United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

General Technical

Report RMRS-GTR-136-vol. 2

September 2004

Restoring Western

Ranges and Wildlands

Volume 2

Chapters 18-23, Index
Abstract ______________________________________

Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps. 2004. Restoring western

ranges and wildlands. Gen. Tech. Rep. RMRS-GTR-136-vol-2. Fort Collins, CO: U.S.

Department of Agriculture, Forest Service, Rocky Mountain Research Station. Pages 295–

698 plus index.

This work, in three volumes, provides background on philosophy, processes, plant

materials selection, site preparation, and seed and seeding equipment for revegetating

disturbed rangelands, emphasizing use of native species. The 29 chapters include guidelines

for planning, conducting, and managing, and contain a compilation of rangeland revegetation

research conducted over the last several decades to aid practitioners in reestablishing healthy

communities and curbing the spread of invasive species. Volume 2 contains chapters 18-23

plus the index.

Keywords: rehabilitation, revegetation, plant ecology, seed, plant communities, wildlife

habitat, invasive species, equipment, plant materials, native plants

A

B

A—Curlleaf mountain mahogany.

B—Arrowleaf balsamroot.

C—Bluebunch wheatgrass seed

production field.

D—Antelope bitterbrush achene.

E—Squirreltail.
Restoring Western Ranges

and Wildlands

i

Compilers

Stephen B. Monsen

Richard Stevens

Nancy L. Shaw

Volume 2

Chapters 18–23, index

D E

C
The Compilers _____________________________________

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

Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah

Richard Stevens, Project Leader (retired), Utah Division of Wildlife Resources, Great Basin

Research Center, Ephraim, Utah

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

Service, Rocky Mountain Research Station, Boise, Idaho

The Authors _______________________________________

James N. Davis is Project Leader, Big Game Range Trend Program, Utah Division of Wildlife

Resources, Shrub Sciences Laboratory, Provo, Utah.

Sherel K. Goodrich is Ecologist, U.S. Department of Agriculture, Forest Service, Ashley

National Forest, Vernal, Utah.

Kent R. Jorgensen was Wildlife Biologist (deceased), Great Basin Research Center, Utah

Division of Wildlife Resources, Ephraim, Utah.

Carlos F. Lopez was Soil Scientist, U.S. Department of Agriculture, Forest Service, Rocky

Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah.

E. Durant McArthur is Project Leader and Research Geneticist, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo,

Utah.

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

Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. He is currently

a private natural resources consultant and volunteer worker for the Rocky Mountain Research

Station.

David L. Nelson was Plant Pathologist (retired), U.S. Department of Agriculture, Forest

Service, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah. He

continues contractual and volunteer work for the Rocky Mountain Research Station.

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

Mountain Research Station, Boise, Idaho.

Richard Stevens was Project Leader (retired), Great Basin Research Center, Utah Division

of Wildlife Resources, Ephraim, Utah.

Arthur R. Tiedemann was Project Leader and Range Scientist (retired), U.S. Department of

Agriculture, Rocky Mountain Research Station, Shrub Sciences Laboratory, Provo, Utah.

John F. Vallentine was Professor (Emeritus), Department of Integrative Biology, Brigham

Young University, Provo, Utah.

Gordon A. Van Epps was Associate Professor (retired), Snow Field Station, Utah Agricultural

Experiment Station, Utah State University, Ephraim, Utah.

Bruce L. Welch is Plant Physiologist, U.S. Department of Agriculture, Rocky Mountain

Research Station, Shrub Sciences Laboratory, Provo, Utah.

Steven G. Whisenant is Professor, Department of Rangeland Ecology and Management,

Texas A&M University, College Station, Texas.

G. Richard Wilson was Director (retired), Utah State Seed Laboratory, Utah Department of

Agriculture, Salt Lake City, Utah.

ii
iii

Contents

Volume 1: Page

Foreword by E. Durant McArthur .............................................................................iii

1. History of Range and Wildlife Habitat Restoration in the

Intermountain West .................................................................................. 1

2. The Intermountain Setting ............................................................................ 7

3. Research Background ................................................................................ 15

4. Basic Considerations for Range and Wildland Revegetation and

Restoration............................................................................................. 19

5. Restoration or Rehabilitation Through Management or

Artificial Treatments ............................................................................... 25

6. Climate and Terrain .................................................................................... 33

7. Assessing Soil Factors in Wildland Improvement Programs ...................... 39

8. Controlling Plant Competition ..................................................................... 57

9. Mechanical Plant Control ........................................................................... 65

10. Herbicides for Plant Control ....................................................................... 89

11. Vegetative Manipulation with Prescribed Burning .................................... 101

12. Seedbed Preparation and Seeding Practices .......................................... 121

13. Incorporating Wildlife Habitat Needs into Restoration and

Rehabilitation Projects ......................................................................... 155

14. Nutritive Principles in Restoration and Management ............................... 175

15. Plant Pathology and Managing Wildland Plant Disease Systems ........... 187

16. Management of Restored and Revegetated Sites ................................... 193

17. Guidelines for Restoration and Rehabilitation of Principal

Plant Communities ............................................................................... 199

Volume 2:

18. Grasses .................................................................................................... 295

19. Forbs for Seeding Range and Wildlife Habitats ....................................... 425

20. Chenopod Shrubs .................................................................................... 467

21. Composite Shrubs .................................................................................... 493

22. Rosaceous Shrubs ................................................................................... 539

23. Shrubs of Other Families ......................................................................... 597

Volume 3:

24. Seed Collection, Cleaning, and Storage .................................................. 699

25. Shrub and Forb Seed Production ............................................................. 717

26. Seed Germination .................................................................................... 723

27. Seed Testing Requirements and Regulatory Laws .................................. 733

28. Establishing Plants by Transplanting and Interseeding ........................... 739

29. Production and Use of Planting Stock ...................................................... 745

References ............................................................................................... 769

