing of

application, illustrates the importance of applying contact

herbicides at the appropriate phenological stage. In Canada,

a spring application of glyphosphate reduced crested wheatgrass by 50 percent, which was adequate control to establish

a native warm season grass seeded at a high application rate

(Bakker and others 1997).

Wilson and Partel (2003) applied multiple herbicide treatments to maximize the mortality of crested wheatgrass in

Canadian grasslands. A total of 13 glyphosphate applications over 6 years significantly reduced cover of crested

wheatgrass; however, the surviving plants in the herbicide

treatment area produced 42 seedheads per m2 compared to

12 seedheads per m2 in the control. Crested wheatgrass

seedlings emerging from the seedbank were not significantly different between the herbicide treatment and control (average density of 284.4 seedlings per m2). Even though

crested wheatgrass was not eliminated with the herbicide

treatments in this study, Bakker and others (2003) reported

that native species diversity and abundance were enhanced

on these study sites. They reported that the careful application of glyphosate by wicking or spraying prior to the active

growth of warm season native species can suppress crested

wheatgrass and promote native species establishment. They

recommend considering cultivation prior to seeding and

applying multiple control treatments (herbicide and intensive grazing) in hot dry years to further reduce crested

wheatgrass competition if herbicide application alone is not

adequate.

Another Canadian project evaluated multi-year application of glyphosphate to reduce crested wheatgrass competition before seeding native species (Ambrose and Wilson

2003). Glyphosphate was applied as a spray in the spring of

the first year and applied with a wick applicator in the 3

subsequent years. Surprisingly, emergence of crested wheatgrass seedlings from the seedbank was not decreased by 4 years

of glyphosphate treatments, due primarily to the tripling in

number of seed heads on surviving plants in the herbicide plots

compared to the control plots. The impacts of releasing

crested wheatgrass from intraspecific competition with

glyphosphate and thereby increasing seed production on

remaining plants must be considered when selecting treatments to reduce crested wheatgrass competition.

When Romo and others (1994) investigated the effects of

a combination of fall burning followed by a spring application of glyphosate on crested wheatgrass mortality in

Canada, they found that burning had little effect on crested

wheatgrass survival, while glyphosphate applied early in

the growing season on the burned crested wheatgrass reduced cover from 78 to 35 percent on one site and from 81 to

55 percent on another site. In another study, Romo and

others (1994) applied mowing in the fall to reduce crested
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Pellant and Lysne Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings

wheatgrass vigor followed by an application of glyphosate on

individual plants and recorded 100 percent crested wheatgrass mortality. They also observed total elimination of

crested wheatgrass with an application of 25 percent

glyphosphate in early spring when two to four leaves per

tiller were present.

Another consideration in using glyphosphate to reduce

crested wheatgrass is the differential effect that this herbicide appears to have on different species of crested wheatgrass. Lym and Kirby (1991) found that ‘Fairway’ crested

wheatgrass was less susceptible to glyphosphate in terms of

yield than was ‘Nordan’ crested wheatgrass. Also glyphosate

generally does not interfere with the establishment of seeded

species (Bakker and others 1997; Masters and Sheley 2001)

since it is bound to the soil once applied and is not available

for uptake by plants. On the negative side, since glyphosphate

is a contact herbicide, it has no residual effect on crested

wheatgrass regrowth and may need to be applied multiple

times in the same growing season or over multiple years,

depending on climatic conditions and plant phenology and

growth patterns.

Paraquat (1,1 ́-dimethyl-4,4 ́-bipyridinium ion) is another

herbicide that has been used to treat crested wheatgrass.

Sneva (1970) found that paraquat applied for 3 consecutive

years did not significantly reduce crested wheatgrass yield

in the fourth year. In this study, clipping crested wheatgrass

to ground level in May of each year was more effective in

reducing the percent of apical meristems than was the

herbicide application. Atrazine and simazine were evaluated as tools to rejuvenate weed infested seedings in Nevada

(Eckert 1979). The reduction of weedy competition in the

stand by these herbicides resulted in slightly more crested

wheatgrass seed production and minimal mortality on treated

compared to control sites. Crested wheatgrass seedling

production was significantly greater in the atrazine treated

plots compared to the control, indicating that the reduction

in weedy competition not only favored seed production but

greatly enhanced seedling establishment.

In summary, herbicides can be very effective in controlling

crested wheatgrass, especially when combined with other

treatments such as burning or mowing. Label restrictions on

their use should be closely followed in order to minimize

adverse effects. If complete crested wheatgrass mortality is

not obtained (usually the case), the seed production on

surviving plants increases significantly and provides significant competition with desirable plants introduced on the

treated areas. As Whisenant (1999) points out, effective

herbicide use requires knowledge of individual site characteristics and knowledge of herbicide effects on the individual

species and the environment.

Mechanical Treatments—Mechanical treatments can

be used to either physically remove crested wheatgrass

biomass (for example, mowing) to reduce plant vigor or cause

mortality, or uproot plants and cause direct mortality (for

example, plowing). Mechanical removal of live crested wheatgrass foliage will be discussed first, followed by an overview

of equipment that can be used to cause direct mortality.

Clipping studies to simulate grazing have been previously

discussed in the Livestock Grazing section of this paper,

and the reader is encouraged to review that information as

it applies to the effects of mowing described in this section.

Lodge (1960) compared mowing in the fall, burning in

spring and fall, and double disking in the fall. He found that

mowing had little effect on floristic composition or in reducing basal area of crested wheatgrass in Canada. Double

disking was the only treatment that significantly reduced

crested wheatgrass basal area (from 6.6 percent on the

control to 2.7 percent in the treatment areas); this treatment

effect disappeared within 2 years. A clipping study to reduce

crested wheatgrass competition was conducted in northern

Utah by Cook and others (1958). They hand-clipped crested

wheatgrass plants at 1- and 3-inch stubble heights throughout the growing season over a 5-year period. The 1-inch

clipping height and more frequent clipping treatments reduced yield, vigor, and seed production of crested wheatgrass more than did the 3-inch clipping height and less

frequent clipping treatments. Seed production, as expressed

by number of spikes per plant, was not significantly affected

by clipping height; however, increasing the frequency of

harvesting decreased the number of spikes produced. Finally, this study documented that frequency and season of

clipping were the most influential factors affecting viable

seed production. At the end of the 5-year study period,

control plants produced 1,834 viable seeds per plant, while

clipping once in mid-June or early July for 5 years reduced

the number of seeds per plant to nearly zero.

Lorenz and Rogler (1962) compared several mechanical

techniques to “renovate” stands of crested wheatgrass in

North Dakota. Plowing in spring eliminated crested wheatgrass production for 2 years, while a spring scarification

treatment (heavy field cultivation that uprooted about onethird of the plants) significantly reduced yields in only the

first year following treatment. In subsequent years the

scarified treatment produced more herbage than the control

plot in one year and similar yields in the remaining years of

the study. The authors urged caution with the plowing

treatment due to the potential for increased wind erosion.

Bakker and others (1997) rototilled crested wheatgrass

plots in May on a sandy site in Canada, reducing cover of

crested wheatgrass from around 40 percent on control plots

to 20 percent on the treatment plots in August of the same

year. Finally, Cox and Anderson (2004) investigated the

effectiveness of two tillage treatments and a herbicide treatment in reducing crested wheatgrass competition prior to

seeding native species. Tillage treatments were done in

February in 2 consecutive years on a crested wheatgrass

seeding in an arid (average annual precipitation of 7 inches)

portion of Utah. The tilling treatment was done with a

cultivator that removed all vegetation and mixed the soil to

a 7-inch depth, while the harrowing treatment was done

with a field harrow that uprooted some, but not all plants.

Tillage was more effective than harrowing in reducing

crested wheatgrass cover in this study. The control plots

averaged 12 to 4 percent crested wheatgrass cover during

the 2-year study compared to 1 to 2 percent cover on tilled

and 4 to 7 percent cover on harrowed plots.

Another category of mechanical equipment, the

interseeders and transplanters, remove plant competition

in narrow bands and seed (interseeder) or plant seedlings

(transplanter) in a one-pass operation (Giunta and others

1975; Stevens 1994; Stevens and others 1981; Wiedemann

2005). Scalping to reduce plant competition is generally

done with either modified disks or a plow pulled behind a
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Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings Pellant and Lysne

tractor. The seeder or transplanter is mounted immediately

behind the disk or plow. Recommended widths for scalping

crested wheatgrass prior to seeding shrub seeds are 40 to 60

inches (Van Epps and McKell 1978). This width should be

adjusted according to density, vigor, and growth form of

existing vegetation, the species to be interseeded, and local

site conditions (Stevens 1994). A side benefit of scalping is

that the scalp captures and holds additional moisture from

snow and rain, which enhances seedling establishment and

growth (Stevens 1994). An indepth description of interseeders

and transplanters can be found in Chapter 28 of Restoring

Western Ranges and Wildlands (Monsen and others 2004b).

Other equipment not specifically addressed in studies

cited above that could be used to reduce crested wheatgrass

competition includes pipe harrows, anchor chains with welded

railroad rails (for example, Ely and Dixie Sager chains), and

the disk chain (Monsen and others 2004a). Effectiveness of

these types of equipment in providing crested wheatgrass

control is expected to be moderate to excellent, although

published studies to support their use for crested wheatgrass control are few.

In summary, the use of mechanical equipment to reduce

crested wheatgrass competition will vary in effectiveness,

dependent upon a wide array of factors. Some cautions on

the use of plows or disks include increased chance of soil

erosion and weed entry, loss of residual native plants and

biological crusts, and treatment costs.

Step 2: Introduce Desired Species

The challenges in controlling crested wheatgrass competition, described in Step 1, must be resolved prior to implementing the seeding or planting treatments outlined in Step 2. The

benefits of increasing plant diversity in grass monocultures

include improved habitat, greater species richness and community diversity, improved aesthetics, more soil cover

(Stevens 1994), and increased diversity of birds, mammals,

reptiles, and insects (Reynolds 1980).

Most of the treatments implemented in the past to increase diversity in crested wheatgrass stands have involved

interseeding or transplanting single species or a few species

such as big sagebrush, rubber rabbitbrush (Chrysothamnus

nauseosus), fourwing saltbush (Atriplex canescens), antelope bitterbrush, Lewis flax (Linum perenne), Palmer penstemon (Penstemon palmeri), western yarrow (Achillea

millefolium), and globemallow (Sphaeralcea spp.) (Monsen

and Shaw 1983; Pendery and Provenza 1987; Stevens 1994).

However, single rows of shrubs or forbs in monocultures of

crested wheatgrass may not meet all of the resource (for

example, sage-grouse habitat) or management objectives for

a particular area.

Step 2 involves the selection of adapted species to plant

and appropriate equipment to implement the planting. It is

essential to select the species and seed mixtures that meet

resource objectives and are adapted to the ecological site(s)

that will be seeded. Nonadapted seeds may respond differently to germination cues and germination may occur at an

inappropriate time, resulting in seeds that fail to germinate

or persist (Meyer 1994). Additional considerations for seed

mixture development include the potential for interspecific

interactions among the species in the seed mixture during

the establishment phase, the ability of plants to coexist, and

the ability of the species to regenerate itself on the site

(Archer and Pyke 1991; Pyke 1994; Pyke and Archer 1991).

If the objective of the crested wheatgrass treatment(s) is

to restore ecosystem functioning and biological diversity to

a site, this will often require the use of native species

(Lesica and Allendorf 1999). Native species introduced into

a crested wheatgrass seeding may facilitate recruitment of

additional native species. For instance, Frischknecht and

Bleak (1957) found that seeded stands of bluebunch wheatgrass were more likely to permit sagebrush seedling recruitment than seeded stands of crested wheatgrass. Introduced

species may also increase the diversity of a crested wheatgrass seeding, improving it as habitat for sage-grouse. Dryland alfalfa (Medicago sativa) and small burnet (Sanguisorba

minor) are introduced forbs that are preferred by sagegrouse that can be successfully reintroduced into crested

wheatgrass seedings. It is important to select site-adapted

species (native or introduced) that are competitive in the

posttreatment environment and that will be maintained

over the long term with livestock management systems.

The selection of a seed mixture should not be done without

consideration of how seed will be distributed during the

planting process. Rangeland drills vary considerably in

their ability to seed native species. If suitable equipment is

not available to properly seed a species in the proposed seed

mix, the mix should be changed or the proper equipment

secured. Another factor to consider is that some site preparation treatments, such as plowing or disking, may create

an unfavorable planting seedbed that requires additional

treatments. Harrowing or cultipacking after these surface

disturbing treatments may be required to mitigate these

unsatisfactory seedbed surfaces (Whisenant 1999).

Direct seeding by drilling or aerially broadcasting seed is

relatively inexpensive, widely applicable, and under appropriate seedbed conditions, provides good plant establishment (Whisenant 1999). Applying seed with a rangeland

drill is considered the best method for establishing species

with large, hard seeds because the seed is placed in contact

with the soil and at an appropriate depth (Hull 1948; Pyke

1994). However, seeding many native species with the standard rangeland drill is problematic given the lack of control

of seeding depth, variable seed coverage with soil, and

absence of a mechanism to improve soil to seed contact.

Surface obstructions such as rocks, steep slopes, and soddy

vegetation also limit the effectiveness of rangeland drills in

establishing any seed mixture, especially native forbs and

grasses. One unknown in the use of rangeland drills to seed

diverse seed mixtures into crested wheatgrass seedings is

the effectiveness of these drills in cutting through the dead

plant crowns and the shallow root mass of the seeding. If this

is a problem, the deep furrow rangeland drill (Hull and

Stewart 1948), which has a double furrow opener, may be

more effective in soddy conditions than the rangeland drill,

which has a single furrow opener. The single disk or double

disk opener on the rangeland drill does create a furrow that

can capture and store water for seedlings. However, the soil

disturbance created by this drill also opens the plant community for the entry of other invasive species.

Another option for ground application seeding into treated

crested wheatgrass stands is the use of a minimum till drill
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Pellant and Lysne Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings

that creates less soil disturbance than the rangeland drill.

The Truax and Amazon drills are minimum till drills that

can place seed at different depths, and their press wheels

improve soil to seed contact. An overview of rangeland drills,

manufacturer’s specifications, and contact information is

the Revegetation Equipment Catalog available online at

http://reveg-catalog.tamu.edu (Wiedemann 2005). This catalog also contains similar information on most of the equipment discussed in this paper. The reader is encouraged to

utilize this Web site for all treatments requiring the use of

equipment.

Aerial broadcasting is often easier and less expensive than

ground application methods because large areas can be

seeded quickly and topography or slopes are generally not a

limiting factor (Monsen 2000). Aerially broadcasting seed

followed by cultipacking, harrowing, or dragging a chain

over the surface, where slope or surface rock is not limiting,

places the seed in contact with the soil; however, seeding

depth is not uniform (Pyke 1994; Stevens 2004). Livestock

trampling has been suggested as another alternative for

covering seed that has been aerially applied. Eckert and

others (1986) found that heavy livestock trampling appeared to favor the emergence of sagebrush and weedy

annual forbs, but was detrimental to the emergence of

perennial grasses and forbs. Aerial seeding native species

mixes into treated crested wheatgrass stands without some

sort of incorporation into the soil is not advised. Given the

high cost of seed and the different seedbed requirements of

native species (seeding rates are generally doubled on aerial

seedings), seeding with rangeland drills is recommended

over aerial seeding with or without seed coverage.

Alternatives to ground or aerial application of seed include transplanting individual plants from existing populations (“wildings”) or planting container stock or bare-root

seedlings grown from seed. In arid and semiarid environments, transplanting young plants may be a more reliable,

albeit a considerably more expensive method for establishing native species in crested wheatgrass seedings. Transplanting young plants bypasses the high-risk germination

and seedling stage. In addition, transplanting may enhance

the success of species that do not establish rapidly from seed

and provide larger plants that are more capable of coping

with competition and herbivory (Archer and Pyke 1991; Van

Epps and McKell 1980; Whisenant 1999). If small islands of

native species are desired in a crested wheatgrass seeding or

if greater native plant diversity is desired in an existing

native species seeding, transplanting wildings, bare-root, or

containerized stock of desired forbs or shrubs may be a good

option. However, costs of treating larger acreages this way

will generally be prohibitive.

The use of livestock to disseminate seed of desired species

via dung (Auman and others 1998; Doucette and others

2001; Ocumpaugh and others 1996; Welch 1985) into crested

wheatgrass seedings is another option since livestock preferentially graze these areas. Seeds ingested by cattle are

deposited in a moist, nutrient-rich medium that may facilitate germination and establishment of ingested seeds and

may result in patches of desirable species (Archer and Pyke

1991). Fecal-seeding offers a nonintrusive, relatively low

cost method of seeding small areas (Archer and Pyke 1991;

Shinderman and Call 2001). Seeding response can be slow

and sporadic, and there is the potential for the introduction

and spread of exotic species by livestock (Auman and others

1998; De Clerck-Floate 1997; Lyon and others 1992; Pleasant and Schlather 1994). A study by Auman and others

(1998) found that cattle dung provided favorable conditions

for the germination of crested wheatgrass as well as

cheatgrass. Also, livestock grazing would need to be closely

monitored to ensure that livestock did not overutilize and

eliminate the very plants they were dispersing (Archer and

Pyke 1991).

Step 3: Posttreatment Management

The long-term success of any project implemented to

increase plant diversity in crested wheatgrass seedings is

dependent on applying appropriate management during the

establishment and postestablishment period. Documentation of implementation practices and the effectiveness of

treatments must be conducted via a well designed monitoring program in order to adjust management now and

design more effective projects in the future (for example,

implement an adaptive management program). An adaptive management program is not possible without good

implementation information combined with sound effectiveness monitoring.

Livestock Management—It is essential that livestock

grazing and rest intervals are matched with the phenology

and life history attributes of desired plant species (Archer

and Pyke 1991; Holechek 1983). Grazing should be restricted until plants are adequately established and sexually reproducing (Pyke 1994). Many plants require at least

2 years, and as many as 5 years, to become established with

adequate root systems to endure grazing (Pyke 1994;

Stevens 1994; Vallentine 1989; Vallentine and others 1963;

West and Hassan 1985). Areas seeded to shrubs must be

protected from grazing during the establishment period

(Ganskopp and others 1999; Richardson and others 1984).

Plant seedlings are particularly sensitive to herbivory because they have low nutrient and energy reserves and

shallow, low-density root systems relative to adult plants

(Archer and Pyke 1991; Holechek 1983).

Once the plant establishment period (period of time when

livestock were excluded from the project area) has passed,

an appropriate livestock management plan must be followed to maintain the diversity restored in the crested

wheatgrass seeding (Archer and Pyke 1991). If plant diversity is increased in a crested wheatgrass seeding to benefit

sage-grouse, additional livestock or recreation management

changes may be necessary to maintain structure, composition, and forage quality to meet seasonal habitat requirements. Impacts of livestock grazing can be positive, negative, or neutral to sage-grouse, depending on the timing and

intensity of livestock grazing and which seasonal habitat is

being considered (Crawford and others 2004). Heavy livestock grazing can reduce grass competition and increase

sagebrush density (Crawford and others 2004; Vallentine

1989) or it can decrease big sagebrush seedling survival under

certain management systems (Owens and Norton 1990). In

general, the season and duration of livestock use and the

stocking rate should be managed to promote optimum growth

of forbs, grasses, and sagebrush to maximize habitat values

for sage-grouse (Beck and Mitchell 2000).
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Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings Pellant and Lysne

Monitoring—Monitoring involves the orderly collection

of data, analysis, and evaluation of data. Combined with

experience, monitoring is a powerful tool to improve the

effectiveness of restoration efforts now and into the future.

Implementation monitoring includes summarizing how,

what, where, and when treatments were actually implemented. The timing of treatments, conditions during application of treatments (for example, was the soil dry or frozen

when seeding occurred), and posttreatment events (for example, Mormon cricket density was high the first year

following seeding) are all important factors in evaluating

treatment effectiveness. The origin and percent of pure live

seed of each species in the seed mixture should also be

documented in the project file to improve the accuracy of

seeding establishment interpretations.

Effectiveness monitoring measures the success of the treatments that were implemented relative to the project objectives. Implementation monitoring provides the context to

evaluate the effectiveness of the treatments. It is important

that project objectives be developed before selecting monitoring protocols. The sage-grouse guidelines developed by

Connelly and others (2000) provide a good starting point to

develop sage-grouse habitat objectives in crested wheatgrass

seedings proposed for treatments to increase their diversity.

Monitoring information, if collected appropriately, provides the framework to implement an adaptive management

program to improve restoration practices in the future.

Adaptive management acknowledges uncertainty and imperfect knowledge in implementing projects (Walters 1986),

and encourages research and management to be conducted

simultaneously (Smallwood and others 1999; Walters and

Holling 1990). An adaptive management approach would be

especially helpful in identifying treatments that are effective in reducing crested wheatgrass competition (Step 1). To

maximize the utility of this approach, different treatments

would be implemented and evaluated on the same project,

promoting a better understanding of treatment effectiveness. At a minimum, adequate monitoring data should be

collected to determine if short- and long-term management

objectives are met when restoring diversity of crested wheatgrass seedings.

Summary ______________________

This review identifies some of the actions that can be

taken to increase plant diversity in crested wheatgrass

seedings for sage-grouse and other uses. The importance of

proper planning and posttreatment management has been

stressed as an essential component of a three-step process to

convert parts of existing crested wheatgrass seedings into

more diverse plant communities. This three-step process

and the treatments associated with it could be used as part

of a more ambitious strategy to first convert cheatgrass

monocultures (Allen 1995; Pellant 1990; Tausch and others

1995) into perennial grasslands followed by the steps described above to increase plant diversity in these crested

wheatgrass grasslands, for example, assisted succession as

described by Cox and Anderson (2004). This strategy provides a bridge between the difficult conversion of exotic

annual grasslands into native plant communities.

It is important to remember that crested wheatgrass

seedings have been an important management tool used to

increase livestock production, reduce weed problems and

wildfires, and mitigate soil erosion potential following disturbances since they were first established in the late 1930s.

Regardless of whether our objectives now are to increase

plant diversity in selected crested wheatgrass seedings or as

part of a larger strategy to reduce cheatgrass domination in

the Intermountain region, the application of good science

and professional experience tempered with results from

monitoring studies should guide our actions. Sage-grouse,

other wildlife species, and all resource uses will benefit from

an objective-based approach (both at the site and landscape

levels) to restoring plant diversity to selected crested wheatgrass seedings. As always, social, economic, and political

values will provide the context for these important restoration decisions.

Acknowledgment _______________

The authors thank Nancy Shaw and Danielle Scholten for

the reviews and edits of this manuscript.

References _____________________

Ackigoz, E.; Knowles, R. P. 1983. Long-term storage of grass seeds.

Canadian Journal of Plant Science. 63: 669–674.

Allen, E. B. 1995. Restoration ecology: limits and possibilities in

arid and semiarid lands. In: Roundy, B. A.; McArthur, E. D.;

Haley, J. S.; Mann, D. K., comps. Wildland shrub and arid land

restoration symposium: proceedings. Gen. Tech. Rep. INT-GTR315. Ogden, UT: U.S. Department. of Agriculture, Forest Service,

Intermountain Research Station: 7–15.

Ambrose, L. G.; Wilson, S. D. 2003. Emergence of the introduced

grass Agropyron cristatum and the native grass Bouteloua gracilis in a mixed-grass prairie restoration. Restoration Ecology. 11:

110–115.

Anderson, D. C.; Harper, K. T.; Holmgren, R. C. 1982. Factors

influencing development of cryptogamic soil crusts in Utah

deserts. Journal of Range Management. 35: 180–185.

Anderson, J. E.; Inouye, R. S. 2001. Landscape-scale changes in

plant species abundance and biodiversity of a sagebrush steppe

over 45 years. Ecological Monographs. 71: 531–556.

Angell, Raymond F. 1997. Crested wheatgrass and shrub response

to continuous or rotational grazing. Journal of Range Management. 50: 160–164.

Archer, S.; Pyke, D. A. 1991. Plant-animal interactions affecting

plant establishment and persistence on revegetated rangeland.

Journal of Range Management. 44: 558–565.

Auman, B. S.; Call, C. A.; Wiedmeier, R. D. 1998. Crested wheatgrass establishment in livestock dung deposited on degraded

rangeland vegetation types in the Intermountain West, USA.

Arid Soil Research and Rehabilitation. 12: 317–333.

Bakker, J.; Wilson, S. 2001. Competitive abilities of introduced and

native grasses. Plant Ecology. 157: 117–125.

Bakker, J. D.; Christian, J.; Wilson, S. D.; Waddington, J. 1997.

Seeding blue grama in old crested wheatgrass fields in southwestern Saskatchewan. Journal of Range Management. 50:

156–159.

Bakker, J. D.; Wilson, S. D.; Christian, J. M.; Li, X.; Ambrose; L. G.;

Waddington, J. 2003. Contingency of grassland restoration on

year, site, and competition from an introduced grass. Ecological

Applications. 13: 137–153.

Balph, David F.; Malecheck, John C. 1985. Cattle trampling of

crested wheatgrass under short-duration grazing. Journal of

Range Management. 38: 226–227.

Beck, J. L.; Mitchell, D. L. 2000. Influences of livestock grazing on

sage-grouse habitat. Wildlife Society Bulletin. 28: 993–1002.
90 USDA Forest Service Proceedings RMRS-P-38. 2005

Pellant and Lysne Strategies to Enhance Plant Structure and Diversity in Crested Wheatgrass Seedings

Blaisdell, J. P. 1949. Competition between sagebrush seedlings and

reseeded grasses. Ecology. 30: 512–519.

Bleak, A. T.; Miller, Warren G. 1955. Sagebrush seedling production

as related to time of mechanical eradication. Journal of Range

Management. 8: 66–69.

Broersma, K.; Krzic, M.; Thompson, D. J.; Bomke, A. A. 2000. Soil

and vegetation of ungrazed crested wheatgrass and native rangelands. Canadian Journal of Soil Science. 80: 411–417.

Busso, C. A.; Richards, J. H. 1995. Drought and clipping effects on

tiller demography and growth of two tussock grasses in Utah.

Journal of Arid Environments. 29: 239–251.

Busso, C. A.; Richards, J. H.; Chatterton, N. J. 1990. Nonstructural

carbohydrates and spring regrowth of two cool-season grasses:

interaction of drought and clipping. Journal of Range Management. 43: 336–343.

Caldwell, M. M.; Eissenstat, D. M.; Richards, J. H.; Allen, M. F.

1985. Competition for phosphorus: differential uptake from dualisotope labeled interspaces between shrub and grass. Science.

229: 384–386.

Caldwell, M. M.; Richards, J. H.; Johnson, D. A.; Nowak, R. S.;

Dzurec, R. S. 1981. Coping with herbivory: photosynthetic capacity and resource allocation in two semiarid Agropyron bunchgrasses. Oecologia. 50: 14–24.

Clinton, W. J. 1999. Executive Order 13112—invasive species.

Federal Register. 64: 6183–6186.

Connelly, J. W.; Knick, S. T.; Schroeder, M. A.; Stiver, S. J. 2004.

Conservation assessment of greater sage-grouse and sagebrush

habitats. Unpublished report on file at: Western Association of

Fish and Wildlife Agencies, Cheyenne, WY. 610 p.

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.

Cook, C. W.; Lewis, C. E. 1963. Competition between big sagebrush

and seeded grasses on foothill ranges in Utah. Journal of Range

Management. 16: 245–250.

Cook, C. Wayne. 1973. Forage utilization, daily intake, and nutrient

value of desert range. In: Proceedings: 3rd workshop of the United

States/Australia Rangelands Panel; 1973 March 26–April 5;

Tucson, AZ. Denver, CO: Society for Range Management: 47–50.

Cook, C. Wayne; Stoddart, L. A.; Kinsinger, Floyd E. 1958. Responses of crested wheatgrass to various clipping treatments.

Ecological Monographs. 28: 237–273.

Cox, R. D.; Anderson, V. J. 2004. Increasing native diversity of

cheatgrass-dominated rangeland through assisted succession.

Journal of Range Management. 57: 203–210.

Crawford, J. A.; Olson, R. A.; West, N. E.; Mosley, J. C.; Schroeder,

M. A.; Whitson, T. D.; Miller, R. F.; Gregg, M. A.; Boyd, C. S. 2004.

Ecology and management of sage-grouse and sage-grouse habitat. Journal of Range Management. 57: 2–19.

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

exotic grasses, the grass/fire cycle, and global change. Annual

Review of Ecology and Systematics. 23: 63–87.

De Clerck-Foate, R. 1997. Cattle as dispersers of hounds-tongue on

rangeland in southeastern British Columbia. Journal of Range

Management. 50: 239–243.

de Kroon, H.; Bobbink, R. 1997. Clonal plant dominance under

elevated nitrogen deposition, with special reference to

Brachypodium pinnatum in chalk grassland. In: de Kroon, H.;

van Groenendael, J. The ecology and evolution of clonal plants.

Leiden, The Netherlands: Backhuys Publishers: 359–379.

Dewey, D. R.; Asay, K. H. 1975. The crested wheatgrasses of Iran.

Crop Science. 15: 844–849.

Dillman, A. C. 1946. The beginnings of crested wheatgrass in

North America. Journal of the American Society of Agronomy.

38: 237–250.

Doucette, K. M.; Wittenberg, K. M.; McCaughey, W. P. 2001. Seed

recovery and germination of reseeded species fed to cattle. Journal of Range Management. 54: 575–581.

Drut, M. S.; Crawford, J. A.; Gregg, M. A. 1994. Brood habitat use

by sage-grouse in Oregon. Great Basin Naturalist. 54: 170–176.

Eckert, Richard E., Jr. 1979. Renovation of sparse stands of crested

wheatgrass. Journal of Range Management. 32: 332–336.

Eckert, R. E., Jr.; Peterson, E. E.; Meurisse, M. S.; Stephens, J. L.

1986. Effects of soil morphology on emergence and survival of

seedlings in big sagebrush communities. Journal of Range Management. 39: 414–420.

Eissenstat, D. M.; Caldwell, M. M. 1988. Competitive ability is

linked to rates of water extraction: a field study of two aridland

tussock grasses. Oecologia. 75: 1–7.

Fortier, M. I. 2000. Wyoming big sagebrush establishment on mined

lands: seeding rates and grass competition. Laramie, WY: University of Wyoming. 66 p. Thesis.

Frischknecht, N. C.; Bleak, A. T. 1957. Encroachment of big sagebrush on seeded range in northeastern Nevada. Journal of Range

Management. 10: 165–170.

Frischknecht, N. C.; Harris, L. E. 1968. Grazing intensities and

systems on crested wheatgrass in central Utah: response of

vegetation and cattle. Tech. Bull. 1388. Washington, DC: U.S.

Department of Agriculture. 47 p.

Frischknecht, Neil C. 1978. Biological methods: a tool for sagebrush

management. In: Sagebrush symposium: proceedings; 1978 April;

Logan, UT. Logan: Utah State University, College of Natural

Resources: 121–128.

Ganskopp, D. T.; Svejcar, T.; Taylor, F.; Farstvedt, J.; Painter, K.

1999. Seasonal cattle management in 3 to 5 year old bitterbrush

stands. Journal of Range Management. 52: 166–173.

Giunta, B. C.; Christensen, D. R.; Monsen, S. B. 1975. Interseeding

shrubs in cheatgrass with a browse seeded-scalper. Journal of

Range Management. 28: 398–402.

Hall, D. B.; Anderson, V. J.; Monsen, S. B. 1999. Competitive effects

of bluebunch wheatgrass, crested wheatgrass, and cheatgrass on

antelope bitterbrush seedling emergence and survival. Res. Pap.

RMRS-RP-16. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 7 p.

Heidinga, L.; Wilson, S. D. 2002. The impact of an invading alien

grass (Agropyron cristatum) on species turnover in native prairie.

Diversity and Distributions. 8: 249–258.

Holechek, J. L. 1981. Crested wheatgrass. Rangelands. 3: 151–153.

Holechek, J. L. 1983. Considerations concerning grazing systems.

Rangelands. 5: 208–211.

Hubbard, R. L. 1957. The effects of plant competition on the growth

and survival of bitterbrush seedlings. Journal of Range Management. 10: 135–137.

Hull, A. C., Jr. 1948. Depth, season, and row spacing for planting

grasses on southern Idaho range lands. Journal of the American

Society of Agronomy. 40: 960–969.

Hull, A. C., Jr. 1974. Species for seeding arid rangeland in southern

Idaho. Journal of Range Management. 27: 216–218.

Hull, A. C., Jr.; Holmgren, R. C. 1964. Seeding southern Idaho

rangelands. Res. Pap. INT-10. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Forest and Range

Experiment Station. 31 p.

Hull, A. C., Jr.; Klomp, G. J. 1966. Longevity of crested wheatgrass

in the sagebrush-grass type in southern Idaho. Journal of Range

Management. 19: 5–11.

Hull, A. C., Jr.; Klomp, G. J. 1974. Yield of crested wheatgrass under

four densities of big sagebrush in Southern Idaho. Tech. Bull.

1483. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service. 38 p.

Hull, A. C., Jr.; Pearse, C. K. 1943. How to reseed southern Idaho

rangelands. Res. Pap. INT-2. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Forest and Range

Experiment Station. 22 p.

Hull, A. C., Jr.; Pehanec, J .F. 1947. Cheatgrass—a challenge to

range research. Journal of Forestry. 45: 555–564.

Hull, A. C., Jr.; Stewart, G. 1948. Replacing cheatgrass by reseeding

with perennial grass on southern Idaho ranges. American Society

of Agronomy Journal. 40: 694–703.

Kimball, S.; Shiffman, P. M. 2003. Differing effects of cattle grazing

on native and alien plants. Conservation Biology. 17: 1681–1693.

Krzic, M.; Broersma, K.; Thompson, D. J.; Bomke, A. A. 2000. Soil

properties and species diversity of grazed crested wheatgrass and

native ranges. Journal of Range Management. 53: 353–358.

Laycock, W. A.; Conrad, P. W. 1981. Responses of vegetation and

cattle to various systems of grazing on seeded and native mountain rangelands in eastern Utah. Journal of Range Management.

34: 52–58.
USDA Forest Service Proceedings RMRS-P-38. 2005 91

Lesica, P.; Allendorf, F. W. 1999. Ecological genetics and the

restoration of land communities: mix and match? Restoration

Ecology. 7: 42–50.

Lodge, R. W. 1960. Effects of burning, cultivating, and mowing on

the yield and consumption of crested wheatgrass. Journal of

Range Management. 13: 318–321.

Lodge, R. W.; Smoliak, S.; Johnston, A. 1972. Managing crested

wheatgrass pastures. Publ. 1473. Ottawa: Agriculture Canada.

20 p.

Looman, J.; Heinrichs, D. H. 1973. Stability of crested wheatgrass

pastures under long-term pasture use. Canadian Journal of Plant

Science. 53: 501–506.

Lorenz, Russell J.; Rogler, George A. 1962. A comparison of methods

of renovating old stands of crested wheatgrass. Journal of Range

Management. 15: 215–219.

Lym, R. G.; Kirby, D. R. 1991. Effect of glyphosate on introduced and

native grasses. Weed Technology. 5: 421–425.

Lyon, D. J.; Baltensperger, D. D.; Rush, I. G. 1992. Viability,

germination, and emergence of cattle-fed jointed goatgrass seed.

Journal of Production Agriculture. 5: 282–285.

Marlette, G. M.; Anderson, J. E. 1986. Seed banks and propagule

dispersal in crested wheatgrass stands. Journal of Applied Ecology. 23: 161–175.

Masters, R. A.; Sheley, R. L. 2001. Principles and practices for

managing rangeland invasive plants. Journal of Range Management. 54: 502–517.

Meyer, Susan E. 1994. Germination and establishment ecology of

big sagebrush: implications for community restoration. In: Monsen,

Stephen B.; Kitchen, Stanley G., comps. Proceedings—ecology

and management of annual rangelands symposium: proceedings;

1992 May 18–21; Boise, ID. Gen. Tech. Rep. INT-GTR-313.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 244–251.

Mohammad, Noor; Dwyer, Don D.; Busby, F. E. 1982. Responses of

crested wheatgrass and Russian wildrye to water stress and

defoliation. Journal of Range Management. 35: 227–230.

Monsen, S. B. 2000. Establishment of big sagebrush (Artemisia

tridentata) in semiarid environments. In: Entwistle, P. G.; DeBolt,

A. M.; Kaltenecker, J. H.; Steenhof, K., comps. Sagebrush steppe

ecosystems symposium: proceedings. BLM/ID/PT-001001+1150.

Boise, ID: U.S. Department of the Interior, Bureau of Land

Management: 81-88.

Monsen, S. B.; Shaw, N. L. 1983. Seeding antelope bitterbrush with

grass on south-central Idaho rangelands—a 39 year response. In:

bitterbrush and cliffrose symposium: proceedings. Gen. Tech.

Rep. INT-152. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 126–136.

Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps.

2004a. Restoring Western ranges and wildlands. Gen. Tech. Rep.

RMRS-GTR-136-vol-1. Fort Collins, CO: U.S. Department of

Agriculture, Forest Service, Rocky Mountain Research Station:

1–294 plus index.

Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps.

2004b. Restoring Western ranges and wildlands. Gen. Tech. Rep.

RMRS-GTR-136-vol-3. Fort Collins, CO: U.S. Department of

Agriculture, Forest Service, Rocky Mountain Research Station:

669–884 plus index.

Nelson, J. R.; Wilson, A. M.; Goebel, C. J. 1970. Factors influencing

broadcast seeding in bunchgrass range. Journal of Range Management. 23: 163–170.

Ocumpaugh, W. R.; Archer, S.; Stuth, J. W. 1996. Swithgrass

recruitment from broadcast seed vs. seed fed to cattle. Journal of

Range Management. 49: 368–371.

Olson, B. E.; Richards, J. H. 1988a. Tussock regrowth after grazing:

intercalary meristem and axillary bud activity of tillers of Agropyron desertorum. Oikos. 51: 374–382.

Olson, B. E.; Richards, J. H. 1988b. Annual replacement of the

tillers of Agropyron desertorum following grazing. Oecologia.

76: 1–6.

Owens, M. K.; Norton, B. E. 1990. Survival of juvenile basin big

sagebrush under different grazing regimes. Journal of Range

Management. 43: 132–135.

Pellant, M. 1990. The cheatgrass-wildfire cycle—are there any

solutions? In: McArthur, E. D.; Romney, E. M.; Smith, S. D.; and

Tueller, P. T., comps. Cheatgrass invasion, shrub die-off, and

other aspects of shrub biology and management symposium:

proceedings. Gen. Tech. Rep. INT-GTR-276. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 11–18.

Pendery, B. M.; Provenza, F. D. 1987. Interplanting crested wheatgrass with shrubs and alfalfa: effects of competition and preferential clipping. Journal of Range Management. 40: 514–520.

Pleasant, J. M. T.; Schlather, K. J. 1994. Incidence of weed seed in

cow (Bos sp.) manure and its importance as a weed source for

cropland. Weed Technology. 8: 304–310.

Plummer, A. P.; Hurd, R. M.; Pearse, C. K. 1943. How to reseed Utah

rangelands. Res. Pap. INT-1. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Forest and Range

Experiment Station. 22 p.

Pyke, D. A. 1990. Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential

expansion. Oecologia. 82: 537–543.

Pyke, D. A. 1994. Rangeland seedings and plantings: exotics or

natives. In: Edge, W. D.; Olson-Edge, S. L., eds. Proceedings—

symposium on sustaining rangeland ecosystems. Special Rep

953. Corvallis, OR: Oregon State University: 32–44.

Pyke, D. A.; Archer, S. 1991. Plant-plant interactions affecting plant

establishment and persistence on revegetated rangeland. Journal of Range Management. 44: 550–557.

Reynolds, T. 1980. Save some sage. Idaho Wildlife. 2: 10–13.

Richardson, B. Z.; Monsen, S. B.; Bowers, D. M. 1984. Interseeding

selected shrubs and herbs on mine disturbances in southeastern

Idaho. In: McArthur, E. D.; Welch, B. L., comps. Biology of

Artemisia and Chrysothamnus symposium: proceedings; 1984

July 9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 134–139.

Rittenhouse, L. R.; Sneva, F. A. 1976. Expressing the competitive

relationship between Wyoming big sagebrush and crested wheatgrass. Journal of Range Management. 29: 326–327.

Robertson, J. H.; Neal, D. L.; McAdams, K. R.; Tueller, P. T. 1970.

Changes in crested wheatgrass ranges under different grazing

treatments. Journal of Range Management. 23: 27–34.

Robertson, J. H.; Pearse, C. K. 1943. How to reseed Nevada rangelands. Res. Pap. INT-3. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 22 p.

Rogler, G. A.; Lorenz, R. J. 1983. Crested wheatgrass—early history

in the United States. Journal of Range Management. 36: 91–93.

Romo, J. T. 2005. Emergence and establishment of Agropyron

desertorum (Fisch.) (crested wheatgrass) seedlings in a sandhills

prairie of central Saskatchewan. Natural Areas Journal. 25:

26–35.

Romo, J. T.; Grilz, P. L.; Delanoy, L. 1994. Selective control of

crested wheatgrass (Agropyron cristatum [L.] Gaertn. and A.

desertorum Fisch.) in the Northern Great Plains. Natural Areas

Journal. 14: 308–309.

Roundy, B. A.; Shaw, N. L.; Booth, D. T. 1997. Using native seeds on

rangelands. In: Shaw, N. L.; Roundy, B. A., comps. Using seeds of

native species on rangelands: proceedings. Gen. Tech. Rep. INTGTR-372. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station: 1–8.

Salihi, D. O.; Norton, B. E. 1987. Survival of perennial grass

seedlings under intensive grazing in semi-arid rangelands. Journal of Applied Ecology. 24: 145–151.

Schuman, G. E.; Booth, D. T.; Cockrell, J. R. 1998. Cultural methods

for establishing Wyoming big sagebrush on mined lands. Journal

of Range Management. 51: 226–230.

Schuman, G. E.; Rauzi, F.; Booth, D. T. 1982. Production and

competition of crested wheatgrass-native grass mixtures.

Agronomy Journal. 74: 23–26.

Sharp, L. A. 1986. Crested wheatgrass: its values, problems and

myths. In: Johnson, K. L., ed. Crested wheatgrass: its values,

problems, and myths symposium: proceedings. Logan: Utah

State University: 3–6.

Shinderman, Matthew J.; Call, Christopher A. 2001. Establishment of forbs in cattle dungpats deposited in vegetation gaps
92 USDA Forest Service Proceedings RMRS-P-38. 2005

in a degraded sagebrush community. Ecological Restoration.

29: 99–106.

Simberloff, D. 2003. Confronting introduced species: a form of

xenophobia? Biological Invasions. 5: 179–192.

Smallwood, K. S.; Beyea, J.; Morrison, M. L. 1999. Using the best

scientific data for endangered species conservation. Environmental Management. 24: 421–435.

Sneva, Forrest A. 1970. Paraquat—effects of growing season applications. Journal of Range Management. 23: 451–452.

Springfield, H. W. 1963. Cattle gains and plant responses from

spring grazing on crested wheatgrass in northern New Mexico.

Production Res. Rep. 74. Washington, DC: U.S. Department of

Agriculture, Forest Service. 46 p.

Stevens, R. 1994. Interseeding and transplanting to enhance species composition. In: Monsen, S. B.; Kitchen, S. G., comps. Ecology

and management of annual rangelands: proceedings; 1992 May

18–21; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 300–306.

Stevens, R. 2004. Establishing plants by transplanting and

interseeding. In: Monsen, Stephen B.; Stevens, Richard; Shaw,

Nancy L., comps. Restoring Western ranges and wildlands. Gen.

Tech. Rep. RMRS-GTR-136-vol-3. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research

Station: 739–744.

Stevens, R.; Moden, W. L., Jr.; McKenzie, D. W. 1981. Interseeding

and transplanting shrubs and forbs into grass communities.

Rangelands. 3: 55–58.

Sturges, D. L. 1977. Soil water withdrawal and root characteristics

of big sagebrush. American Midland Naturalist. 98: 257–274.

Svejcar, T. 2003. Applying ecological principles to wildland weed

management. Weed Science. 51: 266–270.

Tausch, R. J.; Chambers, J. C.; Blank, R. R.; Novak, R. S. 1995.

Differential establishment of perennial grass and cheatgrass

following fire on an ungrazed sagebrush-juniper site. In: Roundy,

B. A.; McArthur, D. E.; Haley, J. S.; Mann, D. K., comps. Wildland

shrub and arid land restoration symposium: proceedings. Gen.

Tech. Rep. INT-GTR-315. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 252–257.

Trlica, M. J., Jr.; Cook, C. Wayne. 1972. Carbohydrate reserves of

crested wheatgrass and Russian wildrye as influenced by development and defoliation. Journal of Range Management. 25: 430–435.

U.S. Department of Agriculture, Natural Resources Conservation

Service (USDA NRCS). 2004. The PLANTS database, version 3.5.

Baton Rouge, LA: National Plant Data Center. (Online: http://

plants.usda.gov). [August 29, 2005].

U.S. Department of the Interior (USDI). 2001. National rangeland

inventory, monitoring, and evaluation report—FY 2001. Denver,

CO: U.S. Department of the Interior, Bureau of Land Management, National Science and Technology Center. 12 p.

U.S. Department of the Interior, Bureau of Land Management.

1981: Idaho Instruction Memorandum ID-81-312. Boise, ID: U.S.

Department of the Interior, Bureau of Land Management, Idaho

State Office.

U.S. Department of the Interior, Bureau of Land Management

(USDI BLM). 2000. The Great Basin Restoration Initiative:

healing the land. U.S. Department of the Interior, Bureau of Land

Management. (Online: http://www.fire.blm.gov/library.htm). [August 29, 2005].

Vallentine, J. F. 1989. Range development and improvements. 3d

ed. New York: Academic Press. 524 p.

Vallentine, J. F.; Cook, C. W.; Stoddart, L. A. 1963. Range seeding in

Utah. Extension Circular 307. Logan, UT: Utah State University.

20 p.

Van Epps, G. A.; McKell, C. M. 1978. Major criteria and procedures

for selecting and establishing range shrubs as rehabilitators of

disturbed lands. In: Hyder, D. N., ed. Proceedings of the First

International Rangeland Congress; 1978 August 14–18; Denver,

CO. Denver, CO: Society for Range Management: 352–354.

Van Epps, G. A.; McKell, C. M. 1980. Revegetation of disturbed sites

in the salt desert range of the Intermountain West. Land Rehabilitation Series 5. Logan, UT: Utah Agricultural Experiment

Station.

Walters, C. J. 1986. Adaptive management of renewable resources.

New York: McGraw-Hill.

Walters, C. J.; Holling, C. S. 1990. Large-scale management experiments and learning by doing. Ecology. 71: 2060–2068.

Wambolt, Carl L.; Harp, Aaron J.; Welch, Bruce L.; Shaw, Nancy;

Connelly, John W.; Reese, Kerry P.; Braun, Clait E.; Klebenow,

Donald A.; McArthur, E. Durant; Thompson, James G.; Torell, L.

Allen; Tanaka, John A. 2002. Conservation of greater sage-grouse

on public lands in the Western U.S.: implications of recovery and

management policies. PACWPL Policy Pap. SG-02-02. Caldwell,

ID: Policy Analysis Center for Western Public Lands. 41 p.

Welch, D. 1985. Studies in the grazing of heather moorland in

Northeast Scotland. Journal of Applied Ecology. 22: 461–472.

West, N. E.; Hassan, M. A. 1985. Recovery of sagebrush-grass

vegetation following wildfire. Journal of Range Management. 38:

131–134.

Whisenant, S. G. 1999. Repairing damaged wildlands: a processoriented, landscape-scale approach. New York: Cambridge University Press. 312 p.

Wiedemann, H. 2005. Revegetation equipment catalog. Revegetation Equipment Development Council; U.S. Department of Agriculture, Forest Service; and U.S. Department of the Interior,

Bureau of Land Management. (Online: http://reveg-catalog

.tamu.edu). [August 29, 2005].

Wilson, S. D.; Partel, M. 2003. Extirpation or coexistence? Management of a persistent introduced grass in a prairie restoration.

Restoration Ecology. 11: 410–416.

Young, J. A.; Evans, R. A. 1986. History of crested wheatgrass in the

Intermountain area. In: Johnson, K. L., ed. Crested wheatgrass:

its values, problems, and myths: proceedings. Logan, UT: Utah

State University: 21–25.

Young, J. A.; McKenzie, D. 1982. Rangeland drill. Rangelands.

4: 108–113.

Rose pusseytoes
USDA Forest Service Proceedings RMRS-P-38. 2005 93

Cindy R. Lysne (former USDI Bureau of LandManagement employee) is an

Environmental Specialist, Power Engineers, 1295 S. Eagle Flight Way, Boise,

ID 83709; e-mail: clysne@powereng.com

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Restoring Wyoming Big

Sagebrush

Cindy R. Lysne

Abstract—The widespread occurrence of big sagebrush can be

attributed to many adaptive features. Big sagebrush plays an

essential role in its communities by providing wildlife habitat,

modifying local environmental conditions, and facilitating the reestablishment of native herbs. Currently, however, many sagebrush

steppe communities are highly fragmented. As a result, restoring

big sagebrush is considered a priority in the conservation and

rehabilitation of sagebrush steppe ecosystems. Wyoming big sagebrush can often be difficult to establish, because many environmental factors act to restrict its emergence and persistence. On fire

rehabilitation projects in Idaho, Wyoming big sagebrush seed is

typically aerially broadcast over the soil surface. This method has

had some success; however, several alternative seeding treatments,

such as cultipacking, have resulted in the establishment and persistence of Wyoming big sagebrush. In addition, transplanting bareroot

and containerized stock may be useful for restoring shrub stands in

critical areas.

Keywords: Artemisia tridentata, revegetation, rehabilitation, seeding, shrub-steppe

In the Western United States, big sagebrush (A. tridentata

Nutt.) steppe communities dominate over 60 million ha

(Wambolt and Hoffman 2001) and provide essential habitat

and forage for many species (West 2000). Fragmentation of

sagebrush steppe communities has occurred through excessive livestock grazing, conversion to agricultural cropland,

invasion of exotic plants, and increasing frequency of large

fires (Anderson and Inouye 2001; Knick 1999; Knick and

Rotenberry 1997; Noss and others 1995). More than 350

species of plants and animals associated with sagebrush

ecosystems have been identified as species of conservation

concern due to declining habitats or populations (Wisdom

and others 2003).

Big sagebrush is important because of its wide distribution and the extent of disturbance within its range. It

provides both food and cover for sage-grouse (Centrocercus

urophasianus Bonaparte) year round (Connelly and others

2004). This paper presents a review of the literature on big

sagebrush taxonomy and characteristics, germination requirements, relevance in rehabilitation projects, and methods for improving its establishment in seedings and

transplantings. It will focus primarily on Wyoming big

sagebrush (A. t. Nutt. ssp. wyomingensis Beetle and Young).

Big Sagebrush Taxonomy and

Characteristics _________________

There are five subspecies of big sagebrush. These include

basin big sagebrush (A. t. Nutt. ssp. tridentata), Wyoming

big sagebrush, mountain big sagebrush (A. t. Nutt. ssp.

vaseyana [Rydb.] Beetle), xeric big sagebrush (A. t. Nutt. ssp.

xericensis Winward ex R. Rosentreter & R. Kelsey), and

subalpine big sagebrush (A. t. Nutt. ssp. spiciformis

[Osterhout] Kartesz and Gandhi) (Wambolt and Frisina

2002; West and Young 2000). The distribution of the subspecies is regulated by seasonal precipitation patterns, elevation, and soil conditions (McArthur 2000; McArthur and

others 1979, 1995; Monsen and Shaw 2000).

The dominance and ubiquitous occurrence of big sagebrush can be attributed to many factors. One factor is the

production of seasonally dimorphic leaves. Ephemeral leaves,

larger and often irregularly lobed, develop in spring and are

shed in summer following moisture stress (West and Young

2000). Persistent leaves are typically three-lobed, smaller,

develop in late spring, and remain on the shrubs through

winter (West and Young 2000). A second major factor contributing to the widespread occurrence of big sagebrush is an

efficient two-component root system (West and Young 2000).

Its fibrous root system captures water and nutrients near

the soil surface, permitting plants to take advantage of

summer precipitation (West and Young 2000). The taproot,

in turn, allows for utilization of water and nutrients deep

within the soil profile and below the principal rooting zone

of associated herbaceous species (West and Young 2000).

Several additional adaptive features influence the distribution and persistence of big sagebrush subspecies. These

include, but are not limited to, variable growth forms,

response to fire, the production of allelopathic substances in

roots and leaves, the ability to conduct photosynthesis at low

temperatures, temperature requirements for seed germination, seed dispersal strategies, seed size, and structure and

timing of seed maturation (Blaisdell and others 1982; Kelsey

1986; Meyer and Monsen 1992; Peterson 1995).

Big sagebrush plants are capable of producing seed in

their second year and will continue to produce some seed

annually, except during years of severe moisture stress

(Meyer and Monsen 1992). Plants flower in fall, following

the summer drought period, and the fruits (achenes) mature

from midfall to early winter (Meyer 2003). Achenes are small

(about 1 by 1.5 mm) and shiny with a deciduous pappus

(Meyer 2003). They are dispersed by gravity and wind, but

do not possess any special adaptations for wind dispersal

(Meyer 1994). Seeds may be blown by wind across crusty

snow surfaces and dispersed by animals and water (Tisdale

and Hironaka 1981; Young and Evans 1989a). Maximum
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Lysne Restoring Wyoming Big Sagebrush

dispersal distance of seeds can be up to 30 m; however, most

seeds (85 to 90 percent) fall within 1 m of the shrub canopy

(Meyer 1994; Young and Evans 1989a).

Big sagebrush seeds are surface or near-surface emerging

and are sensitive to microsite conditions (Meyer 1994).

Germination occurs in late winter to early spring, soon after

snowmelt, in areas where snow accumulates (Meyer and

Monsen 1992; West and Young 2000). A semi-gelatinous

pericarp and hypocotyl hairs aid in the adhesion of the

achene to the soil surface and permit the radicle to penetrate

the soil (Young and Martens 1991). The achene’s small size

reduces the surface area for moisture loss (Young and

Martens 1991). Achenes typically exhibit high seed viability

and germination capacity at maturity (Meyer 2003).

Wyoming Big Sagebrush

Wyoming big sagebrush is the most xeric subspecies of big

sagebrush. It generally occurs on shallow soil in areas

receiving 200 to 300 mm of annual precipitation (Cronquist

1994; Monsen and Shaw 2000). Wyoming big sagebrush

plants exhibit a ragged, irregular growth form, and most

plants grow to less than 1 meter in height. The main stem is

often branched into two or three twisted portions at or near

ground level (Winward and Tisdale 1977). Persistent leaves

are narrowly cuneate to cuneate with the margins curved

outward, and exhibit a strong, pungent odor when crushed

(McArthur and others 1979). The plants flower from late

July to September, and seed maturation occurs in October

and November (Monsen and Shaw 2000).

Germination and Establishment

Ecology _______________________

Many environmental factors act to reduce sagebrush establishment and persistence. Seed germination is substantially limited by water stress, and a principal cause of seed

mortality is early or prolonged drought (Meyer 1994). The

successful establishment of large cohorts of big sagebrush

shrubs can result from recruitment pulses that are associated with rare events of highly favorable precipitation

(Watson and others 1997; West and others 1979; Williams

and Hobbs 1989). High seed densities and synchronous

germination can result in intense competition between big

sagebrush seedlings. Intraspecific competition or self-thinning probably accounts for much of the initial mortality

(Meyer 1994). Competition between sagebrush plants within

a stand may also affect flowering and seed set, particularly

in dry years (Meyer 1994).

Competition with herbaceous species may also impact the

success of sagebrush seedings. However, reports on sagebrush seedling competitiveness with seeded wheatgrasses

are contradictory. During the time period when sagebrush

was being controlled on rangelands, managers often remarked on the ability of sagebrush to reestablish in perennial grass seedings (Meyer 2003). Conversely, researchers

have demonstrated that competition with introduced and

native grasses seeded before or with big sagebrush can

reduce Wyoming big sagebrush establishment (Blaisdell

1949; Fortier 2000; Schuman and others 1998). Similarly,

sagebrush seedings in areas with exotic annual grass competition have had little success (Meyer 2003). Competitive

effects are probably related to the inability of sagebrush

seedlings to compete for soil moisture during establishment

(Cook and Lewis 1963; Sturges 1977). Blaisdell (1949) found

higher grass yields on plots that were seeded with grass

prior to or 1 year after sagebrush, and that prior grass

establishment often prevented the establishment of sagebrush seedlings. However, when grasses were seeded 2 or 3

years following sagebrush seeding, grass yields were reduced and grass competition did not have an effect on

sagebrush (Blaisdell 1949).