Appendix 1 Scientific/Common Names .................................................... 847

Appendix 2 Common/Scientific Names .................................................... 866

Index .................................................................................................. Index-1
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The use of trade or firm names in this publication is for reader information and does not

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CAUTION:

PESTICIDES

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not

contain recommendations for their use, nor does it imply that the uses

discussed here have been registered. All uses of pesticides must be

registered by appropriate State and/or Federal agencies before they can

be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals,

desirable plants, and fish or other wildlife—if they are not handled or

applied properly. Use all pesticides selectively and carefully. Follow

recommended practices for the disposal of surplus pesticides and

pesticide containers.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 295

Stephen B. Monsen

Richard Stevens

Nancy Shaw

18

Chapter

Grasses are adapted to a wide range of edaphic and climatic

conditions and are found in nearly all plant communities. In the

Western United States, grasses are seeded on disturbances to

provide forage (Hull and Holmgren 1964; Vallentine 1989), wildlife habitat (Plummer and others 1968), and watershed stability

(Cornelius 1946; Hafenrichter and others 1949; Piper 1934;

Stewart and Young 1939). A number of introduced grass species

proved well-suited to Western rangelands and received extensive

use in early reseeding efforts (Barnes and others 1995;

Hafenrichter and others 1968; Moser and others 1996). Use of

these and other introductions continued over time, but by the late

1900s, greater emphasis was being placed on the use of native

grasses (Roundy and others 1997).

Selection of introduced and native grasses for Western rangelands began in the late nineteenth century. Excessive livestock

grazing across the West followed early European settlement,

resulting in widespread disturbances, weed invasions, and a

need for revegetation species to stabilize watersheds and produce

forage. A few species and accessions were introduced in the West

before 1900 (Hanson 1959). Some arrived by chance (Weintraub

Grasses
296 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Chapter Contents by Species

Agropyron cristatum (Fairway Crested Wheatgrass) ....................................................................... 297

Agropyron dasystachyum (Thickspike Wheatgrass) ......................................................................... 318

Agropyron desertorum (Standard Crested Wheatgrass or Desert Crested Wheatgrass) ............... 320

Agropyron elongatum (Tall Wheatgrass) .......................................................................................... 325

Agropyron intermedium (Intermediate Wheatgrass) ....................................................................... 327

Agropyron sibiricum (Siberian Wheatgrass) ..................................................................................... 332

Agropyron smithii (Western Wheatgrass or Bluestem Wheatgrass) ............................................... 334

Agropyron spicatum (Bluebunch Wheatgrass).................................................................................. 337

Agropyron trachycaulum (Slender Wheatgrass) ............................................................................... 342

Agrostis stolonifera (Redtop or Carpet Bentgrass) ........................................................................... 345

Alopecurus arundinaceus (Creeping Foxtail or Reed Foxtail) ......................................................... 346

Alopecurus pratensis (Meadow Foxtail) ............................................................................................ 346

Aristida purpurea (Red Threeawn, Purple Threeawn) .................................................................... 349

Arrhenatherum elatius (Tall Oatgrass) ............................................................................................. 351

Bouteloua curtipendula (Sideoats Grama) ........................................................................................ 353

Bouteloua eriopoda (Black Grama).................................................................................................... 355

Bouteloua gracilis (Blue Grama) ....................................................................................................... 356

Bromus anomalus (Nodding Brome) ................................................................................................. 358

Bromus ciliatus (Fringed Brome) ...................................................................................................... 358

Bromus carinatus (Mountain Brome, California Brome) ................................................................. 360

Bromus inermis (Smooth Brome, Hungarian Brome) ...................................................................... 363

Bromus riparius (Meadow Brome) .................................................................................................... 365

Dactylis glomerata (Orchardgrass) .................................................................................................... 367

Deschampsia caespitosa (Tufted Hairgrass) ..................................................................................... 369

Distichlis spicata (Inland Saltgrass) ................................................................................................. 371

Elymus canadensis (Canada Wildrye) ............................................................................................... 373

Elymus cinereus (Great Basin Wildrye) ............................................................................................ 374

Elymus glaucus (Blue Wildrye) ......................................................................................................... 377

Elymus junceus (Russian Wildrye) .................................................................................................... 379

Elymus salinus (Salina Wildrye) ....................................................................................................... 381

Elymus simplex (Alkali Wildrye or Low-Creeping Wildrye) ............................................................ 382

Elymus triticoides (Creeping Wildrye or Beardless Wildrye) .......................................................... 383

Festuca arundinacea (Tall Fescue, Reed Fescue, or Alta Fescue) ................................................... 384

Festuca idahoensis (Idaho Fescue or Bluebunch Fescue)................................................................. 386

Festuca ovina (Sheep Fescue or Alpine Fescue)................................................................................ 388

Hilaria jamesii (Galleta or Curly Grass)........................................................................................... 390

Koeleria macrantha (Prairie Junegrass) ........................................................................................... 392

Oryzopsis hymenoides (Indian Ricegrass) ......................................................................................... 393

Phalaris arundinacea (Reed Canarygrass) ....................................................................................... 396

Phleum alpinum (Alpine Timothy or Mountain Timothy) ............................................................... 398

Phleum pratense (Timothy) ................................................................................................................ 399

Poa ampla (Big Bluegrass)................................................................................................................. 401

Poa compressa (Canada Bluegrass) ................................................................................................... 402

Poa fendleriana (Mutton Bluegrass) ................................................................................................. 404

Poa pratensis (Kentucky Bluegrass).................................................................................................. 406

Poa secunda (Sandberg Bluegrass).................................................................................................... 408

Sitanion hystrix (Squirreltail, bottlebrush) ...................................................................................... 410

Sporobolus airoides (Alkali Sacaton)................................................................................................. 413

Sporobolus cryptandrus (Gray Sand Dropseed)................................................................................ 414

Stipa columbiana (Subalpine Needlegrass or Columbia Needlegrass) ........................................... 416

Stipa comata (Needle-and-thread) .................................................................................................... 418

Stipa lettermanii (Letterman Needlegrass) ...................................................................................... 419

Stipa thurberiana (Thurber Needlegrass)......................................................................................... 421

Stipa viridula (Green Needlegrass)................................................................................................... 422
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 297

Chapter 18 Grasses

1953); others were selected for specific uses and areas

(Dillman 1946). Field testing and seeding trials for

high-elevation disturbances were initiated at this time.

Based on results of these trials, Williams (1898) described the values and uses of introduced forage grasses

for the Eastern Rocky Mountains.

Seeding trials conducted by the Bureau of Plant

Industry were initiated in 1902 (Cotton 1908). Forest

Service reseeding research utilizing forage species

began in 1907 in cooperation with this agency (Sampson

1913). Ecological studies of grasses native to the National Forests were initiated in about 1911 (USDA

Forest Service 1914). Efforts to evaluate grasses for

range and wildlife uses were expanded by Forest

Service researchers and their cooperators by about

1930 (Stewart and others 1939). Cool-season grasses

received special attention for use on Western rangelands (Koonce 1946; Piper 1934).

Breeding and selection programs were organized

within the USDA Soil Conservation Service, now the

USDA Natural Resources Conservation Service, in

the 1930s (Hafenrichter and others 1968). The need to

stabilize abandoned farmlands during the 1930’s further stimulated the demand for conservation species

(Dillman 1946). Weintraub (1953) published a review

of publications on testing and field planting studies

conducted between 1920 and 1950.

The Soil Conservation Service (now the USDA

Natural Resources Conservation Service) and their

cooperators began releasing grass varieties in the mid

1940s (Alderson and Sharp 1994; Hanson 1959). Some

introduced range and pasture varieties obtained immediate success. Introductions from areas of Europe

and Asia with environmental conditions similar to

those encountered in Western North America proved

useful over wide areas. Testing of drought-tolerant

grasses adapted to arid and semiarid rangelands received increasing emphasis during recent decades

(Barnes and others 1952; Bridges 1942; Christ 1934;

Forsling and Dayton 1931; Koonce 1946; Piper 1934;

Price 1938; Stoddart 1946a). Some of the most important grass varieties developed for use on Western range,

wildland, and conservation plantings are listed in table 1.

Following is a discussion of introduced and native

grasses that have proven most useful for seeding

Western wildlands. The range of adaptability, uses,

and characteristics of important grasses are summarized in table 2. In the past, productive accessions that

were palatable and responded well to grazing were

emphasized in plant selection programs for rangelands. Characteristics such as good seed production,

ease of seed cleaning and seeding, and good seedling

establishment also received priority in the selection

process (Frischknecht 1951; Hanson and Carnahan

1956; Plummer and others 1943). Although important, these emphases often discouraged the use of less

productive species and accessions that might, nonetheless, be adapted to specific situations. With increased efforts to reestablish native plant communities, it became essential to more fully utilize the array

of species and populations adapted to the wide array of

environments encountered on Western rangelands.

Many native species have not been brought into seed

production, but should be seeded in appropriate areas

and maintained through proper management. Research is being conducted to develop seed transfer

guidelines for native grasses of the Intermountain

region to permit increase of adapted seed supplies for

revegetation that will conserve the genetic diversity of

native populations. Thus, at the present and into the

future, greater consideration will be given to native

grasses, forbs, and shrubs.

Grass taxonomy has evolved continuously since

Linnaeus published the first descriptions in 1753.

Extensive revisions, particularly within the Triticeae,

have been made over the last few decades as our

understanding of origins and genetic relationships

increased (Arnow 1987; Barkworth and others 1983).

Löve (1982, 1984) reorganized genera within the

Triticeae based on genomic relationships. Dewey (1982,

1983a, 1984) concurred with this revision as it provided a clearer picture of phylogenetic and biological

closeness. New taxonomic keys, however, continue to

utilize conventional morphological features to separate taxa. We subscribe to the recent changes, but we

have elected to use the more traditional nomenclature

because most land managers, seed dealers, and growers are better acquainted with it. Traditional and

current nomenclatures are presented in table 3.

Agropyron cristatum

Fairway Crested Wheatgrass ______

Description

Fairway crested wheatgrass, also called crested

wheatgrass, is an introduced, long-lived, cool-season,

generally nonrhizomatous bunchgrass that produces

strong tillers and an extensive root system. Stems and

leaves are tightly clustered at the base. Plants are

generally uniform in stature and growth form. Stem

heights within individual plants range from 8 to 20

inches (20 to 50 cm), but may reach 40 inches (1 m).

Stems are generally erect, but some may be bent

slightly at the lower nodes. Leaves develop from the

plant base and stems. Leaf blades are 2 to 8 inches (5

to 20 cm) long and 0.08 to 0.2 inch (2 to 5 mm) wide,

appearing flat or loosely bent along the midrib.