Sagebrush seedlings have high first-year survival rates,

even through summer drought periods, on mine sites where

there is little competition (Meyer 1994). Schuman and others (1998) found that grass competition reduced sagebrush

seedling densities in a mined-land reclamation study using

direct-placed topsoil. They concluded that successful establishment of big sagebrush may require seeding big sagebrush without grasses or with very low grass seeding rates

(Schuman and others 2000).

Commercially available sagebrush seed is often not from

locally or regionally adapted seed sources. Nonadapted

seeds may respond differently to normal germination cues

and germination may occur at an inappropriate time, resulting in seeds that fail to germinate or persist (Monsen and

Meyer 1990). Using seedlots with the source or geographic

origin of the seed verified (Source Identified) and matched to

the site may be a key factor for achieving successful shrub

establishment (McArthur and others 1995; Meyer and

Monsen 1992). Commercially available seed often contains

a mixture of sagebrush subspecies (Dalzell 2004). Currently,

the Association of Seed Analysts does not provide guidelines

or testing methods for differentiating sagebrush subspecies

in purchased seed (AOSA 2003). Applying the big sagebrush

subspecies matched to the restoration site is essential because big sagebrush subspecies exhibit differences in seedling establishment traits (McArthur and others 1995),

growth rates (Welch and McArthur 1984), habitats

(Winward and Tisdale 1977), and moisture (Barker and

McKell 1983; Kolb and Sperry 1999), temperature (Harniss

and McDonough 1976), and germination requirements (Meyer

1994).

Seed bank studies of big sagebrush indicate seed banks

are transient, with very little seed carryover from one year

to the next (Meyer and Monsen 1992). Most of the big

sagebrush seeds produced in autumn are absent from the

soil seed bank by late spring of the following year (Young and

Evans 1989a). Wyoming big sagebrush seeds are, in general, short lived and do not survive fires (Young and Evans

1989a). Young and Evans (1989a) found that no mountain

big sagebrush or basin big sagebrush seedlings emerged

from germination tests of 1,000 soil surface samples taken

from a burned area. In contrast, however, some Wyoming big

sagebrush seeds applied with mulch cover on mined lands in

Wyoming remained viable in the seed bank for up to 4 years

(Schuman and others 1998).

Sagebrush seeds are highly viable with little or no dormancy at dispersal, but may have strong light requirements

for germination (Meyer 2003; Young and Evans 1989b). The

light requirement is removed through stratification (moist
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Restoring Wyoming Big Sagebrush Lysne

chilling), and most seeds are germinable by late winter or

early spring (Meyer 2003).

Use of Big Sagebrush in

Rehabilitation Projects ___________

Reestablishing big sagebrush is considered a priority in

the conservation and rehabilitation of sagebrush steppe

ecosystems (USDI BLM 2002a). In addition to providing

habitat for sage-grouse and other sagebrush obligate species, big sagebrush also plays an essential role in these

communities by directly modifying local environmental conditions, thus providing a more favorable environment for

seed germination and seedling survival (Schlesinger and

Pilmanis 1998). Shrubs also help to retain soil nitrogen,

increase organic matter, and create favorable environments

for microorganisms, resulting in fertile islands or patches

that develop over time (Cross and Schlesinger 1999; West

2000). By trapping blowing snow and moderating temperatures, big sagebrush facilitates the establishment of native

herbs, and their canopy protects native herbs from overutilization (West 2000). Wyoming big sagebrush also develops mycorrhizal fungi associations, which aid in nutrient

extraction and cycling (West 2000).

The establishment of big sagebrush is often difficult due to

poor seed quality (Harniss and McDonough 1976; Young and

Evans 1989a), low seedling vigor, exposure to unfavorable

seedbed conditions (McDonough and Harniss 1974; Meyer

and Monsen 1992), competition with herbaceous species

(Blaisdell 1949; Sturges 1977), and inadequate moisture

(Cook and Lewis 1963; Sturges 1977). Improved seed cleaning, handling, and purchasing requirements have made

higher quality seed easier to obtain (Meyer and Monsen

1992; Olson and others 2000). Also, seedbed conditions can

be manipulated to reduce competition and facilitate seed

germination (McArthur and others 1995; Welch and others

1992). Ultimately, however, environmental factors still play

a central role in determining the success of big sagebrush

restoration projects.

Seeding treatments can have a strong influence on the

emergence and survival of big sagebrush seedlings. On

Bureau of Land Management (BLM) fire-rehabilitation

projects in Idaho, Wyoming big sagebrush seed is typically

aerially broadcast over the soil surface by helicopter (USDI

2002b). Aerial broadcasting is often desirable over other

methods, because large areas can be seeded quickly and the

seed can be placed on the soil surface (Monsen 2000). This

seeding method has had some success (Monsen 2000); however, results from a recent study in southern Idaho indicate

that aerially seeding Wyoming big sagebrush had limited

effect on shrub establishment (Dalzell 2004). In this study,

seeding did not increase the density or cover of Wyoming big

sagebrush on seeded plots compared to adjacent unseeded

plots (Dalzell 2004). In fact, shrubs failed to establish on 23

of the 35 (66 percent) study sites sampled (Dalzell 2004).

Another key factor in the establishment and persistence of

sagebrush seedlings is the timing and amount of winter

snowfall. The recommended time for planting big sagebrush

is in fall, just before the first winter snowfall. This is the time

when big sagebrush would naturally be dispersing seed

onsite (Meyer 1994). Snow cover can facilitate the establishment of big sagebrush—particularly in areas with reliable,

long-term snow cover—by compacting or firming the soil

surface and assisting in keeping the seed in contact with the

soil. However on drier and warmer sites, winter snowfall

may be inadequate to facilitate these physical processes to

ensure successful big sagebrush emergence and establishment (Meyer 2003). Wyoming big sagebrush sites are typically windswept and relatively dry in both autumn and

winter (Meyer and Monsen 1992). These environmental

conditions are not favorable for sagebrush emergence or

establishment.

Increasing Shrub Establishment ___

There are several alternatives to aerial seeding that have

been shown to increase big sagebrush establishment and

persistence. For example, seeding equipment that compacts

the soil surface, such as cultipacking, chaining, and imprinting, can increase big sagebrush seedling establishment.

Monsen and Meyer (1990) obtained significantly greater

initial seedling emergence, compared to broadcasting, by

seeding with the Oyer compact row seeder. This device

compacts the soil and then presses the seed into the surface.

Intermediate seedling emergence results were achieved by

using the Brillion cultipacker seeder (Monsen and Meyer

1990). Using this device, the seed is broadcast over the

surface and pressed into the soil (Pyke 1994). The cultipacker

is a circular cylinder or set of wheels that are rolled over the

soil surface to place the seed in contact with the soil near the

soil surface (Pyke 1994).

The U.S. Department of the Interior, Bureau of Land

Management’s Lower Snake River District in Idaho achieved

successful sagebrush establishment using a seeder that

incorporates a fertilizer spreader, anchor chain or tire drags,

and a vine-roller cultipacker (Boltz 1994). This sagebrush

seeder covered the seed and firmed the soil surface on silt

loams, but it was less effective on gravelly and stony areas

(Boltz 1994).

Another option for establishing big sagebrush is to transplant bareroot or containerized stock. Stock that is 12 to 20

cm tall is transplanted in early spring (McArthur and others

1995). First year survival rates for transplanted stock are

often 80 percent or higher (Welch and others 1992). Seedlings are typically transplanted only in small, critical areas

due to the cost of using planting stock. Transplant stock can

be grown from small amounts of seed from specific areas

similar to the planting sites. Transplant stock is available

locally or regionally from private contracted nurseries and

from USDA Forest Service nurseries.

A similar method, the “mother plant” technique, combines

transplanting and natural seed dispersal. The mother plants

are planted as bareroot or containerized stock on key locations throughout the rehabilitation site. Within 3 to 5 years,

established mother plants mature, disperse seed, and provide an established seed source for unseeded areas (Welch

and others 1992). However, successful sagebrush establishment and subsequent dispersal also depends on the species

composition in the unseeded areas.

Big sagebrush is considered an obligate vesiculararbuscular mycorrhizal (VAM) plant (Wicklow-Howard
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Lysne Restoring Wyoming Big Sagebrush

1994). Arbuscular mycorrhizae can improve the ability of

plants to extract nutrients and water from the soil, thereby

improving the host species’ survival and growth on severely

disturbed lands (Wicklow-Howard 1994). In a greenhouse

study, Stahl and others (1998) found that sagebrush seedlings grown in topsoil with mycorrhizal inoculum exhibited

significantly greater tolerance to drought stress than nonmycorrhizal seedlings. Arid land disturbances such as fire,

mining, overgrazing, off-highway vehicle use, and cultivation significantly reduce the mycorrhizal inoculum potential

(MIP) of the soil (Wicklow-Howard 1989). Efforts to add

commercially available VAM fungal inoculum to the soil or

to use VAM-inoculated plants on disturbed areas have met

with limited success (Wicklow-Howard 1994).

To increase big sagebrush establishment, it is imperative

that alternative seeding methods are considered in lieu of

aerially seeding Wyoming big sagebrush, particularly if the

seed is not adequately covered. Although transplanting

bareroot and containerized stock is regarded as costly, this

expense may be acceptable when considering the current

failure to establish sagebrush using aerial seeding (Dalzell

2004). Because areas that have been depleted of sagebrush

for several years may lack the proper mycorrhizal fungi in

the soil, containerized stock should be inoculated with compatible fungi. Bareroot and containerized stock could be

transplanted in small, critical areas and in areas currently

dominated by introduced seeded grasses. Planting big sagebrush can also facilitate the restoration of highly palatable

selections of sagebrush, such as Gordon Creek Wyoming big

sagebrush, or local germplasms that are best suited for the

site conditions (Welch and others 1992).

Another alternative method is to transplant big sagebrush shrubs on areas where fertile islands existed prior to

burning, focusing on areas where native vegetation and

shrub skeletons remain. The transplanted shrubs will assist in the formation of “islands of fertility” (Cross and

Schlesinger 1999). These shrub islands will serve as habitat islands for animal species by providing shrub cover to

reduce the risk of predation (Longland and Price 1991),

providing temporary refuges to facilitate animal dispersal

and the maintenance of a metapopulation, a group of spatially separated subpopulations that are interlinked and

maintained by occasional dispersal (Longland and Bateman

2002). Areas that are positioned adjacent to the shrub

islands and have remaining native vegetation could be left

unseeded; thus reducing the mechanical disturbance of the

soil surface by some seeding equipment and reducing the

likelihood of exotic plant invasion, biological soil crust destruction, and subsequent wind erosion. In addition, the

islands would serve as a seed source for the replenishment

of native species with unavailable or limited commercial

seed supplies. The shrub islands would not only provide a

seed source for animals to harvest, consume, and disperse,

but also provide a refuge for seed dispersers (Longland and

Bateman 2002). In addition, these islands are sites with high

vesicular arbuscular mycorrhizal activity. Ultimately, the

development of fertile shrub islands would serve as inoculum focal points from which shrubs, VAM, and other species

could spread (Allen 1987).

As previously mentioned, establishment of big sagebrush

seedlings is impacted by competition. Seeding introduced

grasses such as crested wheatgrass (Agropyron cristatum)

and intermediate wheatgrass (Elymus hispidus) with big

sagebrush has prevented shrub seedlings from establishing

(Richardson and others 1986). Direct competition for available soil moisture and nutrients exists between seeded

grasses and big sagebrush because of similar root distributions and growth periods (Cook and Lewis 1963; Sturges

1977). In contrast, studies have shown that stands of native

bunchgrasses permitted big sagebrush recruitment (Booth

and others 2003; Frischknecht and Bleak 1957). Frischknecht

and Bleak (1957) reported that seeded stands of bluebunch

wheatgrass (Pseudoroegneria spicata) were more likely to

permit sagebrush seedling recruitment than seeded stands

of crested wheatgrass. In addition, Booth and others (2003)

found that the native perennial bunchgrass squirreltail

(Elymus elymoides) permitted big sagebrush recruitment

and also suppressed cheatgrass.

Seeds of several important native bunchgrasses are available. However, sources of other bunchgrasses adapted to the

Interior Western United States, such as Great Basin wildrye

(Leymus cinereus), bluebunch wheatgrass, Sandburg bluegrass (Poa sandbergii), bottlebrush squirreltail, and the

needlegrasses (Hesperostipa) are just beginning to be marketed. Additional research is needed to develop appropriate

seedbed preparation methods, planting techniques, and

equipment for the establishment of individual native species

and populations. Also, further research could focus on seeding big sagebrush in mixed seedings of native species and

investigating the ability of these seedings to permit sagebrush establishment and compete with invasive species.

Successful rehabilitation following wildland fire is essential to mitigate the effects exotic, invasive plants have on

ecosystems, decrease the frequency of large fires, provide

suitable wildlife habitat, and halt the conversion of diverse

sagebrush steppe communities to communities dominated

by exotic, invasive plants. To increase big sagebrush establishment, it is imperative that other seeding methods be

considered, utilized, monitored, and evaluated instead of the

commonly used aerial seeding technique. If current sagebrush restoration efforts do not result in more consistent

establishment and persistence of this important shrub,

large areas of sagebrush-steppe may be lost, and rehabilitation may no longer be a viable option (West 2000).

References _____________________

Allen, M. F. 1987. Ecology of vesicular-arbuscular mycorrhizae in

an arid ecosystem: use of natural processes promoting dispersal

and establishment. In: Sulvia, D. M.; Hung, L. L.; Graham, J. H.,

eds. 7th North American conference on mycorrhizae: proceedings;

1987 May 3–8; Gainesville, FL: University of Florida: 133–135.

Anderson, J. A.; Inouye, R. S. 2001. Landscape-scale changes in

plant species abundance and biodiversity of a sagebrush steppe

over 45 years. Ecological Monographs. 71: 531–556.

Association of Official Seed Analysts (AOSA). 2003. Artemisia seed

testing working group workshop. Unpublished document on file

at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, ID. 7 p.

Barker, Jerry R.; McKell, Cyrus M. 1983. Habitat differences

between basin and Wyoming big sagebrush in contiguous populations. Journal of Range Management. 36: 450–454.

Blaisdell, J. P. 1949. Competition between sagebrush seedlings and

reseeded grasses. Ecology. 30: 512–519.
USDA Forest Service Proceedings RMRS-P-38. 2005 97

Restoring Wyoming Big Sagebrush Lysne

Blaisdell, J. P.; Murray, R. B.; McArthur, E. D. 1982. Managing

Intermountain rangelands—sagebrush-grass ranges. Gen. Tech.

Rep. INT-134. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 41 p.

Boltz, M. 1994. Factors influencing postfire sagebrush regeneration

in south-central Idaho. In: Monsen, S. B.; Kitchen, S. G., comps.

Ecology, management, and restoration of intermountain annual

rangelands: proceedings; 1992 May 18–22; Boise, ID. Gen. Tech.

Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 281–290.

Booth, M. S.; Caldwell, M. M.; Stark, J. M. 2003. Overlapping

resource use in three Great Basin species: implications for community invasibility and vegetation dynamics. Journal of Ecology.

91: 36–48.

Connelly, J. W.; Knick, S. T.; Schroeder, M. A.; Stiver, S. J. 2004.

Conservation assessment of greater sage-grouse and sagebrush

habitats. Report. Cheyenne, WY: Western Association of Fish

and Wildlife Agencies. 610 p.

Cook, C. W.; Lewis, C. E. 1963. Competition between big sagebrush

and seeded grasses on foothill ranges in Utah. Journal of Range

Management. 16: 245–250.

Cronquist, A. 1994. Vol. 5. Asterales. In: Cronquist, A.; Holmgren,

A. H.; Holmgren, N. H.; Reveal, J. L.; Holmgren, P. K., eds.

Intermountain flora: vascular plants of the Intermountain West,

U.S.A. Bronx: The New York Botanical Garden. 496 p.

Cross, A. F.; Schlesinger, W. H. 1999. Plant regulation of soil

nutrient distribution in the northern Chihuahuan Desert. Plant

Ecology. 145: 11–25.

Dalzell, C. R. 2004. Post-fire establishment of vegetation communities following reseeding on southern Idaho’s Snake River Plain.

Boise, ID: Boise State University. 112 p. Thesis.

Fortier, M. I. 2000. Wyoming big sagebrush establishment on mined

lands: seeding rates and grass competition. Laramie: University

of Wyoming. 66 p. Thesis.

Frischknecht, N. C.; Bleak, A. T. 1957. Encroachment of big sagebrush on seeded range in northeastern Nevada. Journal of Range

Management. 10: 165–170.

Harniss, R. O.; McDonough, W. T. 1976. Yearly variation in germination of three subspecies of big sagebrush. Journal of Range

Management. 29: 167–168.

Kelsey, R. G. 1986. Emergence, seedling growth and crude turpenoid

concentration in a sagebrush garden. In: McArthur, E. D.; Welch,

B. L., comps. Symposium on the biology of Artemisia and

Chrysothamnus: proceedings; 1984 July 9–13; Provo, UT. Gen.

Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 358–365.

Kolb, K. J.; Sperry, J. S. 1999. Differences in drought adaptation

between subspecies of sagebrush (Artemisia tridentata). Ecology.

80: 2373–2384.

Knick, S. T. 1999. Requiem for a sagebrush ecosystem? Northwest

Science. 73: 53–57.

Knick, S. T.; Rotenberry, J. T. 1997. Landscape characteristics of

disturbed shrubsteppe habitats in southwestern Idaho (U.S.A.).

Landscape Ecology. 12: 287–297.

Longland, W. S.; Bateman, S. L. 2002. The ecological value of shrub

islands on disturbed sagebrush rangelands. Journal of Range

Management. 55: 571–575.

Longland, W. S.; Price, M. V. 1991. Direct observations of owls and

heteromyid rodents: can predation risk explain microhabitat

use? Ecology. 72: 2261–2273.

McArthur, E. D. 2000. Sagebrush systematics and distribution. In:

Entwistle, P. G.; DeBolt, A. M.; Kaltenecker, J. H.; Steenhof, K.,

comps. Sagebrush steppe ecosystems symposium: proceedings;

1999 June 23–25; Boise, ID. BLM/ID/PT-001001+1150. Boise,

ID: U.S. Department of the Interior, Bureau of Land Management: 9–14.

McArthur, E. D.; Blauer, A. C.; Plummer, A. P.; Stevens, R. 1979.

Characteristics and hybridization of important Intermountain

shrubs. III. Sunflower family. Res. Pap. INT-220. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Forest and Range Experiment Station. 82 p.

McArthur, E. D.; Stevens, R.; Monsen, S. B. 1995. Adaptation and

success of big sagebrush and rubber rabbitbrush on disturbed

sites. In: Schuman, G. E.; Vance, G. F., eds. 12th annual national

meeting of the American Society for Surface Mining and Reclamation: proceedings; 1995 June 3–8; Gillette, WY. Lexington,

KY: American Society for Surface Mining and Reclamation:

811–823.

McDonough, W. T.; Harniss, R. O. 1974. Effects of temperature on

germination in three subspecies of big sagebrush. Journal of

Range Management. 27: 204–205.

Meyer, S. E. 1994. Germination and establishment ecology of big

sagebrush: implications for community restoration. In: Monsen,

S. B.; Kitchen, S. G., comps. Ecology, management, and restoration of Intermountain annual rangelands: proceedings; 1992

May 18–22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 244–251.

Meyer, S. E. 2003. Artemisia L. In: Bonner, F. T.; Nisley, R. G., eds.

Woody plant seed manual. Washington, DC: U.S. Department of

Agriculture, Forest Service. Online: http://ntsl.fs.fed.us/wpsm/

[February 15, 2005].

Meyer, S. E.; Monsen, S. B. 1992. Big sagebrush germination

patterns: subspecies and population differences. Journal of Range

Management. 45: 87–93.

Monsen, S. B. 2000. Establishment of big sagebrush (Artemisia

tridentata) in semiarid environments. In: Entwistle, P. G.; DeBolt,

A. M.; Kaltenecker, J. H.; Steenhof, K., comps. Sagebrush steppe

ecosystems symposium: proceedings; 1999 June 23–25; Boise, ID.

BLM/ID/PT-001001+1150. Boise, ID: U.S. Department of the

Interior, Bureau of Land Management: 81–86.

Monsen, S. B.; Meyer, S. E. 1990. Seeding equipment effects on

establishment of big sagebrush on mine disturbances. In: Fifth

Billings symposium on disturbed land rehabilitation. Vol. I.; 1990

March 25–29; Billings, MT. Reclamation Research Unit Publ.

9003. Bozeman: Montana State University: 192–199.

Monsen, S. B.; Shaw, N. L. 2000. Big sagebrush (Artemisia tridentata)

communities—ecology, importance and restoration potential. In:

Schuman, G. E.; Richmond, T. C.; Neuman, D. R., eds. Sagebrush

establishment on mined lands: ecology and research: proceedings; 2000 March 20–24; Billings, MT. Reclamation Research

Unit Publ. 00-01. Bozeman, MT: Montana State University:

255–270.

Noss, R. F.; LaRoe, E. T., III; Scott, J. M. 1995. Endangered ecosystems of the United States: a preliminary assessment of loss and

degradation. Biol. Rep. 28. Washington, DC: U.S. Department of

the Interior, National Biological Service. 81 p.

Olson, R. A.; Gores, J. K.; Booth, D. T.; Schuman, G. E. 2000.

Suitability of shrub establishment on Wyoming mined lands

reclaimed for wildlife habitat. Western North American Naturalist. 60: 77–92.

Peterson, J. G. 1995. Ecological implications of sagebrush manipulation. W-101-R-2. Helena: Montana Department of Fish, Wildlife, and Parks, Wildlife Management Division. 49 p.

Pyke, D. A. 1994. Rangeland seedings and plantings: exotics or

natives. In: Edge, W. D.; Olson-Edge, S. L., eds. Symposium on

sustaining rangeland ecosystems: proceedings; 1994 August 29–

31; La Grande, OR. Special Rep. 953. Corvallis: Oregon State

University: 32–44.

Richardson, B. Z.; Monsen, S. B.; Bowers, D. M. 1986. Interseeding

selected shrubs and herbs on mine disturbances in southeastern

Idaho. In: McArthur, E. D.; Welch, B. L., eds. Symposium on the

biology of Artemisia and Chrysothamnus: proceedings; 1984 July

9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 134–139.

Schlesinger, W. H.; Pilmanis, A. M. 1998. Plant-soil interactions in

deserts. Biogeochemistry. 42: 169–187.

Schuman, G. E.; Booth, D. T.; Cockrell, J. R. 1998. Cultural methods

for establishing Wyoming big sagebrush on mined lands. Journal

of Range Management. 51: 223–230.

Schuman, G. E.; Booth, D. T.; Olson, R. A. 2000. Enhancing Wyoming big sagebrush establishment with cultural practices. In:

Schuman, G. E.; Richmond, T. C.; Neuman, D. R., eds. Sagebrush

establishment on mined lands: ecology and research: proceedings;

2000 March 20–24; Billings, MT. Reclamation Research Unit

Publ. 00-01. Bozeman, MT: Montana State University: 32–42.
98 USDA Forest Service Proceedings RMRS-P-38. 2005

Lysne Restoring Wyoming Big Sagebrush

Stahl, P. D.; Schuman, G. E.; Frost, S. M.; Williams, S. E. 1998.

Arbuscular mycorrhizae and water stress tolerance of Wyoming

big sagebrush seedlings. Soil Science Society of America Journal.

62: 1309–1313.

Sturges, D. L. 1977. Soil water withdrawal and root characteristics

of big sagebrush. American Midland Naturalist. 98: 257–274.

Tisdale, E. W.; Hironaka, M. 1981. The sagebrush-grass region: a

review of the ecological literature. Bull. 33. Moscow: University

of Idaho, Forestry, Wildlife, and Range Experimental Station.

31 p.

U.S. Department of the Interior, Bureau of Land Management

(USDI BLM). 2002a. Management considerations for sagebrush

(Artemisia) in the Western United States: a selective summary of

current information about the ecology and biology of woody North

American sagebrush taxa. Washington, DC: U.S. Department of

the Interior, Bureau of Land Management. 73 p.

U.S. Department of the Interior, Bureau of Land Management

(USDI BLM). 2002b. Unpublished fire records and maps. On file

at: U.S. Department of the Interior, Bureau of Land Management, Idaho State Office, Boise, ID. Various paging.

Wambolt, C. L.; Frisina, M. R. 2002. Montana sagebrush guide.

Helena, MT: Montana Department of Fish, Wildlife and Parks.

71 p.

Wambolt, C. L.; Hoffman, G. R. 2001. Mountain big sagebrush age

distribution and relationships on the northern Yellowstone winter range. In: McArthur, E. D.; Fairbanks, D. J., comps. Shrubland

ecosystem genetics and biodiversity: proceedings; 2000 June

13–15; Provo, UT. RMRS-P-21. Ogden, UT: U.S. Department of

the Interior, Forest Service, Rocky Mountain Research Station:

330–335.

Watson, I. W.; Westoby, M.; Holm, A. McR. 1997. Continuous and

episodic components of demographic change in arid zone shrubs:

models of two Eremophila species from Western Australia compared with published data on other species. Journal of Ecology.

85: 833–846.

Welch, B. L.; McArthur, E. D. 1984. Growth rate of big sagebrush as

influenced by accessions, sites, subspecies, and years. In McArthur,

E. D.; Welch, B. L., comps. Symposium on the biology of Artemisia

and Chrysothamnus: proceedings; 1984 July 9–13; Provo, UT.

Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of

Agriculture, Forest Service, Intermountain Research Station:

342–346.

Welch, B. L.; Nelson, E. D.; Young, S. A.; Sands, A. R.; Wagstaff, F. J.;

Nelson, D. L. 1992. ‘Gordon Creek’—a superior, tested germplasm

of Wyoming big sagebrush. Res. Pap. INT-461. Ogden, UT: Forest

Service, Intermountain Research Station. 7 p.

West, N. E. 2000. Synecology and disturbance regimes of sagebrush

steppe ecosystems. In: Entwistle, P. G.; DeBolt, A. M.; Kaltenecker,

J. H.; Steenhof, K., comps. Sagebrush steppe ecosystems symposium: proceedings; 1999 June 23–25; Boise, ID. BLM/ID/PT001001+1150. Boise, ID: U.S. Department of the Interior, Bureau

of Land Management: 15–26.

West, N. E.; Rea, K. H.; Harniss, R. O. 1979. Plant demographic

studies in sagebrush-grass communities of southeastern Idaho.

Ecology. 60: 376–388.

West, N. E.; Young, J. A. 2000. Intermountain valleys and lower

mountain slopes. In: Barbour, M. G.; Billings, W. D., eds. North

American terrestrial vegetation. 2d ed. Cambridge, UK: Cambridge University Press: 255–284.

Wicklow-Howard, M. C. 1989. The occurrence of vesicular-arbuscular

mycorrhizae in burned areas of the Snake River Birds of Prey

Area, Idaho. Mycotaxon. 34: 253–257.

Wicklow-Howard, M. C. 1994. Mycorrhizal ecology of shrub-steppe

habitat. In: Monsen, S. B.; Kitchen, S. G., comps. Ecology, management, and restoration of Intermountain annual rangelands:

proceedings; 1992 May 18–22; Boise, ID. Gen. Tech. Rep. INTGTR-313. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station: 207–210.

Williams, K.; Hobbs, R. J. 1989. Control of shrub establishment by

springtime soil water availability in an annual grassland.

Oecologia. 81: 62–66.

Winward, A. H.; Tisdale, E. W. 1977. Taxonomy of the Artemisia

tridentata complex in Idaho. Bull. 19. Moscow, ID: University of

Idaho, Idaho Forestry, Wildlife and Range Experimental Station.

15 p.

Wisdom, M. J.; Rowland, M. M.; Suring, L. H.; Schueck, L.; Wolff

Meinke, C.; Wales, B. C.; Knick, S. T. 2003. Procedures for

regional assessment of habitats for species of conservation concern in the sagebrush ecosystem. March 2003 Report, Version 1.

La Grande, OR: U.S. Department of Agriculture, Forest Service,

Pacific Northwest Research Station. 151 p.