Blades are rough, heavily grooved with raised nerves,

and often have a bluish cast. Hairs are sometimes

present on the upper portion of the leaf blade. Auricles

are small and clasping. Ligules are membranous
298 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses Table 1—Selected grass cultivars and germplasms for range and wildlife projects. Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes Bentgrass, redtop Streaker 1982 Combined Jacklin Seed Company, Highly rhizomatous, fine leaves Agrostis stolonifera Illinois collections Post Falls, ID Bluegrass, alpine Gruening 1986 La Cure, NRCS, Palmer, AK Low growing, nonrhizomatous, winter hardy, selected for Poa alpina Switzerland Alaska Bluegrass, big Sherman 1945 Sherman County, OR NRCS, Pullman, WA Early greenup, highly productive Poa ampla Bluegrass, Canada Foothills 2001 Composite of NRCS, Bridger, MT Good forage and seed production, useful on low-fertility Poa compressa Germplasm eight accessions sites of moderate acidity and at high elevations Reubens 1976 Reubens, ID Jacklin Seed Company, Dense, low-maintenance turf for erosion control on Post Falls, ID infertile sites Bluegrass, Kentucky Newport 1958 Lincoln County, OR NRCS, Pullman, WA Vigorous, highly productive, broad climatic tolerance Poa pratensis Bluegrass, Sandberg Canbar 1979 Blue Mountains, WA NRCS, Pullman, WA Good drought tolerance and provides early greenup, Poa secunda competitive with annual weeds P. canbyi ( ) High Plains 2000 Composite from NRCS, Bridger, MT Tall, productive, used in native seed mixes for wildlife Germplasm Montana and Wyoming habitat and reestablishment of native plant communities Service 1989 Whitehorse, Yukon, NRCS, Palmer, AK Erect, vigorous, winter hardy Canada Brome, California Cucamonga 1949 Cucamonga, CA NRCS, Lockeford, CA Rapid development, early maturing, selfseeding, cover crop Bromus carinatus Brome, meadow Regar 1966 Zek, Kars Province, NRCS, Aberdeen, ID Excellent establishment and early development, highly Bromus riparius Turkey productive, tolerant of shade, winter hardy Brome, mountain Bromar 1946 Whitman County, WA NRCS, Pullman, WA Rapid development, erect, leafy, late maturing, heavy Bromus marginatus seed and forage producer Garnet 2000 Garnet, MT UCEPC, Meeker, CO Good stand longevity and head smut resistance Other cultivars are Fleet and Paddock. (con.)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 299

Chapter 18 Grasses Brome, smooth Manchar 1943 Manchuria, China NRCS, Pullman, WA Most widely planted northern type in the West, moderately Bromus inermis spreading type, high yields, good regrowth Northern types Polar 1965 Arctic brome hybrids Alaska AES Winter hardy, high yields, early spring growth and smooth brome sources Southern types Achenbach 1944 Washington County, KS Kansas AES Leafy, vigorous, spreads rapidly, competitive Lincoln 1942 Hungary Nebraska AES Aggressive sodformer, good seedling vigor, drought tolerant Numerous other smooth brome cultivars of both the northern and southern types have been developed for regional uses. Canarygrass, reed Ioreed 1946 Composite clones Iowa AES Very productive, well-suited to wet sites, useful for hay Phalaris arundinacea Other less common cultivars include Auburn, Castor, Frontier, Highland, Rise, and Vantage. Fescue, Arizona Redondo 1973 Los Alamos, NM NRCS, Los Lunas, NM High seed yields, excellent seed production Festuca arizonica Fescue, hard sheep Durar 1949 Union, OR, NRCS, Pullman, WA Leafy, persistent, moderate forage value, adapted to Festuca ovina var. 1934 planting semiarid sites duriuscula Fescue, Idaho Joseph 1983 Composite clones U of Idaho, Moscow, ID Improved seed size, high fertility, improved germination, Festuca idahoensis large growth form (F. ovina var. ingrata) Nezpurs 1983 Composite clones U of Idaho, Moscow, ID Excellent forage attributes, erect, leafy ecotype Fescue, sheep Covar 1977 Konya, Turkey NRCS, Pullman, WA Dwarf, blue green, densely tufted, drought tolerant, Festuca ovina persistent, competitive var. ovina MX86 1989 Germany Jacklin Seed Company, Improved seedling vigor and yields Post Falls, ID Fescue, tall Alta 1940 Composite clones Oregon AES Remains green throughout the summer, high forage yields, Festuca arundinacea cold tolerant Fawn 1964 Named varieties and Oregon AES High forage and seed production, widely adapted foreign introductions Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes (con.)
300 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses Foxtail, creeping (reed) Garrison 1963 Europe NRCS, Bismarck, ND Highly productive, excellent quality forage, Alopecurus heavy seed producer, rhizomatous, adapted to arundinaceus mountain meadows Foxtail, meadow Dan 1987 Wilno, Poland AC, Ottawa, Early maturity, good winter hardiness Alopecurus pratensis Ontario, Canada Galleta Viva 1979 New Kirk, NM NRCS, Los Lunas, NM Improved seedling vigor, high seed production Hilaria jamesii Grama, black Nogal 1971 Socorro County, NM NRCS, Los Lunas, NM Intermediate growth habit, fine stemmed Bouteloua eriopoda Grama, blue Hachita 1980 Hachita, NM NRCS, Los Lunas, NM High seed and herbage production, drought tolerant Bouteloua gracilis Lovington 1963 Lovington, NM NRCS, Los Lunas, NM Excellent seedling vigor, rapid establishment Grama, sideoates Butte 1958 Holt and Platte Nebraska AES Good seedling vigor, winter hardy, long lived Bouteloua curtipendula Counties, NE El Reno 1944 El Reno, OK NRCS, Manhatten, KS Outstanding leafiness, forage production, vigor Haskell 1983 Haskell, TX NRCS, Knox City, TX Strong rhizome production Killdeer 1960 Dunn and Bowman NRCS, Bismarck, ND Outstanding vigor, tolerates cold and dry climates Counties, TX Niner 1984 Socorro County, NM NRCS, Los Lunas, NM No rhizomes, matures evenly Pierre 1960 Pierre, SD NRCS, Bismarck, ND Outstanding vigor, leafiness, good seedling vigor Premier 1960 Mexico Texas AES Upright, leafy type, seedlings develop rapidly Trailway 1958 Holt County, NE Nebraska AES Winter hardy, long lived, late maturing Vaughn 1940 Vaughn, NM NRCS, Los Lunas, NM Good seedling vigor, drought tolerant, easily established Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes (con.)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 301

Chapter 18 Grasses Hairgrass, tufted Norcoast 1981 Alaska Alaska AES High forage production, adapted to semiwet and acid soils Deschampsia caespitosa Nortran 1986 Composite collection, Alaska AES Low-maintenance ground cover Alaska and Iceland Peru Creek 1995 Summit County, CO NRCS, Meeker, CO Adapted to high elevations and acidic soils Muhly, spike El Vado 1973 Park View, NM NRCS, Los Lunas, NM Upright growth form, leafy, free of galls Muhlenbergia wrighti Needlegrass, green Lodorm 1970 Bismarck, ND ARS, Mandan, ND High palatability, good regrowth Stipa viridula Orchardgrass Latar 1957 U.S.S.R. NRCS, Pullman, WA Late maturing, abundant leaves, highly productive Dactylis glomerata Paiute 1983 Ankara, Turkey NRCS, Aberdeen, ID Drought hardy, easily established, moderately productive Potomac 1954 Combined Washington Crop Highly productive, persistent, leafy type collections Improvement Association Numerous other pasture cultivars are available Reedgrass, bluejoint Sourdough 1976 Alaska NRCS, Palmer, AK Winter-hardy, robust, cool-season perennial developed Calamagrostis for Alaska canadensis Ricegrass, Indian Nezpar 1978 Whitebird, ID NRCS, Aberdeen, ID Excellent seedling vigor, low-percentage hard seeds Oryzopsis hymenoides Paloma 1974 Pueblo, CO NRCS, Los Lunas, NM Soil stabilization, drought tolerant, highly palatable Rimrock 1996 Billings, MT NRCS, Bridger, MT Good seed retention, low shattering, large stature Sacaton, alkali Salado 1982 Claunch, NM NRCS, Los Lunas, NM Good seedling vigor and forage production Sporobolus airoides Saltalk 1981 Erick, OK NRCS, Knox City, TX Highly palatable, good germination Squirreltail, bottlebrush Sand Hollow 1996 Gem County, ID Self-pollinating, grows at low temperatures Sitanion hystrix (con.) Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes
302 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses Timothy Climax 1947 Combined AC, Ottawa, Highly productive, very leafy, Phleum pratense progeny Ontario, Canada establishes quickly Drummond 1940 Northern Europe McGill University, Shade tolerant, provides late-season forage Montreal,Quebec, Canada Numerous pasture cultivars are available. Wheatgrass, beardless Whitmar 1946 Whitman County, WA NRCS, Pullman, WA Long-lived, highly productive, drought resistant Agropyron spicatum ssp. inerme

(Pseudoroegneria

spicata ssp. inermis)

Wheatgrass, bluebunch Anatone 2003 Asotin County, WA Breeder, seed USDA FS, Ricky Mountain Research Station,

Agropyron spicatum, maintenance Provo, UT; NRCS , Aberdeen, ID Good seedling vigor,

Pseudoroegneria recommended for semiarid sites

spicata Goldar 1989 Asotin County, WA NRCS, Aberdeen, ID Good seedling vigor, adapted to semiarid sites

P-7 2001 Multiple origin polycross ARS, Logan, UT Recommended for mesic to semiarid sites

of 25 populations

Wheatgrass, fairway Douglas 1994 U.S.S.R./Iran/Turkey NRCS, Aberdeen, ID Excellent seedling vigor and summer greenness

crested

Agropyron cristatum Ephraim 1983 Ankara, Turkey NRCS, Aberdeen, ID Low growing, slightly rhizomatous, drought tolerant

Fairway 1927 U.S.S.R. AC, Saskatoon, Productive, leafy types

Saskatchewan, Canada

Parkway 1969 Fairway germplasm AC, Saskatoon, High seed and forage yields, leafiness, increased vigor

Saskatchewan, Canada

Ruff 1974 Commercial lots Nebraska AES Early, cool season, adapted to low rainfall areas

and Nebraska AES

Wheatgrass, hybrid NewHy 1989 Quackgrass and ARS, Logan, UT Vigorous, highly productive, high salinity tolerance

Agropyron repens x bluebunch wheatgrass

A. spicatum hybrid

Elytrigia repens ( x

Pseudoroegneria spicata)

(con.)

Table 1 (Con.)

Common and Cultivar or Release Breeder

scientific names germplasm date Origin seed maintenancea Attributes
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 303