Young, J. A.; Evans, R. A. 1989a. Dispersal and germination of

big sagebrush (Artemisia tridentata) seeds. Weed Science. 37:

201–206.

Young, J. A.; Evans, R. A. 1989b. Reciprocal common garden studies

of the germination of seeds of big sagebrush (Artemisia tridentata).

Weed Science. 37: 319–324.

Young, J. A.; Martens, E. 1991. Importance of hypocotyl hairs in

germination of Artemisia seeds. Journal of Range Management.

44: 438–442.

Engelmann Aster
USDA Forest Service Proceedings RMRS-P-38. 2005 99

Nancy L. Shaw is Research Botanist and Ann DeBolt is Botanist, U.S.

Department of Agriculture, Forest Service, Rocky Mountain Research Station, Boise, ID 83702; e-mail: nshaw@fs.fed.us and adebolt@fs.fed.us. Roger

Rosentreter is State Botanist, U.S. Department of the Interior, Bureau of

Land Management, Idaho State Office, Boise, ID 83709; e-mail:

roger_rosentreter@blm.gov

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Reseeding Big Sagebrush:

Techniques and Issues

Nancy L. Shaw

Ann M. DeBolt

Roger Rosentreter

Abstract—Reestablishing big sagebrush on rangelands now dominated by native perennial grasses, introduced perennial grasses, or

exotic annual grasses, particularly cheatgrass (Bromus tectorum),

serves to stabilize soil, improve moisture availability and nutrient

recyling, increase biological diversity, and foster community stability

and resiliency. A first priority in reseeding is identifying the subspecies of big sagebrush native to the site and procuring adapted,

high-quality seed of that subspecies from a similar site. Seed should

be planted on firm seedbeds and pressed into the soil to provide good

seed-to-soil contact. Competition from invasive species and other

seeded species must be minimized by site preparation practices and

use of appropriate seeding strategies and equipment. Precipitation

is often a major factor in determining seeding success on drier sites.

Postseeding monitoring and careful management are necessary to

maintain stands and provide feedback for improving future seeding

efforts. Additional research and technological developments are

required to better estimate and maintain big sagebrush seed quality, provide required seedbed conditions, and reestablish mixed

seedings of big sagebrush and associated natives.

The sagebrush (Artemisia spp.) biome encompasses approximately 63 million ha of the Western United States, but

little of this area has remained unaltered since Euro-American settlement. Vast tracts have been lost to agriculture,

urbanization, and other human activities. Of the remaining

area, it has been estimated that 50 to 60 percent has been

converted to nonnative annual grasslands or contains exotic

annual grasses in the understory (West 2000). Even though

more than 70 percent of the sagebrush-steppe is publicly

owned, less than 3 percent is protected in National Parks or

other Federal reserves (Knick and others 2003). The increasingly rapid and widespread degradation, fragmentation,

and, in some areas, near total loss of sagebrush has resulted

in its being rated one of the most imperiled ecosystems in

North America (Noss and Peters 1995). Some have advocated that a regional objective of no net loss of sagebrush be

adopted to prevent further declines in biodiversity (Paige

and Ritter 1999; West 2000).

More than 20 sagebrush species and subspecies occur

within the sagebrush biome (Goodrich, this proceedings;

Rosentreter, this proceedings). It is spatially complex, with

variable soils, topography, parent materials, climates, landscape patterns, and disturbance histories (Miller and

Eddleman 2001). Sagebrush populations display a strong

alliance to certain habitats, with morphological specializations and adaptations evolving along environmental gradients (Schultz 1986). Prior to Euro-American settlement, fire

regimes were equally complex across this region and contributed significantly to landscape heterogeneity. With the

shift in fire regimes that has occurred over the past 100

years, largely due to the spread of nonnative plant introductions into voids created by postsettlement livestock grazing,

this once complex landscape has become increasingly homogeneous. All of these factors contribute to the enormous

difficulty that land managers experience in attempts to

restore native plant communities where natural recruitment is often limited by a lack of propagules, drought, a

competitive exotic understory, disruption of hydrologic functioning, and changes in soil structure and biota as a result of

past disturbances.

Early seeding success with introduced grasses contributed to their widespread use for soil stabilization and to type

conversion of sagebrush landscapes for increased forage

production; the latter is an objective that dominated our use

of this biome for much of the twentieth century (Holechek

and others 1998). From the 1930s into the 1970s, an estimated 2 to 6 million ha of sagebrush habitat was burned,

sprayed, or treated mechanically to reduce sagebrush (Braun

1998; Vale 1974). Due to health concerns, use of 2,4-D and

2,4,5-T was curtailed in the 1980s, but other treatments

continued through that decade. The total acreage impacted

is unknown, but it has been estimated to exceed 20 to 25

percent of the total remaining sagebrush-dominated landscape (Braun 1998).

Concern for big game habitat loss increased as these large

treatments continued. Monocultures of any one species do

not constitute healthy or desirable rangelands (Stevens and

others 1981), and generalist animals such as grasshoppers

(Orthoptera), deer mice (Peromyscus spp.), horned larks

(Eremophila alpestris), and introduced chukars (Alectoris

chukar) occur in seedings of introduced grasses (Maser and

others 1984). Public concern led to the increased use of

browse species in wildlife habitat treatments. Blaisdell

(1972) reported that research in shrub ecology had contributed to the identification of about 75 shrubs as promising

for improving big game habitat (see Plummer and others

1968). Four shrubs, big sagebrush (A. tridentata), fourwing
100 USDA Forest Service Proceedings RMRS-P-38. 2005

Shaw, DeBolt, and Rosentreter Reseeding Big Sagebrush: Techniques and Issues

saltbush (Atriplex canescens), antelope bitterbrush (Purshia

tridentata), and rubber rabbitbrush (Chrysothamnus

nauseosus), were considered primary species to be promoted. The use of big sagegrush and other shrubs in rangeland rehabilitation treatments on Federal lands has gradually increased since the mid to late 1980s. More recently, the

decline of sage-grouse and other sagebrush obligate species

has given additional impetus to restoration of big sagebrush

habitats. Although an additional two decades have passed,

we still have much to learn about restoring this landscape

dominant and its associated species to disturbed lands. Here

we provide a review of recent big sagebrush restoration

literature and recommendations for reestablishment and

management of this species and its communities.

Natural Regeneration of Big

Sagebrush _____________________

Most Artemisia species, subspecies, and ecotypes are easily killed by fire. They do not resprout and therefore must

regenerate from seed. Of the five subspecies in the big

sagebrush complex (table 1), only subalpine big sagebrush

(A. t. spp. spiciformis) can resprout from root crowns or lower

stem bases after being top-killed by burning (Winward

1985). Fire passing through a Wyoming big sagebrush (A. t.

ssp. wyomingensis) plant will usually kill it (Britton and

Clark 1985).

A big sagebrush plant may produce 500,000 seeds in a

typical year (Welch and others 1990), but annual production

varies greatly (Young and Evans 1975). Big sagebrush seeds

are small and exceedingly light; those of basin big sagebrush

(A. tridentata ssp. tridentata) are generally lighter (0.018 g/

100 seeds) than those of mountain and Wyoming big sagebrush (0.025 g/100 seeds) (Meyer and others 1987). Big

sagebrush seeds are dispersed primarily by gravity. Maximum dispersal distances are only about 30 m from the

parent plant; 85 to 90 percent of all seeds fall within 1 m of

the edge of the mother plant (Wagstaff and Welch 1990;

Young and Evans 1989). Consequently, long-distance dispersal by wind is ineffective in recolonizing large burns or

other disturbances (Meyer 1994).

Artemisia seeds rarely survive in the soil for more than a

year (Caldwell 1978; McDonough and Harniss 1974; Walton

and others 1986). However, some seed may carry over if

buried and not exposed to light (Hassan and West 1986;

Meyer 1994; Meyer and Monsen 1990; Richardson and

others 1986). Schuman and others (1998) found that Wyoming big sagebrush seed survived up to 4 years when applied

with mulch on mine spoils in Wyoming.

Rapid reestablishment of most big sagebrush subspecies

is more likely on sandy or gravelly soils that are well suited

for supporting the species. Big sagebrush returns more

slowly on fine-textured soils that have a greater potential for

production of herbaceous species (Blaisdell and others1982;

Hironaka and others1983). Xeric big sagebrush (A. t. ssp.

xericensis) is the only taxon in the big sagebrush complex

adapted to fine-textured clay soils.

Natural postfire reestablishment of big sagebrush has not

been widely documented. During years of low precipitation,

few Wyoming big sagebrush plants may establish, and it

may take many years before recolonization takes place.

Even under favorable conditions, site recovery may require

60 to 100 years. On dry Wyoming big sagebrush sites,

several years may pass before conditions favoring establishment of new seedlings occur (Clifton 1981; Lowe-Dalzell and

others 2003; Wambolt and Payne 1986; West and Hassan

1985; Young and Evans 1978). Because of these factors, big

sagebrush must be artificially reseeded on sites where seed

sources have been lost.

Postfire, Pretreatment Site

Evaluation _____________________

Prior to treatment, it is imperative that a site evaluation

be conducted to assure that artificial restoration measures

are needed and that natural recovery will not occur within

an acceptable time frame (fig. 1). If recovery is not anticipated without seeding, the preburn density of exotic annuals

and the postburn seed bank of these species must be estimated to determine the potential for restoring the site, the

overall objectives must be established, and the approach for

accomplishing the seeding or planting must be selected.

The characteristics of various ecological sites and their

distribution within a given management area should be

thoroughly understood. Site characteristics vary according

to the potential natural community, species present, soil

depth and texture, effective precipitation, erosion potential,

elevation, aspect, and other factors (National Research Council 1994). Burned big sagebrush sites that receive less than

250 mm of annual precipitation, particularly where the

understory is cheatgrass (Bromus tectorum) dominated,

have a low probability of regenerating naturally and providing preburn cover and structure in a reasonable length of

time. This is due to inadequate seed supplies on surviving

plants or in the soil seed bank and the combined effects of low

and erratic precipitation and herbaceous competition from

exotic annuals (Boltz 1994). It is these lands that are in the

most urgent need of restoration, but are the most risky to

treat. For such sites, a greater investment of time and money

will be required, and priorities, objectives, and resource

availability are particularly important considerations.

Adapted species and subspecies must be planted using

procedures that remove competition and create suitable

seedbeds. Developing measures to remove or diminish competition is difficult, but failure to implement all proven site

improvement measures significantly reduces the chance of

success (Monsen and McArthur 1995; Stevens and Monsen

2004). For example, herbicide application or the use of

container or bareroot transplant stock may be necessary

Table 1—Big sagebrush complex.

 Common name Scientific name

Subalpine big sagebrush Artemisia tridentata ssp. spiciformis

Basin big sagebrush A. t. ssp. tridentata

Mountain big sagebrush A. t. ssp. vaseyana

Few-flowered mountain A. t. ssp. vaseyana f. pauciflora

 big sagebrush

Wyoming big sagebrush A. t. ssp. wyomingensis

Xeric big sagebrush A. t. ssp. xericensis
USDA Forest Service Proceedings RMRS-P-38. 2005 101

Reseeding Big Sagebrush: Techniques and Issues Shaw, DeBolt, and Rosentreter

Figure 1—Site evaluation form.

Site Evaluation Form

Name _________________________________________________________ Date ___________________________________________

Fire name _____________________________________________________ Fire No. ________________________________________

Date fire started _______________________________________________ Date fire controlled ______________________________

District or Forest ______________________________________________ Elevation _______________________________________

Acres burned and ownership: ______________ total acres ____________ public _____________ State ____________ private

Preburn vegetation types and estimated acres of each: __________________________________________________________

____________________________________________________________________________________________________________

____________________________________________________________________________________________________________

____________________________________________________________________________________________________________

Preburn ecological site(s) and estimated acres of each:

Range/ecological condition: ___________________________________________________________________________________

Precipitation zone(s) _________________________________________________________________________________________

Fire severity: ___________________ acres low _____________________ acres moderate ________________________ acres high

Soil series/name _________________________________Soil depth according to survey _______________________________

Soil description and texture _____________________________________________________________________________________

______________________________________________________________________________________________________________

Current land use(s) ____________________________________________________________________________________________

Grazing season of use/type of system (specific dates) ______________________________________________________________

______________________________________________________________________________________________________________

Acres/AUM _____________________________________________No. pastures ___________________________________________

Range condition ________________________________________________________________________________________________

_______________________________________________________________________________________________________________

Is use pattern map available? __________________________________________ (if so, please attach copy)

Key wildlife seasonal habitat?___________________________________________________________________________________

Noxious weeds? Species and occupied acres (attach map) __________________________________________________________

______________________________________________________________________________________________________________

Resource objectives: ____________________________________________________________________________________________

______________________________________________________________________________________________________________

Fencing (describe preburn and identify additional needs) __________________________________________________________

______________________________________________________________________________________________________________

______________________________________________________________________________________________________________

Recommended treatments (include for noxious weeds) ____________________________________________________________

______________________________________________________________________________________________________________

______________________________________________________________________________________________________________

Seed mix(es), rates, method of application, PLS cost:

Drill seeded—Seed mix 1 Drill seeded—Seed mix 2 Aerial seeded—Seed mix 3

species/subspecies/rate/cost species/subspecies/rate/cost species/subspecies/rate/cost

Totals: Totals: Totals:
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Shaw, DeBolt, and Rosentreter Reseeding Big Sagebrush: Techniques and Issues

under such conditions. Evaluation forms such as figure 1

may be used to facilitate the decisionmaking process.

Stevens (2002) recommends the following four steps be

followed in selecting taxa for a seeding: (1) develop a list of

species and ecotypes that would occur on the proposed

planting site; (2) from this list, determine which species have

a significant amount of high quality seed available for

planting; (3) of these available species, determine those that

are compatible as young developing plants and that will

ensure ecological development of a desired plant community; and (4) evaluate the final species list to determine if

project objectives can be achieved or whether the initial

objectives require revision.

To successfully reestablish big sagebrush, the subspecies

present preburn must be determined and utilized in the

seeding effort. Remnant plants may be identified using the

descriptions provided by Goodrich (this proceedings) and

Rosentreter (this proceedings). Subspecies and populations

of big sagebrush have evolved in distinct environments.

Common garden studies have revealed differences in adaptive characteristics such as drought or frost tolerance

(McArthur and Welch 1982; Meyer and Monsen 1990; Welch

and others 1992); movement of populations to different

climatic or edaphic conditions is not advised (Mahalovich

and McArthur 2004; Monsen 2000). Specific ecotypes may be

especially important on droughty sites or mineral soils.

Matching treatment site characteristics, such as soil type

and elevation, with the seed source is critical, but this has

frustrated land managers and in some cases been impossible

during large fire years when seed is in high demand and

production low. Consequently, it is all the more imperative

that the correct big sagebrush subspecies be used.

Seed Biology and Technology _____

Seed Harvesting and Conditioning

Big sagebrush flowers in summer and is wind pollinated.

Large numbers of tiny flowers develop on spikes, racemes, or

panicles, with individual plants producing hundreds of thousands of achenes (Welch and others 1990) in years with

favorable weather conditions. Seeds (achenes) ripen in late

fall and are usually dispersed within a few weeks of reaching

maturity, depending upon weather conditions and subspecies. Seed of mountain big sagebrush generally ripens earlier than seed of basin big sagebrush or Wyoming big sagebrush, and considerable variability in date of ripening will

be found within individual plants and populations. In addition, seed production varies widely from year to year based

on weather conditions, herbivory, seed predation, and other

factors (Wagstaff and Welch 1991; Young and others 1989).

Because of these factors, seeds should be checked carefully

with a hand lens before harvesting to ensure that adequate

quantities of sound seeds are present to justify harvesting.

Seed harvested too early will be immature and not viable.

Delaying the issuing of permits until seed has matured has

been suggested as one means of discouraging early harvest

on public lands (AOSA 2003). Seed harvested too late, after

dispersal of most sound seeds, will include large quantities

of poorly developed seeds and fruit and flower parts. An

additional complication is the frequent occurrence of basin

big sagebrush and Wyoming big sagebrush mosaics with the

basin big sagebrush growing in deeper soils or along roadways or along riparian areas. It is essential that care be

taken to collect only the target subspecies.

Seed is hand harvested by beating or stripping the inflorescences into seed hoppers, boxes, bags, or other containers.

Harvesting should be done when the humidity is low because

the fruits separate more easily from the inflorescences when

dry. Average moisture content of fully ripened seeds of big

sagebrush has not been examined carefully. Moisture content of seed and debris is often high when seed is harvested

from plants that are covered with snow or frost in late fall.

Seed is initially dried to a moisture content of 18 to 20

percent before cleaning to protect seed viability and to

reduce the volume of material to be conditioned (AOSA

2003). Appropriate drying techniques and rates and their

effects on seed quality require further investigation; rapid

seed deterioration with improper handling is considered a

major obstacle to maintaining big sagebrush seed viability

beyond the first year (AOSA 2003).

Purity of harvested seed lots is extremely low due to the

presence of inflorescence branches, leaves, bracts, poorly

developed fruits, and other debris. Seed is cleaned using a

barley debearder or hammermill to break up the inflorescences and other debris. Screening and fanning then removes trashy material. Big sagebrush seed is generally

cleaned to 10 to 15 percent purity (Stevens and others 1996),

but purities of 80 percent or more can be obtained by further

cleaning with an air screen separator. Cleaning to a purity

of 35 percent has been suggested as a means of reducing bulk

and cost for shipping and storage, increasing the consistency

and accuracy of seed sampling and seed quality testing,

improving the regulation of seed moisture content in storage, and facilitating seed metering through seeding devices

(AOSA 2003; Welch 1995).

For current seeding practices, purity of 10 to 12 percent

and viability of 85 to 95 percent is recommended by Meyer

(2005). Lambert (2005) recommended 14 percent purity and

80 percent viability as minimum standards for USDI Bureau of Land Management purchases of big sagebrush. If all

large debris is removed, seed cleaned to this level can be

seeded through broadcast seeders, rangeland drills, Hansen

browse seeders, and other standard seeding devices (Shaw

and Monsen 1990). Maximum allowable moisture content

can also be listed in purchase specifications.

Seed Storage and Longevity

Following late fall harvest, big sagebrush seed must be

dried, cleaned, and tested before it can be sold. Consequently, a considerable amount of newly harvested seed is

not marketed before the late fall seeding period, but must be

held over in storage for at least 1 year. Storing seed at

moisture contents of 6 to 8 percent and a temperature below

10 °

C (Meyer 2005) may lengthen viability to as much as 5

years. Storage of seed under adverse conditions, even for

short periods, can negatively affect seed quality and translate into a rapid decline in viability and vigor. Thus it is

advised that seed be tested for viability before purchasing or

seeding in order for prices and seeding rates to be determined accurately.
USDA Forest Service Proceedings RMRS-P-38. 2005 103

Reseeding Big Sagebrush: Techniques and Issues Shaw, DeBolt, and Rosentreter

Seed Testing

Testing of big sagebrush seed lots is plagued by a number

of problems stemming from the small seed size, low purity

levels, and the large size of marketed seed lots. Additional

research is urgently needed to provide guidelines that will

aid users in maintaining and more accurately measuring

seed quality. Such guidelines would reduce problems related to marketing, handling, and seeding big sagebrush.

Problems in assessing purity arise from sampling procedures at the warehouse and in the laboratory. Seed lots are

often large and heterogeneous. Big sagebrush seed does not

flow and samples must be drawn from bags by hand, a

technique that introduces more variability than use of a

trier or probe; thus purity of samples drawn from a single

seedlot for submission to the seed laboratory can differ

substantially. Initial samples drawn from large seed lots

may be too large to submit to the laboratory and will require

further subsampling, thus introducing additional variability. When the submitted sample reaches the seed laboratory, the working sample is obtained by dividing the sample

by hand, a less reliable technique than use of mechanical

dividers used for seeds that flow readily. The AOSA (2003)

suggested variability in samples might be reduced by limiting seed lot size and by marketing big sagebrush seed at

purities in the 35 percent range. Seeds might then be

classified as flowable, and triers and mechanical dividers

could be used for sampling, thus improving sampling

consistency.

Purity testing for big sagebrush is slow and costly. In

addition to sampling problems, the small seed size, large

amounts of debris present, and problems related to selection

of pure seed increase the time required for completion of

tests and reduce the accuracy of results (AOSA 2003). Again,

increasing purity levels to the 35 percent range would

reduce the bulk of seed and debris that must be examined,

remove many of the seeds that are small, nonviable, or

poorly developed; speed the testing process considerably;

and reduce the variability of results.

Procedures for testing viability of members of the genus

Artemisia are provided by AOSA (2000). Results can be

obtained quickly, depending primarily on the laboratory’s

backlog. AOSA germination tests are available for big sagebrush, black sagebrush (A. nova), and Louisiana sagebrush

(A. ludoviciana) (AOSA 2000). The germination test for

Louisiana sagebrush requires 14 days. Germination tests

for big sagebrush and black sagebrush require 21 days;

dormant seedlots require a 14-day prechill. Meyer (2005)

recommends testing nongerminating seeds for viability as

not all dormant seed will respond to the short prechill.

Sampling problems and identification of pure seed reduces

consistency of results.

Seed shipped for purity and germination or viability

testing may be packaged in paper bags or containers. Seeds

shipped for moisture testing should be packed in plastic

bags to maintain the water content at the same level as the

seed lot. Use of the International Seed Testing Association

rule for testing moisture content (drying at 105 oC for 16

hours) should be specified for determining seed water content (AOSA 2003).

Germination and Seedling Establishment

The level of seed dormancy and the light requirement for

germination vary widely among big sagebrush seed sources

and tend to decline with afterripening in dry storage or with

a moist prechill. Compared with seed from lower elevations,

seed from high elevations generally requires a longer field or

laboratory stratification to release dormancy and reduce the

light requirement (Meyer 2005). Seeds that have lost their

dormancy germinate rapidly under favorable moisture conditions. Likewise, germination under snow occurs slowly at

high elevations, while only a short period of snow cover may

facilitate rapid germination of low elevation seed sources

(Meyer and Monsen 1990; Young and others 1990). Germination is highly erratic on dry and windy sites where snow

cover is less reliable; seeds from such locations may be capable

of germinating rapidly, even at low temperatures, when

moisture conditions are favorable (Meyer 1994; Meyer and

Monsen 1992). Favorable microsites for germination are

provided if seeds are placed at or near the soil surface and

pressed into a firm, but not compacted seedbed. This provides

the exposure to light required for germination and good seed

to soil contact for improving water uptake. Imbibed seed

produces a layer of mucilaginous material that improves

adhesion to the soil. In addition, the hairs that develop on

emerging hypocotyls also aid in water uptake and soil

contact (Walton and others 1986; Young and Martens 1991).

Although seedlings sometimes establish in large numbers

due to high seed production, favorable weather, and appropriate microsite conditions, most seedlings are generally

lost to late frosts or drought, disease, inter- or intraspecific

competition, herbivory, or other factors. Seeding methods or

techniques that provide favorable microsites or improve

snow or water catchment, as well as the presence of mature

shrubs that can function as nurse plants improve establishment (Monsen and others 1992).

Seeding Considerations

Artificial seeding should only be pursued when the objective is to reestablish shrubs more rapidly than would occur

by natural recovery (Shaw and Monsen 1990). However,

some circumstances such as severe site conditions or degradation, complex ownership patterns, absence of crucial habitat, small size of a treatment area relative to others in need

of seeding, budgetary constraints, or some combination of

these factors may render seeding impractical. Decisions are

best made following a field site evaluation (fig. 1).

Site preparation and seeding techniques that reduce early

competition from both annual grasses and seeded species

and provide suitable microsite conditions for germination

and early growth (Meyer 1994) must be selected. On degraded sites, extensive site preparation and weed control

will be necessary when dense stands of annuals are present

preburn or are expected to develop. Not only will preparation

of a firm seedbed be required, but also control of exotic

annual grasses, primarily cheatgrass (Bromus tectorum L.)

and medusahead wildrye (Taeniatherum caput-medusae L.),

will be necessary.

Recommended big sagebrush seeding rates range from

0.11 to 0.22 kg per ha pure live seed (PLS) (Meyer 1994;
104 USDA Forest Service Proceedings RMRS-P-38. 2005

Shaw, DeBolt, and Rosentreter Reseeding Big Sagebrush: Techniques and Issues

Monsen 2000); increases of up to 50 percent are recommended for broadcast seeding (Welch and others 1986).

Seeding rates should be calculated on a PLS basis using

results of a recent viability or germination test (Meyer 1994).

Because of the extreme variation in microsites, moisture

availability, and temperature conditions encountered by

seeds and seedlings as well as a lack of research and

monitoring data, more definitive recommendations are not

possible.

Due to its small size, big sagebrush seed is usually mixed

with a carrier, thus cleaning to a higher purity than the

commercial lot average of 10 to 20 percent PLS may not be

cost effective or necessary (Meyer 1994). However, newer

drills may be able to seed lots with higher purities at

acceptable rates, thus reducing the bulk of seed lots required

for individual projects; additional research is required to

examine this possibility.

Seeding in late fall or early winter is recommended, as this

is when big sagebrush naturally disperses and soil surfaces

are more likely to be moist and firm; it also permits the

stratification required to attain vigorous germination if

adequate moisture is present. Spring seeding should be

avoided (Meyer 1994).

Seeding Techniques

Big sagebrush seed should be planted on a firm seedbed

with only a light covering of soil. Smooth, compacted seedbeds do not offer good seed to soil contact. Rough seedbeds

may slough and bury seeds too deeply. Big sagebrush can be

seeded with other species to increase diversity; however,

seeding requirements and the relative seedling growth rates

of each species must be considered when writing a seeding

plan. Due to their earlier maturity, seeded grasses establishing with big sagebrush have an initial advantage and suppress big sagebrush seedlings. Dense stands of seeded grasses

may entirely suppress big sagebrush seedlings or prevent

big sagebrush reestablishment for an indefinite period

(Blaisdell 1949). Due to these concerns and as a general rule

of thumb, grass should be seeded at low rates (3.6 to 5.4 kg/ha)

if big sagebrush establishment is one of the treatment

objectives.

Seeding has frequently been accomplished by aerial broadcasting to keep seeds near the soil surface and to plant large,

rough areas rapidly. Ground broadcasting using mechanical

seeders or hand seeding is also commonly used. Coverage of

broadcast seed using chains, harrows, rails, or other implements is recommended (Stevens and Monsen 2005). Lysne

(this proceedings) found that in southern Idaho, big sagebrush seeded aerially and not covered failed to establish on

23 of 35 fire rehabilitation projects examined, while natural

regeneration occurred on about one-fourth of the projects.

Elevation on her sites ranged from 810 to 1,640 m and

annual precipitation from 150 to 305 mm. Overall big sagebrush density did not differ between seeded and nonseeded

portions of these projects. As alternative treatments for this

area, Lysne (this proceedings) and Lysne and Pellant (2004)

recommended seeding methods that create a firm seedbed

and press the seed into the soil, thus at least some sagebrush

seed is placed near the soil surface. They suggested use of

equipment such as the Oyer compact row seeder (Monsen

and Meyer 1990), Brillion cultipacker seeder (Monsen and

Meyer 1990), Jarbidge big sagebrush seeder (Boltz 1994), or

land imprinter (Monsen 1988; Haferkamp and others 1987).

Big sagebrush can be seeded through drills if seed is

dropped on or near the soil surface and covered lightly by

press wheels (Lambert, this proceedings) or by pulling an

implement such as a cultipacker behind the drill. For drill

seedings, Richardson and others (1986) recommended that

big sagebrush be planted in separate rows from grass and

forb species. Otherwise, due to their rapid development,

grasses and forbs will compete directly with the slower

growing shrub seedlings (Richardson and others 1986) for

water and other resources.

New drills equipped with multiple seedboxes, seeding

depth regulators for each drop, and surface compaction

attachments offer greater flexibility for planting different

species in separate rows (Boltz 1994; Wiedemann 2005).

Addition of a fluffy seed box to the rangeland drill has also

increased options for seeding sagebrush. The Utah Division

of Wildlife Resources has recently begun purchasing big

sagebrush seed cleaned to 30 percent purity with a minimum of 80 percent germination giving a PLS of 24 percent.

Seed harvested and purchased in late autumn is placed in

cold storage by December or January and seeded the next

autumn. Use of Truax drills or rangeland drills with a fluffy

seed box permits use of this seed without addition of a carrier

and has reduced problems associated with seed testing

(Vernon 2005).