Chapter 18 Grasses Wheatgrass, hybrid SL1 1991 Hybrid ARS, Logan, UT Large culms, productive, improved seedling vigor Agropyron spicatum x A. dasystachyum Wheatgrass, hybrid CDII 1996 10-clone synthetic ARS, Logan, UT Leafy, highly productive, adapted to arid sites crested derived from Hycrest Agropyron cristatum x A. desertorum Hycrest 1984 Central Asia ARS, Logan, UT Excellent seedling vigor and early maturity, highly competitive with annuals Kirk 1987 Tetraploid selection AC, Saskatoon, Improved vigor, fertility, seed development, reduced awns Saskatchewan, Canada Wheatgrass, intermediate Amur 1952 Manchuria, China NRCS, Los Lunas, NM Excellent forage traits, good establishment attributes Agropyron intermedium (Thinopyrum Chief 1961 U.S.S.R. and AC, Saskatoon, Improved forage and seed yields intermedium) selection from Ree Saskatchewan, Canada Clarke 1980 20-clone synthetic AC, Swift Current, Drought tolerant, winter hardy, high seed yields Saskatchewan, Canada Greenar 1945 U.S.S.R. NRCS, Pullman, WA Vigorous, highly productive, late maturing, good regrowth Oahe 1961 Selection from South Dakota AES Vigorous, rhizomatous, drought tolerant Ree Reliant 1991 6-clone synthetic ARS, Mandan, ND Late maturation, high forage production Rush 1995 Germany NRCS, Aberdeen, ID Excellent seedling vigor, highly productive, good ground cover Slate 1969 Blend of Amur Nebraska AES Erect culms, strongly spreading and Nebraska 50 Tegmar 1968 Turkey NRCS, Aberdeen, ID Drought tolerant, dwarf strain, late maturing, sodforming (con.) Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes
304 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses Wheatgrass, pubescent Greenleaf 1966 Commercial AC, Lethbridge, Sod forming, high seedling vigor and forage production, Agropyron intermedium seed lots Alberta, Canada winter hardy (Thinopyrum intermedium, A. trichophorum) Luna 1963 U.S.S.R. NRCS, Los Lunas, NM Adapted to warm temperatures and dry sites, excellent seedling vigor, highly productive Topar 1953 U.S.S.R. NRCS, Aberdeen, ID Vigorous, drought tolerant, sodformer Wheatgrass, Siberian P27 1953 U.S.S.R. NRCS, Aberdeen, ID Drought resistant, good seedling vigor, persistent Agropyron sibiricum (A. fragile) Vavilov 1994 U.S.S.R. NRCS, Aberdeen, ID Excellent seedling vigor, highly productive Wheatgrass, slender Adanac 1990 Climax, AC, Saskatoon, Highly productive, adapted to saline soils Agropyron trachycaulum Saskatchewan, Saskatchewan, Canada Elymus trachycaulus ( ) Canada Primar 1946 Beebe, MT NRCS, Pullman, WA Vigorous, early growing, long-lived, high yield, alkali tolerant Pryor 1988 South-central MT NRCS, Bridger, MT Tolerant of drought and salinity, vigorous seedlings Revenue 1970 Revenue, AC, Saskatoon, Excellent establishment, high seed and forage yields Saskatchewan, Saskatchewan, Canada Canada San Luis 1984 Rio Grande County, CO NRCS, Meeker, CO Excellent establishment and longevity, adapted to high elevations Wheatgrass, Snake River Secar 1980 Lewiston, ID NRCS, Pullman, WA Dryland ecotype, drought tolerant, tufted bunchgrass Elymus wawawaiensis (Agropyron spicatum, E. lanceolatus spp. wawawai) Wheatgrass, standard Nordan 1953 Central Asia ARS, Mandan, ND Good seedling vigor, large seeds, high yields, cold hardy crested Agropyron desertorum Summit 1953 U.S.S.R. AC, Saskatoon, Excellent forage traits, high seed yields Saskatchewan, Canada (con.) Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes
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Chapter 18 Grasses Wheatgrass, tall Alkar 1951 U.S.S.R. NRCS, Pullman, WA Tall, robust type, tolerant of wet and saline soils Agropyron elongatum (Thinopyrum ponticum) Jose 1965 Australia NRCS, Los Lunas, NM Drought and salt tolerant, early maturing, fine leaves Largo 1961 Turkey NRCS, Los Lunas, NM Highly productive, large bunch type Orbit 1966 PI 98526 and AC, Swift Current, Winter hardy, withstands flooding, good seed and local strain Saskatchewan, Canada forage yield Platte 1972 Nebraska 98526 Nebraska AES Winter hardy bunchgrass with short, spreading rhizomes, tolerant of alkali soils Wheatgrass, thickspike Bannock 1995 Composite NRCS, Aberdeen, ID Rhizomatous, highly productive, drought tolerant Agropyron dasystachyum Elymus lanceolatus (var. lanceolatus, Critana 1971 Havre, MT NRCS, Bridger, MT Excellent seedling vigor, strongly rhizomatous A. riparium) Schwendimar 