Interseeding is another approach to establishing big sagebrush and other slow growing shrubs. This technique involves disking, plowing, or spraying to remove strips of

established vegetation such as introduced grass seedings or

invasive species. Shrubs and other species that are slow to

establish are then seeded using a Hanson seeder or thimble

seeder. Interseeders have been constructed to accomplish

mechanical removal of existing vegetation and seeding in

one pass (Stevens and others 1981; Wiedemann 2005). The

Hansen seeder has also been used to drop big sagebrush seed

and other shrubs ahead of the wheels of a tractor or the

tracks of a caterpillar. The wheels or tracks create a firm

seedbed and press the seed into the soil. Seed of a variety of

species can be planted using this method as seeds are placed

over a range of depths. Grasses and larger seeded forbs are

seeded through the drill.

Hydroseeding is generally impractical for large rangeland

rehabilitation or restoration projects. This technique is

labor intensive and expensive. In addition, many sites are

difficult to access with hydroseeding equipment or water

trucks. Moreover, good seed to soil contact is generally not

provided by incorporating the seed into the mulch on dry

sites.

Establishing early seral native grasses and shrubs such as

rubber rabbitbrush on burned or otherwise disturbed sites

may reduce annual weed density and permit establishment

of big sagebrush seeded at a later time. Summer precipitation occurring following the senescence of native herbaceous

species may enhance big sagebrush establishment. Meyer

and Monsen (1990) found evidence that previously establishing rubber rabbitbrush may have facilitated colonization of big sagebrush on a mined site in Nevada. Naturally

reestablishing rubber rabbitbrush (Chrysothamnus

nauseosus) invaded mine spoils at the Beacon Pit Mine that

had not been covered with topsoil (Meyer 1994). Ten years
USDA Forest Service Proceedings RMRS-P-38. 2005 105

Reseeding Big Sagebrush: Techniques and Issues Shaw, DeBolt, and Rosentreter

following disturbance, more than 60 percent of the rubber

rabbitbrush plants were in the adult size class, while about

70 percent of the big sagebrush plants were less than 30 cm

tall. This suggests that initial establishment of the rubber

rabbitbrush may have ameliorated site conditions and facilitated big sagebrush establishment. Planting disturbed areas to rabbitbrush to enhance big sagebrush establishment

is feasible and ecologically practical (Monsen 2000). Rabbitbrush can be established either by seeding or transplanting.

It is capable of establishing and spreading to sites occupied

by cheatgrass. Rabbitbrush plants aid in trapping snow,

moderating temperature extremes, and accumulating litter,

all beneficial for the big sagebrush seedling environment.

Nursery Stock

To enhance shrub and forb communities in the Intermountain West, container-grown and bareroot stock have

proven effective for increasing diversity (Stevens 1994).

Because of the expense, the usefulness of transplanting

seedlings may be limited to small, critical areas, if high

shrub densities are required. Bareroot planting stock should

be from 12 to 20 cm tall and over-wintered in a nonheated

nursery bed or lathhouse (Welch and others 1986). Long and

Trimmer (2004) reported that at the Lone Peak Conservation Nursery, 1-0 big sagebrush seedlings are root pruned at

a depth of 30 cm in August. Following February or March

lifting, seedlings are graded to specifications of a minimum

of 15 cm shoot length and 4 mm collar diameter.

Transplanting should generally be done in spring when

soil moisture and the chance of storms are greatest, temperatures are low, and frost heaving has ceased (Deitschman

1974; Stevens 1981; Welch and others 1986).

While transplanting is more expensive than direct seeding, success is often much greater and more evident. The

more edaphically or climatically severe the site, the greater

is the need for transplanting (Stevens 1981).

Everett (1980) found that planting containerized shrubs,

including mountain big sagebrush, in late winter (February)

was a viable method of establishing vegetation in the harsh

environment of roadside cutbanks in the Sierra Nevada

foothills (Everett 1980). Initial establishment was highly

dependent on quality of planting stock and weather

conditions. In every instance where small or insufficiently

hardened planting stock was used, survival rates declined

drastically (Everett 1980). Tiedemann and others (1976)

reported on the importance of adequate transplant size for

survival of shrubs in eastern Washington.

Stevens and others (1981) found that bareroot stock of

many native shrubs, including mountain big sagebrush,

could be planted successfully with a hand-fed tree planter in

scalps 0.6 m on a side and 0.2 m deep made in heavy grass

sod. The transplanting rate varied between 10 and 18 plants

per minute depending on plant species, size and condition of

plants, soil type, and surface conditions. Shrubs were planted

at spacings of 0.9 to 2.4 m. Establishment was greater for

bareroot stock than for container-grown stock. Bareroot

stock with roots 15 to 30 cm long and tops at least 8 cm long

were most successful.

Seeding may also be accomplished by employing the

“mother plant” concept—big sagebrush transplants are

planted on a 15- by 15-foot grid. These “mother plants”

mature and produce enough seed in 3 to 5 years to supply the

seed for natural dispersal throughout site if native grasses

are reestablished to reduce weedy competition (Welch and

others 1986). Mechanical or chemical treatments may be

necessary to reduce competition in strips or scalps at the

time of planting.

Postseeding Management

On Bureau of Land Management lands, seedings are

typically excluded from livestock grazing for two growing

seasons to allow establishment. Stevens (1994) found that

grazing pressure must be removed from newly planted or

seeded areas for a minimum of 2 years. However, others

have suggested that longer periods of rest from livestock and

wildlife use are probably needed, particularly when attempting to reestablish shrubs such as big sagebrush(Fisser

1981). Protected plants develop more rapidly and natural

spread from seed is hastened with longer protection. Shaw

and Monsen (1990) stated that 2 to 3 years of protection from

livestock grazing reduces seedling losses from grazing or

trampling. Richardson and others (1986) compared grazed

and ungrazed treatments 7 years after seeding mountain big

sagebrush on a mid-elevation site in southeastern Idaho.

They found significantly lower big sagebrush densities in

the grazed treatment, an effect they attributed, in part, to

trampling.

Stevens and others (1996) reported that 20 to 40 percent

of big sagebrush transplants in seed orchards produce seed

by the second year, and 80 to 90 percent by the third and

fourth years. When seed orchards are established from seed,

10 percent of the plants can be expected to produce seed by

the second year, 30 to 50 percent by the third year, and 80 to

90 percent by the fourth to fifth year. On wildland sites,

longer periods may be required, particularly under drought

conditions.

Plummer and others (1968) stated that planted areas must

not be overgrazed. Until seeded stands have become established and suppressed natives have had an opportunity to

recover and become reproductive, livestock grazing should

be light if permitted at all. After range restoration has been

accomplished, grazing should be conservative. Either fencing or management of animals is often necessary to give

young plants adequate time to attain mature stature.

Protection fences are installed to protect a new seeding

from grazing and trampling during the establishment

period and to manage established seedings (Interagency

Burned Area Emergency Stabilization and Rehabilitation

Handbook 2002).

Management is also required to restrict off-highway vehicles and other human activities that may impact seeded

areas. Early weed control may be required to reduce the risk

of seeding failure and spread or recovery of invasive species.

Monitoring Seeding Establishment

Appropriate monitoring protocols (for example, Elzinga

and others 1998) should be selected to measure the extent to

which seedings are successful in meeting management goals

and to provide for adaptive management. Establishment

and monitoring of unseeded controls and grazing exclusion

plots on seeded and unseeded areas permit evaluation of
106 USDA Forest Service Proceedings RMRS-P-38. 2005

Shaw, DeBolt, and Rosentreter Reseeding Big Sagebrush: Techniques and Issues

seeded species establishment, natural recovery, and the

impacts of livestock grazing (Lysne, this proceedings; Lysne

and Pellant 2004). Regular monitoring during the first few

years postseeding, records of seed lot history (origin, quality,

storage conditions) and seeding techniques applied, a site

description, site conditions during the time of seeding, and

weather records are invaluable for evaluating monitoring

results, establishing the time required for individual species

to reach reproductive status, determining readiness for

grazing, and suggesting modifications for future seeding or

planting efforts.

Conclusions and

Recommendations ______________

Big sagebrush and associated native species can be seeded

on sites where seed sources have been lost and natural

recovery is not expected to occur. Careful planning; acquisition of adapted, high quality seed; selection of seeding

techniques appropriate for individual species as well as the

combination of species selected; and careful post-seeding

monitoring and management are all required to maximize

seeding success, permit recovery of remnant native species,

and maintain established seedings. Additional research and

improved technology are required to solve problems related

to maintaining and evaluating seed quality, providing seeding techniques that place big sagebrush seed in appropriate

microsites for germination, and protect them from herbaceous competition, whether from co-seeded or invasive species. Low and erratic precipitation on drier big sagebrush

sites often limits seeding success.

A major obstacle to the increased use of big sagebrush is

the problem of obtaining adequate seed supplies of the

required subspecies from adapted sites when needed. The

difficulty of identifying the subspecies in individual seed

lots, seed lots containing mixtures of subspecies, limited

shelf life of big sagebrush seed, and inadequate cold storage

space contribute to this problem. Efforts to delineate seed

transfer zones for Artemisiataxa (Mahalovich and McArthur

2004) and a recent research initiative to select and manage

wildland stands of Wyoming big sagebrush for seed production seek to address this issue. In situ conservation and

protection of selected big sagebrush stands in areas where

reseeding is likely to be required could increase the availability and quality of adapted seed.

Current literature and knowledge on seeding and establishment of big sagebrush subspecies have been summarized by Stevens and others (2004), McArthur and Stevens

(2004), Lysne and Pellant (2004), and others. Seasonal

habitat requirements for sage-grouse and recommendations

for restoring degraded sage-grouse habitats are described in

a number of publications including Connelly and Braun

(1997), Connelly and others (2000), Crawford and others

(2004), and Wambolt and others (2002). The SAGEMAP

Project Web site (http://sagemap.wr.usgs.gov/sage_grouse.htm)

provides a library of texts and databases for all aspects of

shrub steppe and sage-grouse management in the Intermountain West.

References _____________________

Association of Official Seed Analysts (AOSA). 2000. Tetrazolium

testing handbook. Contribution 29. Lincoln, NE: Association of

Official Seed Analysts. nonpaginated.

Association of Official Seed Analysts (AOSA). 2003. AOSA Artemisia seed testing working group. Unpublished document on file at:

U.S. Department of Agriculture, Forest Service, Rocky Mountain

Research Station, Boise, ID. nonpaginated.

Blaisdell, J. P. 1949. Competition between sagebrush seedlings and

reseeded grasses. Ecology. 30: 512–519.

Blaisdell, J. P.; Murray, R. B.; McArthur, E. D. 1982. Managing

Intermountain rangelands—sagebrush-grass ranges. Gen. Tech.

Rep. INT-134. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 40 p.

Blaisdell, James P. 1972. Needs and opportunities for shrub research in the Western United States. In: McKell, Cyrus M.;

Blaisdell, James P.; Goodin, Joe R., tech. eds. Wildland shrubs—

their biology and utilization; July 1971; Logan, UT. Gen. Tech.

Rep. INT-1. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Forest and Range and Experiment Station: 409–413.

Boltz, M. 1994. Factors influencing postfire sagebrush regeneration

in south-central Idaho. In: Monsen, S. B.; Kitchen, S. G., comps.

Proceedings: ecology and management of annual rangelands;

1992 May 18–22; Boise, ID. Gen. Tech. Rep. INT-GTR-313.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 281–290.

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. 67: 134–144.

Britton, C. M.; Clark, R. G. 1985. Effects of fire on sagebrush and

bitterbrush. In: Sanders, K.; Durham, J., eds. Rangeland fire

effects: a symposium. Boise, ID: U.S. Department of the Interior,

Bureau of Land Management, Idaho State Office: 22–26.

Caldwell, M. M. 1978. Physiology of sagebrush. In: The sagebrush

ecosystem: a symposium. Logan, UT: Utah State University: 74–85.

Clifton, Nancy A. 1981. Response to prescribed fire in a Wyoming big

sagebrush/bluebunch wheatgrass habitat type. Moscow: University of Idaho. 39 p. Thesis.

Connelly, J. W.; Braun, C. E. 1997. Long-term changes in sagegrouse Centrocercus urophasianus populations in Western North

America. Wildlife Biology. 3: 229–234.

Connelly, J. W.; Schroeder, M. A.; Sands, A. R.; Braun, C. E. 2000.

Guidelines for management of sage grouse populations and

habitat. Wildlife Society Bulletin. 28: 967–985.

Crawford, John A.; Olson, Rich A.; West, Neil E.; Mosley, Jeffrey C.;

Schroeder, Michael A.; Whitson, Tom D.; Miller, Richard F.;

Gregg, Michael A.; Boyd, Chad S. 2004. Ecology and management

of sage-grouse and sage-grouse habitat. Journal of Range Management. 57: 2–19.

Deitschman, G. H. 1974. Artemisia L. sagebrush. In: Shopmeyer,

C. E., tech. coord. Seeds of woody plants in the United States.

Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 235–237.

Elzinga, Caryl L.; Salzer, Daniel W.; Willoughby, John W.; 1998.

Measuring and monitoring plant populations. BLM Tech. Ref.

1730-1. Denver, CO: U.S. Department of the Interior, Bureau of

Land Management, National Applied Resource Sciences Center.

477 p.

Everett, R. L. 1980. Use of containerized shrubs for revegetating

arid roadcuts. Reclamation Review. 3: 33–40.

Fisser, H. G. 1981. Shrub establishment, dominance, and ecology

on the juniper and sagebrush-grass types in Wyoming. In: Stelter,

L. H.; DePuit, E. J.; Mikol, S. A., comps. Proceedings: shrub

establishment on disturbed arid and semiarid lands; 1980 December 2–3; Laramie, WY. Cheyenne: Wyoming Fish and Game

Department: 23–28.

Haferkamp, M. R.; Ganskopp, D. C.; Miller, R. F.; Sneva, F. A. 1987.

Drilling versus imprinting for establishing crested wheatgrass in

the sagebrush-bunchgrass steppe. Journal of Range Management. 40: 524–530.
USDA Forest Service Proceedings RMRS-P-38. 2005 107

Reseeding Big Sagebrush: Techniques and Issues Shaw, DeBolt, and Rosentreter

Hassan, M. A.; West, N. E. 1986. Dynamics of soil seed pools in burned

and unburned sagebrush semi-deserts. Ecology. 67: 269–272.

Hironaka, M.; Fosberg, M. A.; Winward, A. H. 1983. Sagebrushgrass habitat types of southern Idaho. Bull. 35. Moscow: University

of Idaho, Forest, Wildlife, and Range Experiment Station. 44 p.

Holechek, Jerry L.; Pieper, Rex D.; Herbel, Carlton H. 1998. Range

management principles and practice. 4th ed. Upper Saddle River,

NJ: Prentice-Hall, Inc. 624 p.

Interagency burned area emergency stabilization and rehabilitation handbook. 2002. Washington, DC: U.S. Department of the

Interior, Bureau of Land Management, and U.S. Department of

Agriculture, Forest Service. [Online]. Available: http://

fire.r9.fws.gov/ifcc/esr/handbook/4PolicyGuidance.htm [7 May

2004].

Jones, T. A.; Johnson, D. A. 1998. Integrating genetic concepts into

planning rangeland seedings. Journal of Range Management. 51:

594–606.

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.

Lambert, Scott. 2005. List of native grasses, forbs, and shrubs of the

Great Basin region. Boise, ID: U.S. Department of the Interior,

Bureau of Land Management. nonpaginated.

Long, Brandon; Trimmer, Edie. 2004. Propagation protocol for

bareroot sagebrush (Artemisia spp.). Native Plants Journal. 5:

149–151.

Lowe-Dalzell, C.; Eldridge, D.; Pellant, M.; Wicklow-Howard, M.

2003. Post-fire establishment of sagebrush on Idaho rangelands

in the Western United States. In: Allsopp, N.; Palmer, A. R.;

Milton, S. J.; Kirkman, K. P.; Kerley, G. I. H.; Hurt, C. R.; Brown,

C. J., eds. Proceedings of the VIIth International Rangeland

Congress; 2003 July 26–August 1; Durban, South Africa. Irene,

South Africa: Document Transfer Technologies: 370–372.

Lysne, Cindy; Pellant, Mike. 2004. Establishment of aerially seeded

big sagebrush following southern Idaho wildfires. Tech. Bull.

2004-01. Boise, ID: U.S. Department of the Interior, Bureau of

Land Management, Idaho State Office. 14 p.

Mahalovich, Mary F.; McArthur, E. Durant. 2004. Sagebrush (Artemisia spp.) seed and plant transfer guidelines. Native Plants

Journal. 5: 141–148.

Maser, C.; Thomas, J. W.; Anderson, R. G. 1984. Wildlife habitats in

managed rangelands—the Great Basin of southeastern Oregon.

The relationship of terrestrial vertebrates to plant communities

and structural conditions. Gen. Tech. Rep. PNW-172. Portland,

OR: U.S. Department of Agriculture, Forest Service, Pacific

Northwest Forest and Range Experiment Station. 54 p.

McArthur, E. D.; Stevens, Richard. 2004. Composite shrubs. In:

Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps.

Restoring Western ranges and wildlands. Gen. Tech. Rep. RMRSGTR-136-Vol-2. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 493–538.

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.

McDonough, W. T.; Harniss, R. O. 1974. Effects of temperature on

germination in three subspecies of big sagebrush. Journal of

Range Management. 27: 204–205.

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; 1992 May 18–22; Boise, ID. Gen. Tech.

Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Research Station: 244–251.

Meyer, S. E.; McArthur, E. D.; Monsen, S. B. 1987. Infraspecific

variation in germination patterns of rangeland shrubs and its

relationship to seeding success. In: Frasier, G. W.; Evans, R., eds.

Seed and seedbed ecology of rangeland plants: proceedings of

symposium; 1987 April 21–23; Tucson, AZ: U.S. Department of

Agriculture, Agricultural Research Service: 82–92.

Meyer, S. E.; Monsen, S. B. 1990. Seed-source differences in initial

establishment for big sagebrush and rubber rabbitbrush. In:

McArthur, E. D.; Romney, E. M.; Smith, S. D.; Tueller, P. T., eds.

Symposium, cheatgrass invasion, shrub die-off and other aspects

of shrub biology and management; 1989 April 5–7; Las Vegas, NV.

Gen. Tech. Rep. INT-276. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 200–208.

Meyer, S. E.; Monsen, S. B. 1992. Big sagebrush germination

patterns: subspecies and population differences. Journal of Range

Management. 45: 87–93.

Meyer, S. E.; Monsen, S. B.; McArthur, E. D. 1990. Germination

response of Artemisia tridentata to light and chill: patterns of

between-population variation. Botanical Gazette. 152: 176–183.

Meyer, Susan E. 2005. Artemisia L. In: Bonner, F. T.; Nisley, R. G.,

eds. Woody plant seed manual. Agric. Handb. Washington, DC:

U.S. Department of Agriculture, Forest Service. [Online]. Available: http://ntsl/fs.fed.us/wpsm [2005, January 14].

Miller, Richard F.; Eddleman, Lee L. 2001. Spatial and temporal

changes of sage-grouse habitat in the sagebrush biome. Corvallis:

Oregon State University, Agricultural Experiment Station. 35 p.

Monsen, S. B. 1988. Comparison of different planting equipment

and seeding rates to establish Wyoming big sagebrush. Progress

Report. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station; Boise, ID: U.S. Department of the Interior, Bureau of Land Mangement, Boise District.

20 p.

Monsen, S. B. 2000. Establishment of big sagebrush (Artemisia

tridentata) in semiarid environments. In: Entwistle, P. G.; DeBolt,

A. M.; Kaltenecker, J. H.; Steenhof, K., comps. Proceedings:

sagebrush steppe ecosystems; 1999 July 21–23; Boise, ID. Pub.

ID-PT-001001+1150USDI. Boise, ID: U.S. Department of the

Interior, Bureau of Land Management, Idaho State Office: 81–86.

Monsen, S. B.; McArthur, E. D. 1995. Implications of early Intermountain range and watershed restoration practices. In: Roundy,

B. A.; McArthur, E. D.; Haley, J. S.; Mann, D. K., comps. Proceedings: wildland shrub and arid land restoration symposium; 1993

October 19–21; Las Vegas, NV. Gen. Tech. Rep. INT-GTR-315.

Ogden, UT: U.S. Department of Agriculture, Forest Service,

Intermountain Research Station: 16–25.

Monsen, S. B.; Meyer, S. E. 1990. Seeding equipment effects on

establishment of big sagebrush on mine disturbances. In: Fifth

Billings symposium on disturbed land rehabilitation. Vol. I.

Hardrock waste, analytical, and revegetation; 1990 March 25–

30; Billings, MT. Reclamation Res. Unit Publ. 9003. Bozeman:

Montana State University: 192–199.

Monsen, S. B.; Meyer, S. E.; Carlson, S. C. 1992. Sagebrush establishment enhanced by snowfencing. In: Rangeland Technology Equipment Council 1992 annual report; 1992 February 9; Spokane, WA.

Missoula, MT: U.S. Department of Agriculture, Forest Service,

Missoula Technology and Equipment Development Center: 6–8.

National Research Council, Committee on Rangeland Classification, Board on Agriculture. 1994. Rangeland health—new methods to classify, inventory, and monitor rangelands. Washington,

DC: National Academy Press. 200 p.

Noss, R. F.; Peters, R. L. 1995. Endangered ecosystems. A status

report on America’s vanishing habitat and wildlife. Washington,

DC: Defenders of Wildlife. 132 p.

Paige, C.; Ritter, S. A. 1999. Birds in a sagebrush sea: managing

sagebrush habitats for bird communities. Boise, ID: Partners in

Flight Western Working Group. 47 p.

Plummer, A. P.; Christensen, D. R.; Monsen, S. B. 1968. Restoring

big game range in Utah. Publ. 68-3. Salt Lake City: Utah Division

of Fish and Game. 183 p.

Richardson, B. Z.; Monsen, S. B.; Bowers, D. M. 1986. Interseeding

selected shrubs and herbs on mine disturbances in southeastern

Idaho. In: McArthur, E. D.; Welch, B. L., eds. Proceedings—

symposium on the biology of Artemisia and Chrysothamnus; 1984

July 9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 134–139.

Schultz, L. M. 1986. Taxonomic and geographic limits of Artemisia

subgenus Tridentatae (Beetle) McArthur (Asteraceae:

Anthemideae). In: McArthur, E. D.; Welch, B. L., eds. Proceedings—symposium on the biology of Artemisia and Chrysothamnus;

1984 July 9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 20–29.

Schuman, G. E.; Booth, D. T.; Cockrell, J. R. 1998. Cultural methods

for establishing Wyoming big sagebrush on mined lands. Journal

of Range Management. 51: 223–230.
108 USDA Forest Service Proceedings RMRS-P-38. 2005

Shaw, DeBolt, and Rosentreter Reseeding Big Sagebrush: Techniques and Issues

Shaw, N. L.; Monsen, S. B. 1990. Use of sagebrush for improvement

of wildlife habitat. In: Fisser, H. G., ed. Proceedings—Wyoming

shrublands: aspen, sagebrush and wildlife management. Laramie:

University of Wyoming: 19–35.

Stevens, R. 1981. Techniques for planting shrubs on wildland

disturbances. In: Stetler, L. H.; DePuit, E. J.; Mikol, S. A., eds.

Proceedings—shrub establishment on disturbed arid and semiarid lands; 1980 December 1–2; Laramie, WY. Cheyenne: Wyoming Game and Fish Department: 29–36.

Stevens, R. 1994. Interseeding and transplanting to enhance species composition. In: Monsen, S. B.; Kitchen, S. G., comps. Proceedings—ecology and management of annual rangelands; 1992

May 18–22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT:

U.S. Department of Agriculture, Forest Service, Intermountain

Research Station: 300–306.

Stevens, R. 2002. Selecting adapted species, developing seeding

combinations, seeding rates, and methods of treatment to facilitate establishment and recovery of native species. In: Restoration

and management of sagebrush/grass communities workshop;

2002 November 4–8; Elko, NV. [Place of publication unknown]:

[Publisher unknown]: n.p. Abstract.

Stevens, Richard; Jorgensen, Kent R.; Davis, James N. 1981. Viability of seed from thirty-two shrub and forb species through fifteen

years of warehouse storage. Great Basin Naturalist. 41: 274–277.

Stevens, Richard; Jorgensen, Kent R.; Young, Stanford A.; Monsen,

Stephen B. 1996. Forb and shrub seed production guide for Utah.

Logan: Utah State University Extension. 51 p.

Stevens, Richard; Monsen, Stephen B. 2004. Guidelines for restoration and rehabilitation of principal plant communities. In: Monsen,

Stephen B.; Stevens, Richard; Shaw, Nancy L., comps. Restoring

Western ranges and wildlands. Gen. Tech. Rep. RMRS-GTR-136-

Vol-1. Fort Collins, CO: U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station: 199–294.

Tiedemann, Arthur R.; Klock, Glen O.; Mason, Lee L.; Sears, Donald

E. 1976. Shrub plantings for erosion control in eastern Washington—progress and research needs. Res. Note PNW-279. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific

Northwest Research Station. 11 p.

Vale, T. R. 1974. Sagebrush conversion projects: an element of

contemporary environmental change in the Western United

States. Biological Conservation. 6: 272–284.

Vernon, J. 2005. [Personal communication]. March 7. Ephraim, UT:

Utah Division of Wildlife Resources, Great Basin Research Center.

Wagstaff, F. J.; Welch, B. L. 1990. Rejuvenation of mountain big

sagebrush on mule deer winter ranges using onsite plants as a

seed source. In: McArthur, E. Durant; Romney, Evan M.; Smith,

Stanley D.; Tueller, Paul T., comps. Proceedings—symposium on

cheatgrass invasion, die-off, and other aspects of shrub biology

and management; 1989 April 5–7; Las Vegas, NV. Gen. Tech.

Rep. INT-276. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 171–174.

Wagstaff, F. L.; Welch, B. L. 1991. Seedstalk production of mountain

big sagebrush enhanced through short-term protection from

heavy browsing. Journal of Range Management. 44: 72–74.

Walton, T. P.; White, R. S.; Wambolt, C. L. 1986. Artemisia reproductive strategies: a review with emphasis on plains silver

sagebrush. In: McArthur, E. D.; Welch, B. L., eds. Proceedings—

symposium on the biology of Artemisia and Chrysothamnus; 1984

July 9–13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station: 67–74.

Wambolt, C. L.; Payne, G. F. 1986. An 18-year comparison of control

methods for Wyoming big sagebrush in southwestern Montana.

Journal of Range Management. 39: 314–319.

Wambolt, Carl L.; Harp, Aaron J.; Welch, Bruce L.; Shaw, Nancy;

[and others]. 2002. Conservation of greater sage-grouse on public

lands in the Western U.S.: implications of recovery and management policies. Caldwell, ID: Policy Analysis Center for Western

Public Lands. 41 p.

Welch, B. L. 1995. Beyond twelve percent purity. In: Roundy, B. A.;

McArthur, E. D.; Haley, J. S.; Mann, D. K., comps. Proceedings:

wildland shrub and arid land restoration symposium; 1993 October 19–21; Las Vegas, NV. Gen. Tech. Rep. INT-GTR-315. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 126–129.

Welch, B. L.; McArthur, E. D.; Nelson, D. L.; Pederson, J. C.; Davis,

J. N. 1986. ‘Hobble Creek’—a superior selection of low-elevation

mountain big sagebrush. Res. Pap. INT-370. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station. 10 p.

Welch, B. L.; Nelson, E. D.; Young, S. A.; Sands, A. R.; Wagstaff, F. J.;

Nelson, D. L. 1992. ‘Gordon Creek’—a superior, tested germplasm

of Wyoming big sagebrush. Res. Pap. INT-461. Ogden, UT: U.S.

Department of Agriculture, Forest Service, Intermountain Research Station. 7 p.

Welch, B. L.; Wagstaff, F. J.; Williams, R. L. 1990. Sage grouse

status and recovery plan for Strawberry Valley, Utah. Res. Pap.

INT-430. Ogden, UT: U.S. Department of Agriculture, Forest

Service, Intermountain Research Station. 10 p.

West, N. E. 2000. Synecology and disturbance regimes of sagebrush

steppe ecosystems. In: Entwistle, P. G.; DeBolt, A. M.; Kaltenecker,

J. H.; Steenhof, K., comps. Proceedings—sagebrush steppe ecosystems; 1999 July 21–23; Boise, ID. Pub. ID-PT-001001+1150USDI.