1994 The Dalles, OR NRCS, Pullman, WA Weakly rhizomatous, adapted to dry sites Sodar 1954 Grant County, MT NRCS, Aberdeen, ID Drought resistant, strongly rhizomatous, excellent ground cover Thickspike 1994 The Dalles, OR NRCS, Aberdeen, ID Vigorous sodformer and controls weeds Wheatgrass, western Arriba 1973 Flagler, CO NRCS, Los Lunas, NM Rapid germination, excellent establishment, good seed Agropyron smithii and forage yields Elytrigia smithi ( , Pascopyrum smithii) Barton 1970 Heizer, KS NRCS, Manhattan, KS Strongly rhizomatous, leafy, highly productive Flintlock 1975 Nebraska and Kansas AES, Lincoln, NE Spreading type, early spring growth, drought tolerant Rodan 1983 Mandan, ND ARS, Mandan, ND Highly productive, excellent pasture type, vigorous Rosana 1972 Forsyth, MT NRCS, Bridger, MT Excellent seedling vigor, close sod, leafy Walsh 1982 Alberta and AC, Lethbridge, Drought tolerant, dense sodformer, high yields, Saskatchewan Alberta, Canada tolerant of flooding and salinity (con.) Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes
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Chapter 18 Grasses Wildrye, Altai Eejay 1989 U.S.S.R. AC, Indian Head, Highly productive, good-quality late-season herbage, good Elymus angustus Saskatchewan, Canada seedling vigor (Leymus angustus) Pearl 1989 U.S.S.R. AC, Swift Current, Highly productive, good production in late summer, good Saskatchewan, Canada seedling vigor, drought tolerant Prairieland 1976 U.S.S.R. AC, Swift Current, Excellent producer, good-quality late-season herbage, Saskatchewan, Canada free of leaf spot Wildrye, beardless Rio 1991 Stratford, CA NRCS, Lockford, CA Excellent seed viability, vigorous spreader or creeper Elymus triticoides (Leymus triticoides) Shoshone 1980 Riverton, WY NRCS, Bridger, MT Leafy, fine stemmed, highly productive, rhizomatous Wildrye, blue Arlington 1995 Arlington, WA NRCS, Corvallis, OR Vigorous, disease-free, high-quality seed Elymus glaucus Wildrye, Canada Mandan 1946 Mandan, ND ARS, Mandan, ND Easily established, short stature, high seed and Elymus canadensis forage production, leafy, persistent Wildrye, Dahurian Arthur 1989 China AC, Swift Current, Short lived, highly productive first 13 years of stand Elymus dahuricus Saskatchewan, Canada establishment, erect, green foliage James 1989 China AC, Indian Head, Highly productive, good stand establishment Saskatchewan, Canada Wildrye, Great Basin Magnar 1979 Saskatchewan, NRCS, Aberdeen, ID Large, robust, extremely productive, excellent Elymus cinereus Canada ground cover, good seed quality (Leymus cinereus) Trailhead 1991 Roundup, MT NRCS, Bridger, MT Vigorous, drought tolerant, early spring forage Wildrye, mammoth Volga 1949 U.S.S.R. NRCS, Meeker, CO Tall, persistent, useful for dune stabilization Elymus giganteus (Leymus racemosus) Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes (con.)
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Chapter 18 Grasses Table 1 (Con.) Common and Cultivar or Release Breeder scientific names germplasm date Origin seed maintenancea Attributes Wildrye, Russian Bozoisky- 1984 U.S.S.R. ARS, Logan, UT Improved seedling vigor, highly productive Elymus junceus Select Psathyrostachys (juncea) Cabree 1976 Six clones from AC, Lethbridge, Improved seed retention, seedling vigor, and forage Manyberries, Alberta, Alberta, Canada and seed yield Canada seeding Mankota 1991 Six clone synthetic, Mandan, ND Excellent fall and winter grazing, seedling establishment, U.S.S.R. and and forage production Mandan, ND Mayak 1971 Twenty-clone AC, Swift Current, Productive, good late-summer herbage, resistant to synthetic Saskatchewan, Canada leaf spot disease Swift 1978 Twenty-six clone AC, Swift Current, Improved seedling vigor, highly productive, good seed synthetic Saskatchewan, Canada quality and yield Tetracan 1988 Colchicine induced AC, Swift Current, Large seeds, improved seedling vigor tetraploids Saskatchewan, Canada Vinall 1960 Five clone ARS, Mandan, ND Provides early greenup and fall grazing, high seed synthetic production a NRCS = USDA Natural Resource Conservation Service, formerly USDA Soil Conservation Service; AES = Agricultural Experiment Station; AC = Agriculture Canada; ARS = USDA Agricultural Research Service; UCEPC = Upper Colorado Environmental Plant Center.
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Chapter 18 Grasses