Boise, ID: U.S. Department of the Interior, Bureau of Land

Management, Idaho State Office: 15–26.

West, N. E.; Hassan, M. A. 1985. Recovery of sagebrush-grass vegetation following wildfire. Journal of Range Management. 38: 131–134.

Wiedemann, H. T. 2005. Revegetation equipment catalog. College

Station, TX: Revegetation Equipment Technology Committee,

U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior, Bureau of Land Management. [Online].

Available: http://reveg-catalog.tamu.edu/ [2005, March 6].

Winward, A. H. 1985. Fire in the sagebrush-grass ecosystem—the

ecological setting. In: Sanders, K.; Durham, J., eds. Rangeland

fire effects—a symposium; 1984 November 27–29; Boise, ID.

Boise, ID: U.S. Department of Interior, Bureau of Land Management, Idaho State Office: 2-6.

Young, J. A.; Evans, R. A. 1975. Germinability of seed reserves in a

big sagebrush community. Weed Science. 23: 358–364.

Young, J. A.; Evans, R. A. 1978. Population dynamics after wildfires

in sagebrush grasslands. Journal of Range Management. 31:

283–289.

Young, J. A.; Evans, R. A. 1989. Dispersal and germination of big

sagebrush (Artemisia tridentata) seeds. Weed Science. 37:

201–206.

Young, J. A.; Evans, R. A.; Palmquist, D. E. 1989. Big sagebrush

(Artemisia tridentata) seed production. Weed Science. 37: 47–53.

Young, J. A.; Evans, R. A.; Palmquist, D. E. 1990. Soil surface

characteristics and emergence of big sagebrush seedlings. Journal of Range Management. 43: 358–367.

Young, J. A.; Martens, E. 1991. Importance of hypocotyls hairs in

germination of Artemisia seeds. Journal of Range Management.

43: 358–366.

Big sagebrush
USDA Forest Service Proceedings RMRS-P-38. 2005 109

Carl L. Wambolt is Professor of Range Science, Department of Animal and

Range Sciences, Montana State University, Bozeman, MT 59717 U.S.A.,

FAX: (406) 994-5589, e-mail: cwambolt@montana.edu

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Sagebrush-Ungulate

Relationships on the Northern

Yellowstone Winter Range

Carl L. Wambolt

Abstract—Sagebrush (Artemisia) taxa have historically been the

landscape dominants over much of the Northern Yellowstone Winter Range (NYWR). Their importance to the unnaturally large

ungulate populations on the NYWR throughout the twentieth

century has been recognized since the 1920s. Sagebrush-herbivore

ecology has been the focus of research on the NYWR for 2 decades.

Specific research topics include general ungulate use of the NYWR,

forage relationships involving sagebrush taxa, the effects of browsing

and sagebrush taxa on forage production, the results of long-term rest

from severe browsing on the NYWR, sagebrush plant and community

characteristics related to herbivory, and the relationships of sagebrush taxa to fire on the NYWR. Sagebrush taxa have been very

important on the NYWR but are generally in a state of decline.

The Northern Yellowstone Winter Range (NYWR) is dominated by big sagebrush (Artemisia tridentata) taxa. Big

sagebrush occupies relatively snow-free portions of the

NYWR, making it accessible to ungulates throughout the

winter. Heavy foraging use of sagebrush by the large ungulate populations of the region has been discussed by naturalists for almost a century (Wright and Thompson 1935) (fig.

1). My research has confirmed that early naturalists were

right to be concerned about sagebrush habitat on the NYWR.

Those investigations have determined the mechanisms that

influence sagebrush-herbivore interactions on the NYWR

and their effects.

Area Description ________________

The NYWR is an extensive area covering about 100,000 ha

along the lower elevations in Northern Yellowstone National Park (YNP), and extends northward into Montana

along the Yellowstone River drainage (fig. 2). The Lamar

and Gardiner River drainages, important lowland areas

within YNP, are relatively free of snow and provide reliable

winter foraging for ungulates. During the winter, 80 percent

of all ungulates are found on the NYWR within YNP. In

addition, there have been as many as 2,544 Rocky Mountain

mule deer and 8,626 Rocky Mountain elk wintering on the

portion of the NYWR north of the YNP boundary in Montana

during the last decade.

Figure 1—Photo from Wright and Thompson (1935).

The original caption read: “End of the range. The right

side of the picture shows elk winter range within

Yellowstone National Park. The fence is the boundary.

The left side of the picture shows the rank growth of

sagebrush just outside the park. Here we can compare

the original and the present state of the range. There

can be no doubt of what has happened. (Photograph

taken June 1, 1932, near Gardiner, Wildlife Division

No. 2501.)”

The climate is favorable in this area for ungulate foraging

during winter. The mean annual precipitation is approximately 280 mm (1,616 m) at Gardiner, MT, 400 mm (1,899 m)
110 USDA Forest Service Proceedings RMRS-P-38. 2005

Wambolt Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter RangeFigure 2—The Northern Yellowstone Winter Range, Montana and Wyoming (National Park Service Map).
USDA Forest Service Proceedings RMRS-P-38. 2005 111

Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter Range Wambolt

at Mammoth, and 410 mm (1,912 m) at Tower Falls. About

half of the moisture each year is received as snow, the peak

occuring in spring and early summer.

The Gardiner Basin, an area near the town of Gardiner,

MT, includes the boundary line area (BLA), and has less

snow than the majority of the NWYR that lies to the east in

YNP. Therefore, the Gardiner Basin is especially important

for pronghorn and mule deer. Elk are usually able to negotiate deeper snow at higher elevations. Elk and bison tend to

dominate the foraging over the remaining portions of the

NYWR, but there is no strong evidence that bison have had

an impact on the shrubs of sagebrush habitats. Therefore,

elk are the only browsing ungulate found in significant

numbers during winter on the portions of the NYWR outside

the Gardiner Basin.

Not only is big sagebrush the largest vegetation type on

the NYWR, but perhaps more importantly, sagebrush taxa

are the dominant vegetative form on the portions of the

NYWR that are most valuable as winter range for ungulates

(DeSpain 1990). Sagebrush communities also furnish important security and thermal cover for ungulates and other

animals (Wambolt and McNeal 1987).

The variety of sagebrush habitats within NYWR includes

the Wyoming big sagebrush- (Artemisia tridentata

wyomingensis) bluebunch wheatgrass (Agropyron spicatum)

type that is found primarily at lower elevations in the

Gardiner Basin. Mountain big sagebrush- (A. t. vaseyana)

Idaho fescue (Festuca idahoensis) habitat type dominates

the majority of the NYWR (Wambolt and Sherwood 1999).

Other important sagebrush are basin big sagebrush (A. t.

tridentata) and black sagebrush (A. nova). Important subdominant plants are the sprouting shrubs often found in

sagebrush habitats including rubber rabbitbrush

(Chrysothamnus nauseosus), green rabbitbrush (C.

viscidiflorus), and gray horsebrush (Tetradymia canescens).

These shrubs are found throughout the NYWR in various

sagebrush habitat types.

Results and Discussion __________

Sagebrush Forage Relationships

The relationships of secondary compounds found in the

foliage of sagebrush taxa to foraging by NYWR ungulates

was studied. The relationships between crude terpenoid

content of sagebrush and in vitro organic matter digestibility (IVOMD) as well as preference by mule deer was considered for the four dominant sagebrush taxa found on the

NYWR. Basin big sagebrush was the most easily digested by

mule deer followed in order of decreasing digestibility by

Wyoming big sagebrush, mountain big sagebrush, and black

sagebrush. The IVOMD for all four taxa generally increased

from January to April as crude terpenoids decreased in the

foliage. When the crude terpenoids were removed from the

sagebrush foliage the IVOMD increased. However, terpenoid removal resulted in few differences in IVOMD among

the taxa and collection dates during the winter (Striby and

others 1987). Wild mule deer from the NYWR were compared to domestic sheep and steers for their ability to digest

sagebrush. There were no significant differences found among

the three species of animals; therefore, it appears that there

should be no significant differences between the wild ungulates (mule deer, elk, and pronghorn) found on the NYWR.

Personius and others (1987) isolated 31 compounds from

sagebrush foliage and found that seven of them were particularly important for preference indicators among the four

sagebrush taxa. In order to separate the individual plants

within a taxon into browse-form classes (use classes), seven

other compounds were helpful. The compounds above were

used by Bray and others (1991) to differentiate mule deer

preference among sagebrush taxa during feeding trials. This

study determined if the deer discriminated against forage

treated with the individual compounds. During this feeding

trial, mule deer selected significantly higher quantities of

untreated forage compared to the same forage treated with

the suspect compounds (table 1). These studies found that

volatility of compounds had little influence on preference at

the concentrations found in nature. Both volatile and nonvolatile compounds were found to be responsible for deterring foraging on the four taxa. The combination of nonvolatile sesquiterpene lactones with certain volatiles explained

why one sagebrush taxon is often preferred over another.

Site characteristics that might be responsible for elk and

mule deer foraging on the NYWR were studied (Wambolt

and McNeal 1987). Most portions of the Gardiner Basin —

even the timbered areas—were used by elk. However, elk fed

mostly in mountain big sagebrush habitat types. The mule

deer in the same area preferred to forage in the Wyoming big

sagebrush type when it was found adjacent to steep and dry

topography at lower elevations. These areas tended to furnish good security and thermal cover while also meeting

foraging needs. Elk distribution and concentration on the

NYWR vary with wind, snow depth, temperature, and snow

crusting that might expose or conceal forage. In addition,

hunting north of YNP affects animal distribution on the

winter range.

Mule deer and elk preferences for the four sagebrush taxa

were studied over 10 winters of varying severity in the

Gardiner Basin (Wambolt 1996). This study was purposely

long term in order to avoid anomalies over shorter periods

that might lead to erroneous conclusions. Approximately

2,500 leaders were tagged on 244 plants each winter over

the 10-year period to determine if browsing had occurred.

Table 1—Mule deer preference for seven

compounds found in four sagebrush taxa. All seven significantly

influence intake and are listed in

decreasing order as a foraging

deterant.

Compound

1,8-cineole a

Black sage NVCTF b

Wyoming big sage NVCTF

p-cymene a

Basin big sage NVCTF

Methacrolein a

Mountain big sage NVCTF

a

 Volatile.

b

 Nonvolatile crude terpenoid fraction.
112 USDA Forest Service Proceedings RMRS-P-38. 2005

Wambolt Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter Range

Overall, the two ungulates browsed large amounts of the

four sagebrush taxa, particularly considering that the 10-

year period was below average for winter severity. In addition, mule deer diets averaged 52 percent big sagebrush over

the 10-year study. The diets were determined by

microhistological techniques on composite samples of feces

collected early each spring. Both ungulates displayed a

distinct preference among the four taxa. The amount of

sagebrush use varied with winter and taxon, as expected.

Use reached 91 percent for the preferred taxon, mountain

big sagebrush, which averaged 56 percent over the 10-year

study (fig. 3). Average use of Wyoming big sagebrush was 39

percent and basin big sagebrush was 30 percent. The least

preferred taxon was black sagebrush at 17 percent. The

results of our study of actual use by wild ungulates supported the findings of Personius and others (1987) and Bray

and others (1991) that sagebrush terpenoids affect herbivory.

Regression models were developed to determine the production of winter forage for the three big sagebrush subspecies on the NYWR (Wambolt and others 1994). Models

available in previous literature did not consider the variation among sagebrush subspecies nor use classes (developmental differences from past browsing) that might be present.

During this study (Wambolt and others 1994), the consideration of taxon and form class increased the Ra

2 an average of

10 percent with values to 0.90. These models help with the

determination of carrying capacity, detecting trends in forage production, and measuring plant responses to different

management options. In addition, they clearly show the

importance of taxon recognition and the role of past

browsing history on present forage production of sagebrush

taxa (table 2).

Ecology of Big Sagebrush on the NYWR

The question of whether Wyoming big sagebrush plants

that had been protected from browsing for 35 years would

exhibit similar growth characteristics to browsed plants was

considered (Hoffman and Wambolt 1996). To test this hypothesis, paired comparisons in and out of a 2-ha National

Park Service (NPS) exclosure constructed in 1957 were

made. Because heavy browsing had occurred outside the

exclosure for many years, plants there had no terminal

leader growth. Plants inside the exclosure were dominated

by terminal growth, and axial long shoots were rare. Therefore, to further investigate differences between protected and

unprotected plants it was necessary to compare terminal

Figure 3—Elk browsing mountain big sagebrush on the Northern Yellowstone winter range.

Table 2—Variables selected to model big sagebrush winter forage production (g).

Variables

Average seedhead weight (g)

Height (cm)

Average cover (cm)

Crown depth (cm)

Circular area (cm2)

Major axis (cm)

Minor axis (cm)
USDA Forest Service Proceedings RMRS-P-38. 2005 113

Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter Range Wambolt

leaders of protected plants inside the exclosure to axial long

shoots on browsed plants. The unbrowsed plants had a

consistently higher production level than browsed plants

(table 3). The average production per plant was 10 g with

browsing and 45 g with protection. The browsed plants had

large amounts of dead crown as well, although no measurements were taken of this parameter. Production of seedheads

by big sagebrush plants was the parameter with the greatest

difference between browsed and unbrowsed plants. The

browsed plants averaged 0.08 seedheads per plant, where

the unbrowsed plants averaged 60.3 per plant. Almost all

terminal leaders on browsed plants were removed so flowering stems would have to be initiated from elsewhere

(Hoffman and Wambolt 1996). This loss of seedhead production from browsing has undoubtedly resulted in reproductive

declines for NYWR sagebrush, as these taxa do not reproduce

asexually.

The importance of average seedhead production as expressed by their weight was discovered to be important for

the construction of regression models to predict sagebrush

production of winter forage from the three big sagebrush

subspecies found on the NYWR (Wambolt and others 1994).

Models improved for all three taxa when average seedhead

weight was included for the lightly used (browse form class)

plants. Because heavily used plants did not produce many

inflorescences, the addition of average seedhead weight to

the model for those plants was not beneficial. Separation of

browse-form classes and inclusion of average seedhead

weight in the models both recognized the impact browsing

has had on the annual production and reproduction of

NYWR sagebrush.

Snow cover is naturally light on the NYWR, but some

snow falls each winter and may protect small sagebrush

plants for several years before ungulates find them available for foraging. Among the mountain big sagebrush plants

that established between the years 1978—1992, nearly half

(47 percent) germinated in 1988. That was the year of the

Yellowstone fires. That was also a year that offered relatively good seed production due to good spring moisture

followed by a winter with considerably more snow than other

winters during the 15-year period. The snow protected the

seedlings from herbivores and provided an insulated environment for their welfare during this usually precarious

establishment period. These conditions also coincided with

a 35 to 40 percent reduction in elk that occurred during that

most severe winter (1988—1989) of the 15-year period. This

also benefited the seedlings that were allowed to establish

during the next several years of reduced elk numbers.

Fire Relationships

The NYWR sagebrush taxa have no mechanisms to cope

well with fire. Therefore, when fire has occurred on NYWR

sagebrush habitats, they have been an additive injury, along

with intense herbivory, to sagebrush plants. A wildfire in the

Gardiner Basin was studied (Wambolt and others 1999) to

determine, among other things, the rate of reestablishment

of sagebrush and rabbitbrush taxa under continuous browsing following fire. Nineteen years after the fire, recovery of

the three subspecies of big sagebrush was at levels between

1 and 20 percent (canopy coverage) of that in adjacent unburned sagebrush stands. This was found despite the fact that

the adjacent stands were already in decline from historically

heavy browsing (Wambolt 1996; Wambolt and Sherwood

1999). The recovery of Wyoming big sagebrush was less

than that of mountain or basin big sagebrush in this burn.

Similar relationships were also found for big sagebrush

density and production when comparing burned areas to

unburned portions.

Wambolt and others (1999) also studied seven prescribed

burns conducted by the USDA Forest Service on mountain

big sagebrush sites within the Gardiner Basin. These burns

had been conducted 10 to 14 years before they were studied.

The burns were compared with 33 unburned sites in the

Basin to determine recovery following burning. The canopy

coverage and densities of the sagebrush on the unburned

sites averaged 12 and 15 times greater than on burned sites.

This confirms that burning accelerated the browsing-induced decline (Wambolt 1996; Wambolt and Sherwood 1999)

of NYWR sagebrush. Sagebrush not directly eliminated by

fire were also subjected to unusually heavy browsing on

surviving or reestablishing shrubs due to the loss of normally available forage from the fires.

Rens (2001), when studying the effects of fire on the

NYWR, considered the effect of elk browsing on the recovery

from fires on the Black-tailed Deer Plateau. This investigation also found significant differences in the development of

protected and browsed shrubs. One decade after burning

mountain big sagebrush, canopy cover averaged 20 percent

with protection from elk browsing and 9.7 percent where

browsed. Similar differences were found in density and

production of protected and browsed mountain big sagebrush following fire on the Black-tailed Deer Plateau.

Browsing Impacts

Wambolt (1996) found that 35 percent of the heavily

browsed mountain big sagebrush plants died between 1982

and 1992. Most of the plants that survived the decade of

heavy browsing developed a form that clearly showed the

effects of their past heavy use, and contained a high percentage of dead crown. The amount of dead crown found in the

three big sagebrush subspecies was in direct proportion to

the amount of browsing received by each taxon. Among the

plants that survived the heavy browsing, the percentage of

dead crown from mountain big sagebrush, Wyoming big

sagebrush, and basin big sagebrush, was 58.7, 45.4, and 30.1

percent, respectively (Wambolt 1996).

The impact of herbivory on sagebrush across the NYWR

was studied by Wambolt and Sherwood (1999) using NPS

Table 3—Average differences between browsed and unbrowsed

Wyoming big sagebrush plants.

Browsed Unbrowsed

Production (g/plant) 10.0a 44.7b

Seedheads per plant 0.08a 60.3b

Leader length (mm) 22.9a 22.3a

Leader dry weight (g) 0.02a 0.02a

Values followed by different letters are significantly different (P <0.01).
114 USDA Forest Service Proceedings RMRS-P-38. 2005

Wambolt Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter Range

exclosures that were constructed in 1957 and 1962. The

exclosures were originally constructed to learn what would

happen under protection from the heavy herbivory that

was occurring throughout the northern part of Yellowstone

Park. Wambolt and Sherwood (1999) compared shrub parameters between browsed and unbrowsed plants with the

aid of these exclosures. They discovered a significant difference between the development of protected and browsed big

sagebrush communities across the NYWR.

Since the period of exclosure construction in 1957 and

1962, there has been a significant difference in the development of protected and browsed big sagebrush communities.

Average big sagebrush canopy cover on protected sites was

202% greater (P ≤ 0.0027) than on browsed sites over the 19-

paired sites. The average big sagebrush cover for all 19 sites

was 19.7% inside and 6.5% outside the exclosures. This relationship was universal on sites with Wyoming big sagebrush

or mountain big sagebrush, flat to very steep topographies,

and all aspects and precipitation levels (Wambolt and Sherwood

1999).

Wambolt and Sherwood (1999) found that Wyoming big

sagebrush sites that were protected by the exclosures averaged almost 10 times more sagebrush cover than where

browsing had continued unabated (fig. 4). Where mountain

big sagebrush was the dominant cover type, the same figure

was almost three times as much sagebrush cover where

protected.

The density of big sagebrush plants was affected in a

similar manner by ungulate browsing. Across the NYWR,

big sagebrush plants were twice as numerous where they

had not been browsed in exclosures as where browsing had

occurred throughout the interval since exclosure construction. This amounted to an average density of 30.5 plants per

60 m2 inside and 15.3 per 60 m2 outside the exclosures. In

addition, individual mountain big sagebrush plants produced 88 percent more winter forage where protected. A

similar production figure was not practical to obtain from

Wyoming big sagebrush plants even though the differential

between protected and browsed plants was even greater

than it was for mountain big sagebrush. The Wyoming big

sagebrush plants could not be quantified for this character

because browsed plants were so reduced in size that their

parameters were not suitable for inclusion in the models.

Summary ______________________

Both the historical evidence and our recent studies clearly

show a significant decline of sagebrush on the NYWR. This

decline has already impacted ungulates that rely on sagebrush habitats for nutritional and other requirements (fig. 5).

Among the ungulates, it is likely that elk will be the least

impacted from the decline in the sagebrush habitat types

Figure 4—National Park Service exclosure near Gardiner, MT, erected in 1957 after Wyoming big sagebrush

was nearly eliminated by browsing. Shrubs inside show some recovery (photo taken September 2001).
USDA Forest Service Proceedings RMRS-P-38. 2005 115

Sagebrush-Ungulate Relationships on the Northern Yellowstone Winter Range Wambolt

because of their abilities to use other types when available.

However, it is clear that elk browsing has been responsible

for declines of sagebrush over the majority of the NYWR

(Wambolt and Sherwood 1999). It is logical to assume that

numerous fauna will be affected by the decline of any native

vegetative type. This is especially true with one as extensive

as the sagebrush type in the region of the NYWR. The

decline of an important habitat type should be given serious

consideration when land management opportunities present

themselves.

References _____________________

Bray, R. O.; Wambolt, C. L.; Kelsey, R .G. 1991. Influence of

sagebrush terpenoids on mule deer preference. Journal of Chemical Ecology. 17: 2053–2062.

Hoffman, T. L.; Wambolt, C. L. 1996. Growth response of Wyoming

big sagebrush to heavy browsing by wild ungulates. In: Barrow,

J. R.; McArthur, E. D.; Sosebee, R. E.; Rausch, R. E., comps.

Proceedings—symposium on shrubland ecosystem dynamics in a

changing environment. Gen. Tech. Rep. INT-GTR-338. Ogden,

UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 242–245.

Personius, T. L.; Wambolt, C. L.; Stephens, J. R.; Kelsey, R. G. 1987.

Crude terpenoid influence on mule deer preference for sagebrush.

Journal of Range Management. 40: 84–88.

Rens, Reyer J. 2001. Elk effects on sagebrush-grassland after fire on

Yellowstone’s northern range. Bozeman: Montana State University. Thesis.

Striby, K. D.; Wambolt, C. L.; Kelsey, R. G.; Havstad, K. M. 1987.

Crude terpenoid influence on in vitro digestibility of sagebrush.

Journal of Range Management. 40: 244–248.

Wambolt, C. L. 1996. Mule deer and elk foraging preference for 4

sagebrush taxa. Journal of Range Management. 49: 499–503.

Wambolt, C. L. 1998. Sagebrush and ungulate relationships on

Yellowstone’s northern range. Wildlife Society Bulletin. 26:

429–437.

Wambolt, C. L.; Creamer, W. H.; Rossi, R. J. 1994. Predicting big

sagebrush winter forage by subspecies and browse form class.

Journal of Range Management. 47: 231–234.

Wambolt, C. L.; Frisina, M. R.; Douglass, K. S.; Sherwood, H. W.

1997. Grazing effects on nutritional quality of bluebunch wheatgrass for elk. Journal of Range Management. 50: 503–506.

Wambolt, C. L.; Hoffman, T. 2001. Mountain big sagebrush age

distribution and relationships on the Northern Yellowstone winter range. In: McArthur, E. D.; Fairbanks, D. J., comps. Proceedings: shrubland ecosystem genetics and biodiversity. Proc. RMRSP-21. Fort Collins, CO: U.S.Department of Agriculture, Forest

Service, Rocky Mountain Research Station: 330–335.

Wambolt, C. L.; Hoffman, T.; Sherwood, H. 2000. Browsing effect on

Wyoming big sagebrush by antelope and other ungulates. In:

Fischer, J. K., ed. Proceedings: 17th biennial pronghorn antelope

workshop; Lake Tahoe. California Department of Fish and Game:

73–76.

Wambolt, C. L.; Hoffman, T. L.; Mehus, C. A. 1999. Response of

shrubs in big sagebrush habitats to fire on the Northern

Yellowstone winter range. In: McArthur, E. D.; Ostler, W. K.;

Wambolt, C. L., comps, Proceedings: shrubland ecotones. Proc.

RMRS-P-11. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station: 238–242.

Wambolt, C. L.; McNeal, A. F. 1987. Selection of winter foraging

sites by elk and mule deer. Journal of Environmental Management. 25: 285–291.

Wambolt, C. L.; Sherwood H. W. 1999. Sagebrush response to

ungulate browsing in Yellowstone. Journal of Range Management. 52: 363–369.

Wright, G. M.; Thompson, G. H. 1935. Fauna of the National Parks

of the United States: wildlife management in the National Parks.

Fauna Series No. 2. Washington, DC: U.S. Government Printing

Office. 142 p.

Figure 5—Even large ungulates use sagebrush for cover. This is one of a dozen elk standing in a patch

of basin big sagebrush on the NYWR. Only a portion of the elk’s head is visible in the center of photo.
116 USDA Forest Service Proceedings RMRS-P-38. 2005

Nancy Hampton is an Ecologist, Idaho National Laboratory/Idaho State

University, P.O. Box 1625, Idaho Falls, ID 83415-2213; e-mail: hampnanc@isu.edu

In: Shaw, Nancy L.; Pellant, Mike; Monsen, Stephen B., comps. 2005. Sagegrouse habitat restoration symposium proceedings; 2001 June 4–7; Boise, ID.

Proceedings RMRS-P-38. Fort Collins, CO: U.S. Department of Agriculture,

Forest Service, Rocky Mountain Research Station.

Insects of the Idaho National

Laboratory: A Compilation

and Review

Nancy Hampton

Abstract—Large tracts of important sagebrush (Artemisia L.)

habitat in southeastern Idaho, including thousands of acres at the

Idaho National Laboratory (INL), continue to be lost and degraded

through wildland fire and other disturbances. The roles of most

insects in sagebrush ecosystems are not well understood, and the

effects of habitat loss and alteration on their populations and

communities have not been well studied. Although a comprehensive survey of insects at the INL has not been performed, smaller

scale studies have been concentrated in sagebrush and associated

communities at the site. Here, I compile a taxonomic inventory of

insects identified in these studies. The baseline inventory of more

than 1,240 species, representing 747 genera in 212 families, can be

used to build models of insect diversity in natural and restored

sagebrush habitats.

Introduction ___________________

The Idaho National Laboratory (INL), formerly the Idaho

National Engineering and Environmental Laboratory, is

located in a cool desert ecosystem characterized by shrubsteppe vegetation communities typical of the northern Great

Basin and Columbia Plateau region. Established in 1949 to

carry out nuclear energy research and related activities,

public access to the 570-thousand-acre INL facility has been

restricted for over 50 years. As a consequence, large remnants of relatively undisturbed sagebrush-steppe are still

preserved in the interior portion of the site (Anderson 1999).

In recognition of the ecological importance of INL lands, the

facility was designated as a National Environmental Research Park in 1975 (DOE 1985).

As these important habitats continue to be lost and

degraded, interest in the status and condition of remaining

sagebrush communities has grown (Entwistle and others

2000; Knick 1999; Knick and Rotenberry 1997). Much is

unknown about these ecosystems, and there is an immediate need to establish baselines, fill information voids, and

focus research on critical issues, including restoration

alternatives.

Major portions of the INL have been burned by wildfires

over the past several years, and restoration and recovery of

sagebrush habitat are current topics of investigation (Anderson and Patrick 2000; Blew 2000). Most restoration projects,

including those at the INL, are focused on the reestablishment of vegetation communities (Anderson and Shumar

1989; Williams 1997). Insects also have important roles in

restored communities (Williams 1997) and show promise as

indicators of restoration success in shrub-steppe (Karr and

Kimberling 2003; Kimberling and others 2001) and other

habitats (Jansen 1997; Williams 1997).

The purpose of this paper is to present a taxonomic list of

insects identified by researchers studying cold desert communities at the INL. Insects act as herbivores, decomposers,

pollinators, and predators, and they are major prey for

reptiles, mammals, and birds inhabiting sagebrush communities, including sage-grouse chicks [Centrocercus urophasianus

(Bonaparte)]. However, the function of most insects in sagebrush ecosystems is not well understood (West 1983). As

natural sagebrush communities disappear, remaining habitats such as those at the INL represent important resources

for establishing baseline attributes against which restoration success or indicator measures can be evaluated. A

taxonomic inventory is useful for developing reference models of natural insect diversity and community composition in

sagebrush habitats at the INL.

Methods _______________________

I constructed a baseline taxonomic list of insect species

documented in six major investigations at the INL (Allred

1968a; Allred and Cole 1971; Bohart and Knowlton 1977;

Karr and Kimberling 2003; Stafford 1983, 1987; Stafford

and Johnson 1986; Stafford and others 1986). The initial list

was expanded to include species from 16 smaller studies in

which insects were identified or collected (Allred 1970; Blom

and others 1991; Bromenshenk 1987; Cieminski and Flake

1995; Clark and Blom 1988, 1991, 1992, 1999; Johnson and

Stafford 1986; Merickel and Clark 1994; Stafford and Johnson

1986; Vieth 1983; Wenninger 2001; Winter 1984; Youtie

1986; Youtie and others 1987). A resident collection of INL

voucher specimens for 466 species, primarily collected and

identified by M. P. Stafford, W. H. Clark, and P.E. Blom, was

also incorporated into the list. All insects identified to family

were included. Although over 100 species of other arthropods,

including ticks, mites, spiders, solpugids, and scorpions, have

been collected and identified at the INL (Allred 1968b, 1969,
USDA Forest Service Proceedings RMRS-P-38. 2005 117

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

1970, 1973; Allred and Muma 1971; Karr and Kimberling

2003; Wenninger 2001), only insect species were compiled

for this list. Ordinal names and placement of families and

genera are according to Borror and others (1992).

To minimize the potential for double counting, I included

only the maximum number of unidentified species within

the same family or genus cited in any single reference. The

number of unidentified species was then reduced for each

unique species within the same family or genus cited in

additional references. Thus, hundreds of unidentified specimens were excluded from this inventory and remain to be

examined further.

Taxonomic authority was compiled from original documentation, voucher specimens, and other local and regional

species lists (Haws and others 1988; Horning and Barr

1970). The INL list was not reviewed for synonymy or

misidentifications. Discrepancies in spelling between INL

studies and other authorities were reconciled to Horning

and Barr (1970) where possible. Otherwise, spellings of

Haws and others (1988) or Arnett (2000) were adopted.

Results and Discussion __________

A list of over 1,240 insect species from 17 orders, representing 747 genera in 212 families, was compiled from 22

studies conducted at the INL (table 1). Insect sampling has

been widely distributed across the INL (fig.1), but most

studies were of short duration and focused on associations

Table 1—List of documented insect species at the Idaho National Engineering and Environmental Laboratory.

ORDERa ORDER

Familya Family

Scientific nameb Scientific name

COLLEMBOLA

Onychiuridae

spp. undetermined

Entomobryidae

spp. undetermined

Isotomidae

unident. sp. #1

Sminthuridae

unident. sp. #1

EPHEMEROPTERA

Baetidae

spp. undetermined

Caenidae

spp. undetermined

ISOPTERA

Kalotermitidae

unident. sp. #1

ODONATA

Aeshnidae

unident. sp. #1

Libellulidae

unident. sp. #1

Coenagrionidae

spp. undetermined

PSOCOPTERA

Liposcellidae

unident. sp. #1

PHTHIRAPTERA

Enderleinellidae

Enderleinellus sp.

Hoplopleuridae

Hoplopleura acanthopus

Hoplopleura arboricola

Hoplopleura erratica

Hoplopleura hesperomydis (Osborn)

Hoplopleura minimus

Polyplacidae

Fahrenholzia pinnata

Fahrenholzia sp. #2

Haemodipus setoni

Neohaematopinus inornatus Kellogg

Neohaematopinus laeviusculus

Neohaematopinus marmota

Neohaematopinus pacificus

Neohaematopinus sp. #5

Polyplax auricularis Kellogg and Ferris

Polyplax spinulosa Burmeister

Polyplax sp. #3

Trichodectidae

Geomydoecus sp.

Neotrichodectes interruptofasciatus

Mallophaga

spp. undetermined

ORTHOPTERA

Acrididae

Arphia pseudonictana Thomas

Aulocara elliotti Thomas

Cratypedes lateritius

Hesperotettix viridis Thomas

Melanoplus sanguinipes (Fabricius)

Trimerotropis gracilis

unident. spp. #1, #2

Tettigoniidae

unident. sp. #1

Gryllacrididae

Ceuthophilus maculatus

Gryllidae

Oecanthus sp.

unident. spp. #1, #2

HEMIPTERA

Enicocephalidae

unident. sp. #1

Corixidae

Cenocorixa wileyae

Sigara alternata

Notonectidae

spp. undetermined

Tingidae

Aclypta cooleyi

unident. sp. #2

Miridae

Atomoscelis modestus VanDuzee

Chlamydatus artemisiae

Chlamydatus associatus (Uhler)

(con.)
118 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Chlamydatus obliquus (Uhler)

Chlamydatus sp. #4

Coquilletia insignis Uhler

Coquilletia sp. #2

Deraeocoris bakeri Knight

Deraeocoris brevis Uhler

Deraecoris schwartzi (Uhler)

Hadronema simplex Knight

Hesperocapsus davisi

Illnacorella argentata Knight

Irbisia pacificus (Uhler)

Labopidea sericata (Uhler)

Labops utahensis Slater

Litomeris debilis (Uhler)

Lopidea sp.

Lygus desertinus Knight

Lygus elisus VanDuzee

Lygus hesperus Knight

Lygus sp. #4

Melanotrichus albocostatus

Orectoderus arcuatus

Orthotylus coagulatus (Uhler)

Parthenicus sp.

Phyllopidea hirta

Phyllopidea picta Uhler

Phytocoris sp.

Pilophorus sp.

Plagiognathus sp.

Polymerus diffusus Knight

Psallus pilosulus Uhler

Psallus sp.

Slaterocoris pilosipes

Slaterocoris utahensis

Slaterocoris sp. #3

Stenodema laevigatum (L.)

Stenodema pilosipes

Stenodema virens (L.)

Stenodema vicinum

Thyrillus pacificus (Uhler)

Trigonotylus ruficornis (Geoffroy)

Nabidae

Nabis alternatus (Parshley)

Reduviolus alternatus

Anthocoridae

Orius tristicolor White

Reduviidae

Sinea diadema (F.)

Zelus tetracanthus

unident. sp. #1

Piesmatidae

Piesma sp.

Phymatidae

unident. sp. #1

Corimelaenidae

Corimelaena virilis M. and Mc.

Lygaeidae

Blissus sp.

Emblethis vicarius

Geocoris pallens Stal

Leptoterna sp.

Lygaeus kalmii Stal

Malezonotus sp.

Megalonotus sabicula

Nysius ericae (Schilling) Auctorum

Nysius niger

Nysius raphanus Horvath

Ortholomus scolopax (Say)

Sisamnes claviger Coreidae

Chelinidae vittiger

Rhopalidae

Arhyussus sp.

Corizus punctiventris Dallas

Corizus scutatus (Stal)

Harmostes reflexulus (Say)

Leptocoris trivittatus (Say)

Liorhyssus hyalinus (F.)

Cydnidae

unident. sp. #1

Scutelleridae

Vanduzeina balli

unident. sp. #1

Pentatomidae

Aelia americana Dallas

Chlorochroa sayi Stal

Codophila remota Horvath

Prionosoma podopioides

Rhytidilomia uhleri Stal

HOMOPTERA

Cicadidae

Okanagana annulata Davis

Okanagana bella Davisc

Okanagana luteobasalis Davis

Platypedia putnami lutea Davis

Membracidae

Campylenchia latipes (Say)

Tortistillus wickhami VanDuzee

Aetalionidae

Aetalion sp.

Cercopidae

Clastoptera brunnea Bale

Clastoptera delicata Uhler

Neophilaenus lineatus (L.)

Philaronia sp.

Cicadellidae

Aceratagallia poudris (Stal)

Aceratagallia sp. #2

Athysanella spp. #1, #2

Balclutha sp.

Ballana hebea

Ballana sp. #2

Calladonus montanus VanDuzee

Carsonus aridus

Ceratagallia artemisia Oman

Chlorotettix unicolor Fitch

Circulifer tenellus Baker

Comellus sp.

Dikraneura carneola (Stal)

Empoasca alboneura

Empoasca aspersa G. & B.

Empoasca nigra G. & B.

Exitianus exitiosus

Exitianus sp.

Hecalus viridis (Uhler)

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
USDA Forest Service Proceedings RMRS-P-38. 2005 119

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

Idiocerus sp.

Idiodonus geminatus

Norvellina vermiculata

Parabolocratus viridis Uhler

Texananus sp.

Xerophloea viridis (Uhler)

Delphacidae

Eurysa obesa Beamer

unident. spp. #1 to #3

Fulgoridae

Delphacodes pellucida (F.)

Delphacodes campestris VanDuzee

unident. sp. #1

Psillidae

Aphalara artemesiae Forster

Aphalara calthae (L.)

Aphalara loca Calderwood

Aphalara minutissima Crawford

Calophya triozoma Schwarz

Aphididae

Aphis gregalis Knowlton

Aphis ornata (Gillette & Palmer)

Brachycaudus helichrysi (Kaltenbach)

Durocapillata utahensis Knowlton

Forda marginata (Koch)

Microsiphoniella acophorum (Knowlton & Smith)

Myzus persicae (Sulzer)

Obtusicauda artemisicola

Pleotrichophorus pycnorhysus (Knowlton & Smith)

Pleotrichophorus utensus (Pack & Knowlton)

Uroleucon escalanti Knowlton

Zyxaphis canae (Williams)

Zyxaphis filifoliae Gillette & Palmer

Phylloxeridae

unident. sp. #1

Margarodidae

unident. sp. #1

Coccidae

unident sp. #1

Pseudococcidae

Phenacoccus sp.

THYSANOPTERA

Aeolothripidae

Aeolothrips auricestus Treherne

Aeolothrips fasciatus

Thripidae

Aptinothrips rufus (Gmelin)

Frankliniella occidentalis (Pergande)

Sericothrips sp.

Thrips tabaci

Oelolothripidae

Oelolothrips sp.

Phlaeothripidae

Leptothrips mali (Fitch)

unident. sp. #1

NEUROPTERA

Coniopterygidae

unident. sp. #1

Hemerobiidae

Hemerobius sp.

Kimminsia coloradensis (Banks)

Micromus variolus (Hagen)

Chrysopidae

Chrysopa coloradensis Banks

Chrysopa nigricornis

Eremochrysis punctinervis MacLachlan

Myrmeleontidae

Myrmeleon sp.

COLEOPTERA

Cicindelidae

Cicindela decemnotata Say

Cicindela purpurea

Carabidae

Agonum balesi Gray

Agonum placidum (Say)

Amara apricaria Paykull

Amara farcata LeConte

Amara impuncticollis Say

Amara laticollis LeConte

Amara littoralis

Amara musculus Say

Amara quenseli Schönherr

Apristus sp.

Axinopalpus biplagiatus (Dejean)

Bembidion immaculosum Hatch

Bembidion nebraskense LeConte

Bembidion obscurellum Motschulsky

Bembidion rupicola Kirby

Bembidion timidum LeConte

Bradycellus congener

Calleida viridis

Calosoma luxatum Say

Clivina fossor

Cymindis planipennis LeConte

Dicheirus piceus Menetries

Harpalus amputatus Say

Harpalus basilaris Kirby

Harpalus fraternus LeConte

Harpalus sp.

Lebia vittata (Fabricius)

Microlestes nigrinus (Mannerheim)

Philophuga viridis Dejean

Piosoma setosa LeConte

Pseudomorpha behrensi Horn

Pterostichus sp.

Haliplidae

spp. undetermined

Dytiscidae

Laccophilus decipiens LeConte

Gyrinidae

spp. undetermined

Ptiliidae

spp. undetermined

Leiodidae

Hydnobius sp.

Letodes grassa

Ptomophagus californicus (LeConte)

Silphidae

Nicrophorus hecate Bland

Nicrophorus guttulus Motschulsky

Staphylinidae

Acratona sp

Aleochara sp.

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
120 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Aleocharinae spp. #1, #2

Anotylus sp.

Astenus longiusculus

Bledius strenuus Casey

Bryoporus testaceus

Oxypoda sp.

Philonthus concinnus (Gravenhorst)

Philonthus cruentatus (Gmelin)

Platystethus americanus (Erichson)

Quedius sp.

Tachinus angustatus Horn

Tachyporus canadensis

Pselaphidae

Pilopius sp.

Eucnemidae

Analestes sp.

Hydrophilidae

Berosus fraternus LeConte

Berosus styliferus Horn

Cercyon quisquilius (L.)

Helophorus sp.

Sphaeridium scarabaeoides (L.)

Histeridae

Psiloscelis corrosa Casey

Saprinus lanei

Saprinus lugens Erichson

Saprinus oregonensis LeConte

Xerosaprinus acilinea (Marseul)

Xerosaprinus lubricus (LeConte)

Eucinetidae

unident. sp. #1

Scarabaeidae

Aphodius denticulatus Haldeman

Aphodius distinctus Muller

Aphodius fossor (L.)

Aphodius granarius (L.)

Aphodius hirsutus Brown

Aphodius militaris (?) LeConte

Aphodius vittatus Say

Bolbocerus obesus

Boreocanthon simplex (LeConte)

Cremastocheilus crinitus bifoviatus Van Dyke

Dichelonyx truncata LeConte

Dichelonyx sp. #2

Diplotaxis brevicollis

Diplotaxis obscura LeConte

Diplotaxis subangulata LeConte

Diplotaxis tenebrosa Fall

Glaresis canadensis Brown

Glaresis clypeata

Ochodaeus simplex LeConte

Paracotalpa granicollis (Haldeman)

Phyllophaga sociata

Serica anthracina LeConte

Serica barri Dawson

Trox sp.

Buprestidae

Acmaeodera immaculata Horn

Agrilus politus Say

Agrilus pubifrons Fisher

Agrilus walsinghami Crotch

Anthaxia retifer LeConte

Chrysobothris deleta LeConte

Chrysobothris horningi Barr

Chrysobothris idahoensis Barr

Chrysobothris texana LeConte

Heteroceridae

Lanternarius brunneus (Melsheimer)

Nannularia brunneata (Knull)

Elmidae

spp. undetermined

Elateridae

Aeolus dorsalis

Aeolus mellilus

Agriotella fusca Lane

Ampedus ursinus (Van Dyke)

Anchastus cinereipennis (Eschscholtz)

Cardiophorus spp. #1 to #5

Cardiophorus tumidicollis

Ctenicera noxia (Hyslop)

Ctenicera pruinina Horn

Ctenicera semivittata (Say)

Horistonotus pilosus Lanchester

Hypolithus bicolor (Eschscholtz)

Limonius (?) sp.

Cantharidae

Malthodes spp.

Dermestidae

Dermestes marmoratus Say

Bostrichidae

unident. sp. #1

Cleridae

Enoclerus acerbus Wolcott

Enoclerus barri Knull

Phyllobanus sp.

Trichodes ornatus Say

Melyridae

Amecocerus spp.

Attalus glabrellus Fall

Attalus morulus smithi Hopping

Attalus oregonensis Horn

Collops bipunctatus (Say)

Collops bridgeri Tanner

Collops hirtellus LeConte

Collops punctulatus LeConte

Dasytellus sp.

Hoppingiana nitida Hatch

Neodasytes testaceous

Trichochrous paisleyi

Dasytes cruralis

Dasytellus nigricornis (?) Bland

Nitidulidae

Brachypterolous pulicarius (L.)

Carpophilus pallipennis (Say)

Cryptophagidae

Caenoscelis ferruginea

Cryptophagus cellaris

Phalacridae

Olibrus rufipes LeConte

Phalacrus penicilatus Say

Coccinellidae

Brachyacantha dentipes socialis Casey

Brachyacantha ursina uteella Casey

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
USDA Forest Service Proceedings RMRS-P-38. 2005 121

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

Brumus serptentrionis Weise

Coccinella difficilis Crotch

Coccinella guttata Bennett

Coccinella septempunctata

Coccinella novemnotata degener Casey

Coccinella prolongata Crotch

Coccinella transversoguttata richardsoni Brown

Hippodamia apicalis Casey

Hippodamia convergens Guerin

Hippodamia glacialis lecontei Mulsant

Hippodamia quinquesignata Kirby

Hippodamia tredecimpunctata tibialis (Say)

Hyperaspidius hercules Belicek

Hyperaspidius vittigera LeConte

Hyperaspis lateralis montanica Casey

Hyperaspis postica LeConte

Nephus ornatus

Nephus sordidus (Horn)

Psyllophora vigintimaculata (Say)

Selvadius nunemacheri

Selvadius sp. #2

Scymnus ardelio Horn

Scymnus caurinus Horn

Scymnus marginicollis Mannerheim

Scymnus postpictus Casey

Latridiidae

Melanopthalma americana Mannerheim

Melanopthalma sp. #2

Melandryidae

Physicus sp.

unident. sp. #1

Mordellidae

Mordella atrata Melsheimer

Mordellistena aspersa (Melsheimer)

Mordellistena idahoensis Ray

Mordellistena sericans Fall

Tenebrionidae

Alaudes singularis Horn

Araeoschizus airmeti Tanner

Blapstinus barri Boddy

Blapstinus discolor

Blapstinus substriatus Champion

Coelocnemis punctatus LeConte

Coniontis obesa LeConte

Coniontis ovalis (Say)

Coniontis setosa Casey

Eleodes cordata Eschscholtz

Eleodes elongata

Eleodes extricata cognata Haldeman

Eleodes granulata

Eleodes hispilabris connexa LeConte

Eleodes humeralis

Eleodes nigrina LeConte

Eleodes novoverrucula Boddy

Eleodes obscura Say

Eleodes pilosa Horn

Eleodes rileyi Casey

Embaphion elongatum Horn

Eusattus muricatus

Helops californicus Mannerheim

Helops convexulus LeConte

Helops opacus LeConte

Melanstrus ater (LeConte)

Oxygonodera hispidula (Horn)

Sphaeriontis muricata (Casey)

Alleculidae

Mycetochara procera Casey

Meloidae

Epicauta immerita Walker

Epicauta normalis Werner

Epicauta piceiventris Maydell

Gnathium eremicola Macswain

Lytta vulnerata cooperi LeConte

Nemognatha lutea LeConte

Nemognatha scutellaris

Linsleya sphaericollis (Say)

Oedemeridae

Oxacis bicolor (LeConte)

Anthicidae

Anthicus cervinus LaFerte

Anthicus formicarius LaFerte

Anthicus hastatus Casey

Anthicus nanus

Notoxus serratus LeConte

Notoxus robustus

Notoxus sp. #2

Cerambycidae

Centrodera nevadica nevadica LeConte

Cortodera barri Linsley & Chemsak

Crossidius ater LeConte

Crossidius coralinus LeConte

Crossidius hirtipes allgewahri LeConte

Crossidius punctatus LeConte

Judolia gaurotoides Casey

Mecas bicallosa Martin

Megacheuma brevipennis (LeConte)

Megasemum asperum (LeConte)

Micas bicallosa

Prionus californicus Motschulsky

Bruchidae

Acanthoscelides pauperculus (LeConte)

Acanthoscelides pudis Fall

Acanthoscelides sp. #3

Chrysomelidae

Altica plicipennis

Anisostena californica Van Dyke

Brachycoryna montana (Horn)

Chaetocnema sp.

Crepidodera nana (Say)

Cryptocephalus spurcus LeConte

Dibolia borealis Chevrolat

Disonycha latifrons Shaeffer

Exema conspersa (Mannerheim)

Glyptina atriventris Horn

Glyptoscelis artemisiae Blake

Longistarsis oregonensis

Monoxia consputa LeConte

Monoxia pallida Blake

Monoxia puberula Blake

Pachybrachis caelatus LeConte

Pachybrachis jacobyi Bowditch

Phyllotreta albionica LeConte

Phyllotreta oregonensis

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
122 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Pseudoluporus longulus LeConte

Psylliodes punctulata

Pyrrhalta luteola Muller

Pyrrhalta nymphaeae (L.)

Saxinis saucia LeConte

Scelolyperus nigrovirescens (Fall)

Stenopodius flavidus

Stenopodius vanduzeei Blaisdell

Systena blanda Melsheimer

Trirhabda nitidicollis LeConte

Trirhabda sp. #2

Curculionidae

Acmaegenius granicollis Van Dyke

Anthonomus tenuis Fall

Anthonomus squamosus

Apion sordidum Smith

Brachyrhinus ovatus (L.)

Cercopedeis artemisiae (Pierce)

Ceutorhinchus addunctus Dietz

Ceutorhinchus bakeri Hatch

Ceutorhinchus disturbans Dietz

Ceutorhinchus sp. #4

Cleonidius poricollis

Cleonus kirbyi Casey

Cleonus quadrilineatus (Chevrolat)

Cosmobarius americana Casey

Dinocleus denticollis Casey

Dyslobus alternatus Horn

Epimechus mimicus Dietz

Gyrotus sinuatus Hatch

Miloderoides maculatus Van Dyke

Myrmex vittatus (Horn)

Ophryastes latirostris LeConte

Sitona hispidula (Fabricius)

Sitona lineata

Smicronyx abnormis

Sphenophorus gentilis LeConte

Tostates cinerascens

Trachyphloeni sp.

Tychius tectus LeConte

Tychius mixtus (?) Hatch

Anthribidae

Trigonorhinus sp.

Anobiidae

Ptinis villiger

Xyletynus fucatus LeConte

SIPHONAPTERA

Ctenophthalmidae

Rectofrontia fraterna

Hystrichopsyllidae

Anomiopsyllus amphibolus

Callistopsyllus terinus

Catallagia decipiens

Epiledia stanfordi

Epiledia wenmanni

Hystrichopsylla occidentalis

Megarthroglossus divisus

Meringis hubbardi

Meringis parkeri

Phalacropsylla allos

Phalacropsylla paradisea

Rhadinopsylla sectilis

Stenistomera alpina

Stenistomera macrodactyla

Ceratophyllidae

Foxella ignota

Malaraeus bitterrootensis

Malaraeus euphorbi

Malaraeus telchinum

Megabothris abantis

Megabothris obscurus

Monopsyllus eumolpi

Monosyllus exilis

Monopsyllus wagneri

Opisocrostis labis

Opisocrostis tuberculatus

Opisodasys keeni

Orchopeas leucopus

Orchopeas sexdentatus

Thrassis bacchi

Thrassis francisi

Thrassis howelli

Thrassis pandorae

Leptopsyllidae

Amphipsylla siberica

Odontopsyllus dentatus

Peromyscopsylla hesperomys

Pulicidae

Cediopsylla inaequalis

Pulex irritans

DIPTERA

Tipulidae

spp. undetermined

Bibionidae

Bibio albipennis

Bibiodes sp.

Mycetophilidae

Bodetina (?) sp.

Fungivora sp.

Megalopelmna (?) sp.

unident. spp. #1, #2

Sciaridae

unident. spp. #1 to #4

Cecidomyidae

Rhopalomyia sp.

unident. spp. #1 to #4

Psychodoidea

spp. undetermined

Scatopsidae

Scatopse fusipes Meigan

Dixidae

unident. sp. #1

Cuculidae

Aedes dorsalis Meigan

Simuliidae

Cnephia munus D. & S.

Simulium bivittatum Malloch

Simulium venator D. & S.

Simulium venustum (?) Say

Simulium vittatum Zetterstedt

Simulium spp. #5, #6

Ceratopogonidae

Culicoides sp.

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
USDA Forest Service Proceedings RMRS-P-38. 2005 123

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

Dasyhelea spp. #1, #2

Forcipomyia brevipennis Macquaert

Forcipomyia sp. #2

Chironomidae

Chironomus spp. #1 to #3

Stratiomyiidae

Nemotelus canadensis Loew

Therevidae

Ozodiceromya sp.

Psilocephala spp. #1 to #3

Thereva pseudoculata Cole

Thereva semitaria

Scenopinidae

Scenopinus sp.

Asilidae

Asilus occidentalis Hine

Asilus mesae

Efferia benedicta (Bromley)

Efferia subcuprea (Schaffer)

Heteropogon senilis Bigot

Holopogon seniculus Loew

Lasiopogon sp.

Leptogaster fornicata Martin

Megaphorus martinorum

Ospriocerus abdominalis (Say)

Stenopogon inquinatus Loew

Stenopogon neglectus

Bombyliidae

Aphoebantus mormon Melander

Apolysis arenicola

Exoprosopa caliptera (Say)

Geron sp.

Lepidanthrax inauratus (Coquillet)

Lordotus apicalis (Coquillet)

Mythicomyia armata

Mythicomyia atra Cresson

Mythicomyia rileyi Coquillet

Mythicomyia spp. #4 to #9

Oligodranes acrostichalis Melander

Oligodranes quinquenotatus

Phthirla sulphurea

Prorates arctos

Prorates claripennis Melander

Thyridantrax andrewsi

Toxophora virgata Osten Sacken

Toxophora sp.

Villa molitor Loew

Empididae

Drapetis spp. #1 to #3

Platypalpus sp.

Phoridae

Megaselia spp. #1 to #4

Syrphidae

Eupeodes volucris Osten Sacken

Scaeva pyrasti (L.)

Sphaerophoria philanthus

Pipunculidae

Tomosvaryella sp.

Conopidae

Physocephala texana (Willinston)

Thecophora propinqua (Adams)

Zodion fulvifrons Say

Micropezidae

unident. spp. #1, #2

Psilidae

Psila dimidiata Loew

Otitidae

Euxesta (?) sp.

Oedopa capito Loew

Physiphora demandata (F.)

Tritoxa cuneata

Tritoxa pollinosa Cole

Tephritidae

Aciurina ferruginea Doane

Aciurina luteana Cresson

Aciurina trixa Cresson

Aciurina spp. #4, #5

Euaresta tapetis Coquillet

Eutreta diana

Eutreta oregona Curran

Neaspilota sp.

Neotephrites finalis (Loew)

Oxyna palpalis Coquillet

Paroxyna clathrata (Loew)

Paroxyna corpulenta Cresson

Paroxyna minima Doane

Paroxyna steyskali Novatny

Procecidochares minuta Snow

Procecidochares sp. #2

Tephritis araneosa (Coquillet)

Trupanea bisetosa Coquillet

Trupanea jonesi Curran

Trupanea nigricornis Coquillet

Milichiidae

Leptometopa halteralis Coquillet

Madiza glabra (F.)

Neophyllomyza quadricornis Melander

Pholeomyia indecora Loew

Dryomyzidae

unident. sp. #1

Sciomyzidae

unident. spp. #1, #2

Sepsidae

Saltella scutellaris Fallen

Saltella sp. #2

Sepsis biflexuosa

Sepsis neocynipsea Melander & Spuler

Sepsis punctum F.

Lauxaniidae

Camptoprosopella borealis Shewell

Chamaemyiidae

Leucopis americana Malloch

Leucopis flavicornis Aldrich

Pseudodinea nitens Melander & Spuler

unident. spp. #1 to #3

Heleomyzidae

Heleomyza sp.

Pseudoleria sp.

Trixoscelidae

Trixoscelis sp.

Sphaeroceridae

Leptocera sp.

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
124 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Drosophilidae

unident. sp. #1

Ephydridae

Hydrellia griseola (Fallen)

Leptopsilota sp.

Mosillus bidentatus Cresson

Phylligria debilis Loew

Psilopa sp.

Chloropidae

Chlorops rubicundus Adams

Chlorops sordidellus

Conioscinella sp.

Goniopsita oophaga

Hippelates particeps Becker

Hippelates pusio Loew

Incertella sp.

Meromyza pratorum Meigan

Meromyza saltatrix (L.)

Neoneura flavifacies Collin

Neoneura polita Sabrosky

Olcella punctifrons Becker

Oscinella frit (L.)

Oscinella inserta Becker

Siphonella sp.

Thaumatomyia annulata (Walker)

Thaumatomyia appropinqua (Adams)

Thaumatomyia glabra (Meigan)

Anthomyiidae

Calythea micropteryx Thomson

Hydrophoria brunneifrons (Zetterstedt)

Hydrophoria divisa (Meigan)

Hylemyia spp. #1 to #3

Leucophora sp.

Pegomya sp.

Scatophaga stercoraria (L.)

Scatophaga sp. #2

Agromyzidae

Agromyza pusilla Meigan

Ceradontha dorsalis Loew

Melanagromyza sp.

Phytobia sp.

unident sp. #5

Scatophagidae

unident. sp. #1

Muscidae

Fannia sp.

Helina duplicata (Meigan)

Helina multisetosa

Helina troene Walker

Helina spp. #3, #4

Lasipcs septentrionalis Stein

Musca domestica L.

Muscina stabulans Fallen

Orthellia caesarion (Meigan)

Quadrolaria laetifica Robineau-Desvoidy

Schoenomyza dorsalis Loew

Calliphoridae

Calliphora lilae Walker

Phormia regina (Meigan)

Protophormia terrenovae (Macquaert)

Sarcophagidae

Hilarella hilarella (Zetterstedt)

Robineauella sp.

Sarcophaga spp. #1 to #3

Senotainia trilineata (Wulp)

Senotainia vigilans Allen

Sphenometopa tergata Coquillet

Taxigramma heteroneura (Meigan)

Cuterebridae

Cuterebra jellisoni

Tachinidae

Acemya tibialis Coquillet

Anthrycia cineria (Coquillet)

Anthrycia sp.

Bennettia compta (Fallen)

Blondelia (?) sp.

Catagoniops facialis (Coquillet)

Dinera grisescens (Fallen)

Euphorcera sp.

Exorista mella Walker

Gonia albagenae Morrison

Hyalomyia aldrichii Townsend

Lespesia archippivora (Riley)

Leucostoma simplex Fallen

Lydella radicis Townsend

Microchaetina valida Townsend

Norwickia latifacies Tothill

Norwickia latigena Tothill

Norwickia robinsoni

Paradidyma simulans Townsend

Paradidyma singularia Townsend

Patellea sp.

Peleteria malleola Bigot

Periscopsia cinerosa Coquillet

Periscepsia helymus (Walker)

Promasiphia (?) sp.

Siphosturmia maltana Reinhardt

Sitophaga sp.

Spallanzania sp.

Spathidexia reinhardti Arnaud

Stomatomyia parvipalpis Wulp

Voria ruralis Fallen

TRICHOPTERA

Leptoceridae

spp. undetermined

LEPIDOPTERA

Micropterigidae

unident. spp. #1, #2

Lyonetiidae

Bucculatrix seorsa

Bucculatrix tridenticola Braun

Coleophoridae

Coleophora sp.

Gelichiidae

Aroga websteri Clarke

Chionodes sp.

Tortricidae

Eucosma sp.

Phaneta salmicolorana

Phaneta setonana

Synnoma lynosyrana Walsingham

Plutellidae

Plutella maculipennis Curtis

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
USDA Forest Service Proceedings RMRS-P-38. 2005 125

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

Pterophoridae

unident. sp. #1

Microlepidoptera spp. #1 to #9

Pyralidae

Loxostege commixtalis

Omnaptopteryx occellea Hanson

Hesperiidae

Hesperia harpalus (Edwards)

Hesperia juba (Scuder)

Papilionidae

Papilio sp.

Pieridae

Colias interior

Euchloe ausonides Lucas

Pieris beckerii

Pieris protodice Biusdyvak

Lycaenidae

Lycaena helloides Boisduval

Lycaena rubidus

Plebejus melissa (Edwards)

Nymphalidae

Euphydryas anicia (Doubleday)

Speyeria callippe nevadensis (Edwards)

Vanessa cardui (L.)

Satyridae

Cericyonis oetus oetus (Boisduval)

Saturniidae

Hemileuca hera hera Harris

Sphingidae

Hyles lineata (Fabricius)

Proserpinus clarkiae Boisduval

Arctiidae

Apantesis retilineata

Arctia caja (L.)

Leparctia sp.

Geometridae

Chlorosea sp.

Glaucina nephos

Pero modesto

Plataea linearia

Procherodes amplicineraria Pearson

Semiothisa nubiculata

Synaxis formosa

Noctuidae

Abagrotis nefascia Smith

Apamea occidens (Grote)

Aseptis characta Grote

Autographa californica Speyer

Copablepharon canariana McDonald

Cucullia arizona

Dicestra crotchi

Drasteria mirifica Hy. Edwards

Euxoa auxiliaris Grote

Euxoa costata idahoensis Grote

Euxoa pluralis Grote

Faronta diffusa (Walker)

Heliothis belladonna

Lacinipolia sp.

Rhynchagrotis exertistigma Morrison

Synedoida sp.

HYMENOPTERA

Cephidae

Cephus cinctus Norton

Ceraphronidae

Ceraphron sp.

unident. spp. #1, #2

Braconidae

Agathis californica (?) Rhower

Agathis gibbosa Muesebeck

Apanteles sp. #2

Apanteles yakutatensis (Say)

Aphidius sp.

Bracon hyslopi (Viereck)

Bracon nuperus (?) Cresson

Bracon spp. #3 to #5

Chelonus spp. #1, #2

Chorebus sp.

Diaeretiella spp. #1, #2

Hormius sp.

Macrocentrus ancylivorus Rohwer

Macrocentrus sp. #2

Meteorus leviventris Wesmail

Microplitis plutellae Muesebeck

Microctonus pusillae Muesebeck

Opius (?) sp.

Orgilus sp.

Rogas sp.

Tetrasphaeropyx sp.

Vipio (?) sp.

unident. spp. #1 to #8

Ichneumonidae

Amblyteles spp. #1, #2

Anomalon ejuncidum Say

Anomalon sp.

Banchus nubilus Townes

Campoplex sp.

Chorineaus sp.

Conoblasta sp.

Cryptus asymmetricus Pratt

Cryptus sp. #2

Diadegma plutellae (Viereck)

Diadegma sp. #1

Diplazon laetitorius (F.)

Exetastes sp.

Gelis sp.

Gnypetonoipla (?) sp.

Mesostenus gracilis Cresson

Ophion abnormalis Felt

Ophion purgatus Say

Pseudamblyteles superbus (Provander)

Pseudamblyteles kocheli Swezey

unident. spp. #1 to #8

Mymaridae

Gonatocerus sp.

unident. sp. #1

Trichogrammatidae

unident. sp. #1

Eulophidae

Achrysocharella sp.

Euderus sp.

Chrysocharis sp.

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
126 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Cirrospilus sp.

Diglyphus spp. #1, #2

Elachertus sp.

Entedon sp.

Necremnus duplicatus Gahan

Necremnus sp. #2

Pediobius utahensis (Crawford)

Tetrastichus coerulescens Ashmead

Tetrastichus spp. #2 to #10

Zagrammosoma nigrolineatum Crawford

Zagrammosoma sp. #2

unident. spp. #1 to #13

Elasmidae

Elasmus sp.

Aphelinidae

unident. sp. #1

Encyrtidae

Homalotylus (?) sp.

Oencyrtus sp.

unident. spp. #1 to #15

Eupelmidae

Eupelmus allynii French

Eupelmus sp. #2

Calosota sp.

unident. spp. #1 to #3

Torymidae

Torymus coloradensis

unident. spp. #1 to #11

Pteromalidae

Asaphes sp.

unident. spp. #1 to #37

Eutrichosomatidae

Eutrichosoma mirabile Ashmead

Perilampidae

Perilampus chrysopae Crawford

Perilampus hyalinus Say

Perilampus similis Crawford

Perilampus sp. #4

Eurytomidae

Eurytoma spp. #1 to #5

Harmolita spp. #1, #2

Rileya cecidomyiae Ashmead

Tetramesa elymophaga (Phillips)

Tetramesa sp. #2

unident. spp. #1 to #4

Chalcididae

Haltichella sp.

Spilochalcis albifrons Welsh

Spilochalcis ignoides (?) Kirby

Spilochalcis leptis Burks

Spilochalcis side Walker

unident. sp. #1

Eucoilidae

unident. sp. #1

Figitidae

Melanips coxalis

Trischiza sp.

Cynipidae

Periclistus sp.

Proctotrupidae

Proctotrupes florissantensis Kiefer

Diapriidae

unident. spp. #1, #2

Scelionidae

Gaeus sp.

Gryon sp.

Telenomus spp. #1 to #3

Trissolcus utahensis

Platygasteridae

Platygaster rohweri

Platygaster utahensis

Platygaster spp. #2, #3

Synopeas spp. #1 to #3

Inostemma sp.

unident. sp. #1

Chrysididae

Ceratochrysis perpulchra (Cresson)

Ceratochrysis trachyplenia R. Bohart

Chrysis canadensis Buysson

Chrysis coerulans (F.)

Chrysis coloradica

Chrysis dorsalis (?) Aaron

Chrysis vagabunda

Chrysis sp. #6

Chrysura densa (Cresson)

Cleptes purpuratus

Hedychridium carrilloi R. Bohart & Brumley

Omalus aeneus

Bethylidae

unident. spp. #1, #2

Dryinidae

unident. sp. #1

Sphecidae

Ammophila spp. #1 to #3

Ammoplanops (?) sp. #1

Ancistromma sp.

Astata bakeri Parker

Belomicrus sp.

Bembix amoena Handlirsch

Bembix spinolae Lepeletier

Cerceris minax Mickel

Cerceris nigrescans Simth

Dienoplus sp.

Diodontus spp. #1, #2

Dryudella immigrans William

Dryudella sp. #2

Ectemniis dilectus Cresson

Ectemniis sp. #2

Eucerceris sp.

Glenostictia megacera J. Parker

Gorytes sp.

Mimesa sp.

Miscophus (Nitelopterus) sp.

Nyson sp.

Oxybelus sp.

Philanthus multimaculatus Cameron

Podalonia spp. #1, #2

Prionyx canadensis (Provancher)

Solierella spp. #1 to #3

Sphex ichneumoneus (L.)

Steniolia elegans J. Parker

Stictella megacera Parker

Tachysphex irregularis

(con.)

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name
USDA Forest Service Proceedings RMRS-P-38. 2005 127

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

Table 1—(Cont.)

ORDER ORDER

Family Family

Scientific name Scientific name

Tachysphex tarsatus (Say)

Tachysphex williamsi

Melittidae

unident. sp. #1

Colletidae

Colletes dissoptus Timberlake

Colletes fulgidus Swank

Colletes lutzi Timberlake

Halictidae

Agapostemon texanus Cresson

Dialictus spp. #1, #2

Evylaeus sp.

Halictus farinosus Smith

Halictus ligatus

Halictus tripartitus Cockerell

Sphecodes arvensiformis (?) Cockerell

Sphecodes sp.

Andrenidae

Andrena prunorum Cockerell

Descurainia richardsoni

Perdita spp. #1, #2

Megachilidae

Anthidium emarginatum (Say)

Anthidium placitum Cresson

Anthidium utahense Swenk

Ashmeadiella gillettei Titus

Ashmeadiella opuntiae Cockerell

Dianthidium pudicum decorum Timberlake

Dioxys pomonae Cockerell

Hoplitis producta Michner

Megachile laurita Mitchell

Megachile onobrychidis Cockerell

Megachile parallela Smith

Osmia integra

Stelis sp.

Anthophoridae

Anthophora exigua Cresson

Anthophora ursina Cresson

Ceratina pacifica Cresson

Didadasia enavata Cresson

Epeolus minimus Robertson

Melissodes bimatris (?) LaBerge

Nomada articulata Smith

Nomada suavis Cresson

Synhalonia spp. #1, #2

Tetralonia fulvitarsis Cresson

Triepeolus helianthi (?) Robertson

Apidae

Apis mellifera (L.)

Bombus fervidus (F.)

Bombus huntii Greene

Tiphiidae

Brachycistis spp. #1, #2

Sapygidae

Sapyga pumila Cresson

Tiphia sp.

Mutillidae

Chyphotes sp.

Sphaerophthalma unicolor

Sphaerophthalma spp. #2 to #4

Scoliidae

Campsoscolia alcione Banks

Formicidaed

Camponotus hyatti Emery

Camponotus vicinus May

Camponotus sp. #2

Ephebomyrmex sp.

Formica ciliata Mayr

Formica cinerea canadensis Santschi

Formica fusca L.

Formica gynocrates Snelling and Buren

Formica haemorrhoidalis Emery

Formica hewitti Wheeler

Formica lasioides Emery

Formica laviceps Creighton

Formica manni Wheeler

Formica montana

Formica neogagates Emery

Formica obscuriventris

Formica obtusopilosa Emery

Formica oreas comptula Wheeler

Formica rufa (L.)

Formica subnuda

Formica subpolita Mayr

Formica whymperi

Formicoxenus diversipilosus

Formicoxenus hirticornis

Lasius alienus

Lasius crypticus Wilson

Leptothorax andrei

Leptothorax nevadensis Wheeler

Liometopum luctuosum W. M. Wheeler

Manica mutica

Monomorium minimum

Myrmecocystus mojave

Myrmecocystus testaceus Emery

Myrmica americana Weber

Myrmica lobicornis Emery

Pheidole californica

Pogonomyrmex occidentalis (Cresson)

Pogonomyrmex owyheei Cole

Pogonomyrmex salinus Olsen

Solenopsis molesta Say

Stenamma sp.

Tapinoma sessile Say

Veromessor lobognathus (Andrews)

Pompilidae

Ageniella spp. #1, #2 (?)

Anoplius insolens

Anoplius tenebrosus (Cresson)

Anoplius sp. #3

Aporinellus completus Banks

Aporinellus fasciatus (Smith)

Ceropales sp.

Episyron snowi (Viereck)

Evagetes padrinus (?) (Viereck)

Evagetes parvus

Evagetes sp. #3

Pompilus angularis (Banks)

Vespidae

Ancistrocerus spp. #1, #2

Euodynerus annulatus (Say)

Euodynerus sp. #2

(con.)
128 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Pterocheilus fasciatus Say

Pterocheilus pediculatus

Pterocheilus provancheri

Stenodynerus noticeps (?)

Stenodynerus sp. #2

a Order and Family names are according to Borror and others (1992).

b Spellings are according to Horning and Barr (1970), Haws and others (1988),

and Arnett (2000). Authors are provided only in cases where they were available

from literature cited in the reference list. Authors shown in parentheses indicate

the generic name has changed since the species was originally identified (Borror

and others 1992). Abbreviations: sp. = single species (Borror and others 1992),

spp. = multiple species (Borror and others 1992), unident. = unidentified species

or morphospecies, undetermined = specimens were not identified past family

level, (?) = questionable identification noted by the researcher. c

 Subsequently identified as O. annulata. d Additional taxa and clarifications pending (Clark and Blom, in preparation).

Table 1—(Cont.)

ORDERa

Familya

Scientific nameb

with specific hostplants or particular insect groups. In most

cases, only a few dozen species were collected and identified.

The investigation by Bohart and Knowlton (1977), in which

over 800 species were identified, constitutes the most extensive single inventory conducted at the INL, followed by the

study by Karr and Kimberling (2003) and the multiple-year

investigation conducted by Stafford (1983, 1987) and Stafford

and Johnson (1986).

In comparison, a 3-year survey at nearby Craters of the

Moon National Monument (CMNM) (Horning and Barr

1970) resulted in the identification of nearly 2,100 species,

representing 248 families and 1,144 genera in 19 orders.

Totals for the suborder Raphidoidea were reported separately in the summary of COM insect orders by Horning and

Barr (1970); for the INL list, species Raphidoidea are included in totals reported for Neuroptera (see table 2).

Although about 860 more insects were documented in the

CMNM survey, only 157 of the 212 families, 396 of 747

genera, and 305 of 1,241 species identified at the INL have

also been identified at CMNM. Insects in disturbed habitats

at the INL have been investigated (Karr and Kimberling

2003; Wenninger 2001), but studies have been primarily in

native sagebrush and grassland. While similar communities

were included in the CMNM survey, the larger inventory

likely reflects investigation of a wider variety of habitat

types (Horning and Barr 1970) and greater nighttime collecting efforts.

Insect research at the INL has been focused on terrestrial

species. However, a few aquatic insect families have been

documented in and around industrial waste ponds (Cieminski

and Flake 1995; Millard and others 1978). Additional aquatic

insects have been collected from sections of the Big Lost

River, but specimens have not been sorted and identified (R.

C. Rope, personal communication).

In recent years, large wildfires at the INL have destroyed

thousands of acres of sagebrush habitat, including several

former research sites (fig. 1). The response of insects to fire

has been studied at the INL (Stafford 1983, 1987; Winter

1994), but as the need for restoration of sagebrush habitats

damaged by fire and other disturbance increases, a greater

understanding of insect life cycles, population dynamics,

and changes in species composition over time will be required. This baseline list of insects can be used to identify

groups that have not been well characterized and will help

focus further investigation of insect ecology and function in

natural and restored sagebrush systems.

Acknowledgments ______________

I thank Dr. William H. Clark and Dr. Nancy Huntly for

comments and suggestions that improved this manuscript.

Special thanks are extended to Dr. James R. Karr and Dr.

Diana N. Kimberling for graciously providing collection data

from their INL study. Kenneth S. Moor gave helpful comments on early versions of the manuscript and Dr. Randall

C. Morris assisted with documenting the INL insect voucher

collection. I am also grateful to Dr. Nancy Shaw and Loa

Collins for their help and diligent work to make this publication possible.

References _____________________

Allred, D. M. 1968a. Fleas of the National Reactor Testing Station.

Great Basin Naturalist. 28: 73–87.

Allred, D. M. 1968b. Ticks of the National Reactor Testing Station.

Brigham Young University Scientific Bulletin, Biological Series.

10: 1–29.

Allred, D. M. 1969. Spiders of the National Reactor Testing Station.

Great Basin Naturalist. 29: 105–108.

Allred, D. M. 1970. Mites and lice of the National Reactor Testing

Station. Brigham Young University Scientific Bulletin, Biological Series. 12: 1–17.

Allred, D. M. 1973. Scorpions of the National Reactor Testing

Station, Idaho. Great Basin Naturalist. 33: 251–254.

Allred, D. M.; Cole, A. C., Jr. 1971. Ants of the National Reactor

Testing Station. Great Basin Naturalist. 31: 237–242.

Allred, D. M.; Muma, M. H. 1971. Solpugids of the National Reactor

Testing Station, Idaho. Great Basin Naturalist. 31: 164–168.

Anderson, J. E. 1999. The Idaho National Engineering and Environmental Laboratory: an ecological treasure of the Upper Snake

River Plain. Rangelands. 21: 1–17.

Anderson, J. E.; Patrick, S. M. 2000. Fire ecology of the INL. In: Luft,

A. A.; Warren, R. W., eds. 2000. Annual Technical Report to DOEID: Calendar year 1999. ESRF-037. Idaho Falls, ID: Environmental Science and Research Foundation: 91–93.

Anderson, J. E.; Shumar, M. L. 1989. Guidelines for revegetation of

disturbed sites at the Idaho National Engineering Laboratory.

DOE/ID-12114. Idaho Falls, ID: U.S. Department of Energy,

Radiological and Environmental Sciences Laboratory: 6–12.

Arnett, R. H., Jr. 2000. American insects. Boca Raton, FL: CRC

Press. 1,003 p.

Blew, R. D. 2000. Planting and irrigating effects on vegetation

recovery from fire. In: Luft, A. A.; Warren, R. W., eds. 2000.

Annual Technical Report to DOE-ID: Calendar year 1999. ESRF037. Idaho Falls, ID: Environmental Science and Research Foundation: 89–90.

Blom, P. E., Clark, W. H.; Johnson, J. B. 1991. Colony densities of

the seed harvesting ant Pogonomyrmex salinus (Hymenoptera:

Formicidae) in seven plant communities on the Idaho National

Engineering Laboratory. Journal of the Idaho Academy of Science. 27: 28–36.

Bohart, G. N.; Knowlton, G. E. 1977. Invertebrates. In: Final

environmental impact statement: Safety Research Experiment

Facilities, Idaho National Engineering Laboratory, Idaho. ERDA1552. Idaho Falls, ID: Argonne National Laboratory: D47–D58.

Borror, D. J.; Triplehorn, C. A.; Johnson, N. F. 1992. An introduction

to the study of insects. Orlando, FL: Harcourt Brace & Company.

875 p.
USDA Forest Service Proceedings RMRS-P-38. 2005 129

Insects of the Idaho National Laboratory: A Compilation and Review Hampton

year 1999. ESRF-037. Idaho Falls, ID: Environmental Science

and Research Foundation, Inc.: 64–66.

Clark, W. H.; Blom, P. E. [In preparation]. Annotated checklist of

the ants (Hymenoptera: Formicidae) on the Idaho National Engineering and Environmental Laboratory.

Department of Energy. 1985. Department of Energy, National

Environmental Research Parks. DOE/ER-246. Washington, DC:

U.S. Department of Energy, Office of Energy Research. 48 p.

Entwistle, P. G.; DeBolt, A. M.; Kaltenecker, J. H.; Steenhof, K.

2000. Proceedings: sagebrush steppe ecosystems symposium.

BLM/ID/PT-001001+1150. Boise, ID: U.S. Department of the

Interior, Bureau of Land Management. 145 p.

Haws, B. A.; Roe, A. H.; Nelson, D. L. 1988. Index to information on

insects associated with western wildland shrubs. Gen. Tech. Rep.

GTR-INT-248. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 296 p.

Horning, D. S., Jr.; Barr, W. F. 1970. Insects of Craters of the Moon

National Monument, Idaho. Misc. Ser. 8. Moscow: University of

Idaho, College of Agriculture. 118 p.

Jansen, A. 1997. Terrestrial invertebrate community structure as

an indicator of the success of a tropical rainforest restoration

project. Restoration Ecology. 5: 115–124.

Bromenshenk, J. J. 1987. Regional monitoring of pollutants with

honey bees. In: Wise, S. A.; Zeiser, R.; Goldstein, G. M., eds.

Progress in environmental specimen banking. National Bureau

of Standards Special Publ. 740. Springfield, VA: National Technical Information Service: 156–170.

Cieminski, K. L.; Flake, L. D. 1995. Invertebrate fauna of wastewater ponds in southeastern Idaho. Great Basin Naturalist. 55:

105–116.

Clark, W. H.; Blom, P. E. 1988. Observations on the relationship

between ants (Hymenoptera: Formicidae: Myrmicinae, Dorylinae)

and Araeoschizus (Coleoptera: Tenebrionidae). Journal of the

Idaho Academy of Science. 24: 34–37.

Clark, W. H.; Blom, P. E. 1991. Observation of ants (Hymenoptera:

Formicidae: Myrmicinae, Formicinae, Dolichoderinae) utilizing

carrion. Southwest Naturalist. 36: 140–142.

Clark, W. H.; Blom, P. E. 1992. Notes on spider (Theridiidae: Salticidae)

predation of the harvester ant, Pogonomyrmex salinus Olson

(Hymenoptera: Formicidae: Myrmicinae), and a possible parasitoid fly (Chloropidae). Great Basin Naturalist. 52: 385–386.

Clark, W. H.; Blom, P. E. 1999. Diversity of the ant fauna at the INL:

implications for waste management. In: Luft, A. A.; Warren, R.

W., eds. 2000. Annual Technical Report to DOE-ID: Calendar

Figure 1—Sampling locations for major insect studies at the INL. NOTE: the INEEL is now the Idaho National Laboratory (INL);

the TRA is now the Reactor Technology Complex (RTC); and the ANL-W is now the Materials and Fuels Complex (MFC).
130 USDA Forest Service Proceedings RMRS-P-38. 2005

Hampton Insects of the Idaho National Laboratory: A Compilation and Review

Johnson, J. B.; Stafford, M. P. 1986. Adult Noctuidae feeding on

aphid honeydew, and a discussion of honeydew feeding by adult

Lepidoptera. Journal of the Lepidopterists’ Society. 39: 321–327.

Karr, J. R., Kimberling, D. L. 2003. A terrestrial arthropod index of

biological integrity for shrub-steppe landscapes. Northwest Science. 77: 202–213.

Kimberling, D. L.; Karr, J. R., Fore, L. S. 2001. Measuring human

disturbance using terrestrial invertebrates in the shrub-steppe of

eastern Washington (USA). Ecological Indicators. 1: 63–81.

Knick, S. T. 1999. Requiem for a sagebrush ecosystem? Northwest

Science. 73: 53–57.

Knick, S. T.; Rotenberry, J. T. 1997. Landscape characteristics of

disturbed shrubsteppe habitats in southwestern Idaho (U.S.A.).

Landscape Ecology. 12: 287–297.

Merickel, F. W.; Clark, W. H. 1994. Tetramorium caespitum(Linnaeus)

and Liometopum luctuosum (W. M. Wheeler) (Hymenoptera:

Formicidae): New State records for Idaho and Oregon, with notes

on their natural history. Pan-Pacific Entomologist. 70: 148–158.

Millard, J. B.; Whicker, F. W.; Markham, O. D. 1978. Gamma

emitting radionuclides of the Test Reactor Area leaching ponds.

In: Markham, O. D., ed. Ecological studies on the Idaho National

Engineering Laboratory Site, 1978 Progress Report. IDO-12087.

Idaho Falls, ID: U.S. Department of Energy, Idaho Operations

Office: 28–46.

Stafford, M. P. 1983. Surface-dwelling Coleoptera inhabiting sagebrush communities in southeastern Idaho. Moscow, ID: University of Idaho. 43 p. Thesis.

Stafford, M. P. 1987. Insect interactions with four species of sagebrush (Artemisia) in a southeastern Idaho high desert rangeland.

Moscow, ID: University of Idaho. 81 p. Dissertation.

Stafford, M. P.; Barr, W. F.; Johnson, J. B. 1986. Coleoptera of the

Idaho National Engineering Laboratory Site: an annotated checklist. Great Basin Naturalist. 46: 287–293.

Table 2—Summary of insect inventories for the Idaho National Laboratory and Craters

of the Moon National Monument.

 Families Genera Species

 Order INLa CMNMb INL CMNM INL CMNM

Collembolac 4 5 _6 _ 6

Thysanura _ 2 _ 2 _ 2

Ephemeropterac 2 4 _4 _ 4

Odonata 3 4 3 6 3 10

Isoptera 1 1 1 1 1 1

Plecoptera _ 4 _ 4 _ 5

Dermaptera _ 1 _ 1 _ 1

Psocoptera 1 2 1 3 1 3

Phthiraptera 4 _ 9 _ 20 _

Orthoptera 4 4 8 14 11 23

Hemiptera 17 20 63 79 79 115

Homoptera 13 14 49 56 65 79

Thysanoptera 4 3 7 8 9 10

Neuroptera 5 5 9 14 10 24

Coleoptera 42 44 187 198 297 324

Strepsiptera _ 1 _ 1 _ 1

Siphonaptera 4 _ 24 _ 38 _

Diptera 46 50 159 286 238 521

Trichopterac 1 6 _ 10 _ 12

Lepidoptera 18 35 49 140 66 218

Hymenoptera 43 43 178 311 403 705

Totals 212 248 747 1,144 1,241 2,064

a Idaho National Laboratory.

b

 Craters of the Moon National Monument (totals from Horning and Barr 1970).

c INL specimens were not identified past family level for these orders.

Stafford, M. P.; Johnson, J. B. 1986. Phytophagous insects of green

rabbitbrush in southeastern Idaho. In: McArthur, E. D; Welch,

B. L., comps. Proceedings—symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9–13; Provo, UT. Gen. Tech.

Rep. GTR-INT-200. Ogden, UT: U.S. Department of Agriculture,

Forest Service, Intermountain Forest and Range Experiment

Station: 211–214.

Veith, R. D. 1983. Dispersal, relative density, and speed of carrion

location of the burying beetle, Nicrophorus hecate Bland (Coleoptera: Silphidae), in southeastern Idaho. Pocatello, ID: Idaho

State University: 42 p. Thesis.

Wenninger, E. J. 2001. Insect community response to plant diversity and productivity in a sagebrush steppe ecosystem. Pocatello:

Idaho State University. 55 p. Thesis

West, N. E. 1983. Western intermountain sagebrush steppe. In:

West, N. E., ed. Temperate deserts and semideserts, Vol. 5.

Ecosystems of the world. Amsterdam: Elsevier: 351-397.

Williams, K. S. 1997. Terrestrial arthropods as ecological indicators

of habitat restoration in southwestern North America. In:

Urbanska, K. M.; Webb, N. R.; Edwards, P. J., eds. Restoration

ecology and sustainable development. Cambridge: Cambridge

University Press: 238–258.

Winter, B. M. 1984. Effects of prescribed burning on avian foraging

ecology and arthropod abundance in sagebrush-grassland. Ames:

Iowa State University. 82 p. Thesis.

Youtie, B. A. 1986. The insect fauna associated with Great Basin

wildrye in southern Idaho. Moscow, ID: University of Idaho. 83 p.

Thesis.

Youtie, B. A.; Stafford, M. P.; Johnson, J. B. 1987. Herbivorous and

parasitic insect guilds associated with Great Basin wildrye

(Elymus cinereus) in southern Idaho. Great Basin Naturalist. 47:

644–651.
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