Table 2—Characteristics of selected grasses.

- - - - - - - - - - - - - - - - - - - - - - - ratingsb- - - - - - - - - - - - - - - - - - - - - - -

Barley, bulbous 4 4 3 4 5 5 4 2 S 5 2 2 3 4 3 5 5 5 3 2 2 4 WM,JP,BS

Barley, meadow 3 4 3 4 4 5 4 3 V 5 4 3 4 3 3 5 4 3 3 4 4 2 SA,A,WM

Bluegrass, big 3 5 5 4 5 5 4 4 S 4 3 4 4 4 3 4 4 4 4 4 3 3 A,MB,JP,PP,MS,R

Bluegrass, Canada 3 5 5 4 4 5 4 4 V 5 3 4 4 4 3 4 4 4 4 4 2 3 SA,A,MB,PP

Bluegrass, Kentucky 3 5 5 4 4 5 4 5 V 5 4 4 2 5 2 4 4 5 5 4 4 5 SA,A,WM,MB,PP,

JP,MS,R

Bluegrass, mutton 2 3 5 4 4 5 3 5 S 5 5 4 4 4 4 4 4 3 3 3 3 3 MB,PP,JP,MS

Bluegrass, Sandberg 3 5 5 4 4 4 4 5 S 5 2 3 4 5 2 5 2 4 4 2 2 5 MB,PP,JP,MS,

BS,WS,SS,C,AW

Brome, mountain 4 5 5 4 5 5 5 4 S 3 4 4 5 3 4 4 4 4 4 4 4 3 SA,A,MB,PP,MS

Brome, nodding 4 5 5 4 5 5 5 4 S 4 4 4 5 3 4 4 4 4 4 5 4 3 SA,A,MB,PP,MS

Brome, smooth 5 5 5 4 5 5 4 5 V 4 4 4 1 5 5 4 4 5 5 5 4 4 SA,A,WM,MB, PP,MS,R

 northern

Brome, smooth 5 5 5 4 5 5 5 5 V 4 4 3 1 5 4 4 3 5 5 3 4 4 SA,A,WM,MB,

 southern PP,JP,MS,BS,R

Brome, subalpine 3 5 5 4 5 4 3 4 S 4 4 4 4 4 4 4 4 3 4 5 4 3 SA,A,WM,MS

Canarygrass, reed 5 3 4 4 3 2 5 5 V 3 5 3 1 5 5 3 3 5 5 2 5 2 SA,A,WM,IS,R

Dropseed, sand 3 3 4 4 3 2 4 4 S 5 3 2 4 4 3 3 2 4 4 2 3 3 MB,PP,JP,BS,

WS,SS,BB,C,AW

Fescue, hard sheep 4 5 5 5 5 4 4 5 S 4 3 2 1 5 2 2 2 5 4 2 2 4 A,MB,PP,JP,MS,BS

Fescue, Idaho 3 5 5 5 5 4 3 3 S 5 3 2 4 4 3 5 4 4 4 2 2 3 MB,PP,MS

Fescue, meadow 3 5 5 5 5 4 4 5 S 4 4 3 2 4 4 5 4 5 5 3 4 2 SA,A,WM,JP,BG,IS

Fescue, sheep 3 5 5 5 5 4 4 4 S 5 3 2 3 4 3 3 2 4 4 2 2 5 SA,A,MB,PP,MS

Fescue, spike 2 4 4 4 3 3 4 3 V 4 3 2 4 4 4 3 3 4 3 3 3 2 MB,PP,JP,MS

Fescue, reed 5 3 3 3 5 4 4 5 S 4 4 3 2 2 4 5 4 4 5 3 5 2 SA,A,WM,JP,BG,IS

Fescue, Thurber 2 3 4 4 4 4 3 3 S 5 2 2 4 5 3 4 3 4 3 2 2 4 SA,A,MB,MS

Foxtail, meadow 5 2 2 2 4 2 4 5 V 4 4 4 2 5 5 5 5 5 5 3 5 2 SA,A,WM,MB,IS

Foxtail, creeping 5 2 2 2 4 2 4 5 V 4 4 4 2 5 5 5 5 5 5 3 5 2 SA,A,WM,MB,IS

Galleta 3 3 3 3 3 4 3 3 V 3 4 2 4 4 3 4 4 3 4 2 1 5 JP,BS,WS,SS,BB

Grama, sideoats 3 4 4 5 4 4 3 3 S 3 4 2 4 4 3 4 2 3 4 2 1 3 JP,BS,WS,SS,BB

Grama, blue 2 4 4 5 4 4 3 4 V 3 4 2 3 5 2 4 2 5 5 3 1 3 JP,BS,WS,SS,BB

Grama, black 2 4 4 5 4 4 3 4 V 3 4 2 3 5 2 4 2 5 5 3 1 3 JP,BS,WS,SS,BB

Hair grass, tufted 3 4 4 4 4 4 4 4 S 3 4 2 4 4 2 2 2 4 3 2 2 3 SA,A,WM,MB,PP,IS

Junegrass, prairie 2 4 5 4 3 4 4 3 S 5 3 3 5 4 2 5 3 4 3 3 2 3 MB,PP,JP,MS, JP,WS,BB

Needle-and-thread 3 2 2 3 3 4 3 5 S 5 4 3 3 5 3 4 2 4 4 2 2 3 MB,PP,JP,BS, WS,C,AW

Needlegrass, green 4 3 2 3 4 4 4 4 S 5 4 3 4 4 4 4 3 4 4 3 3 3 A,MB,PP,MS

Needlegrass, Letterman 3 3 2 3 3 3 3 4 S 3 5 2 4 5 2 2 2 4 4 3 3 2 SA,A,MB,MS

Needlegrass, subalpine 2 2 2 3 3 3 3 3 S 3 5 3 4 5 2 4 3 4 4 4 3 2 SA,A,MS

Needlegrass, Thurber 3 2 2 3 2 3 3 5 S 5 4 3 3 5 4 4 4 4 4 3 2 3 MS,BS,WS,C,AW

Oatgrass, tall 4 3 3 4 5 4 4 4 S 5 4 3 4 3 4 5 4 3 4 5 3 2 SA,A,MB,PP,MS

Oniongrass 2 5 5 5 5 4 5 3 V 4 3 3 4 2 3 4 4 3 3 5 3 2 A,MB,PP,MS

Orchardgrass 5 5 5 5 5 5 5 4 S 5 4 4 3 4 5 5 4 4 5 5 4 5 SA,A,WM,MB, PP,MS

Redtop 3 3 4 4 3 3 3 5 V 3 4 2 3 4 2 3 2 4 3 3 4 3 A,WM,BS,BG,IS

Ricegrass, Indian 4 3 3 4 2 4 3 4 S 5 2 2 4 5 3 4 3 4 4 2 5 4 MB,PP,PJ,MS,BS,

WS,SS,BG,BB,C,AW

(con.)

Species

Seed production

Ease of seed harvesting

Ease of seed cleaning

Ease of seeding

Germination

Seedling vigor

Growth rate

Final establishment

Method of spread

Spring growth

Summer growth

Fall growth

Compatibility with other species

Longevity

Forage production

Spring palatability

Summer palatability

Grazing tolerance

Soil stabilization

Shade tolerance

Flood tolerance

Ecotypic variability

Vegetative typesa
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 309

Chapter 18 Grasses

- - - - - - - - - - - - - - - - - - - - - - - ratingsb- - - - - - - - - - - - - - - - - - - - - - -

Rye, mountain 5 4 4 5 5 5 5 4 S 5 2 5 3 2 5 5 4 4 4 3 2 3 MB,PP,JP,MS,BS,WS,SS,

BB,C,AW

Sacaton, alkali 3 3 4 4 1 2 2 4 V 3 3 3 1 5 2 2 2 5 5 2 4 3 WM,BS,WS,SS,BG,

BB,IS,R

Squirreltail, bottlebrush 4 2 2 3 4 4 4 4 S 5 2 2 4 4 3 4 2 4 4 2 5 5 MB,PP,JP,BS,WS,SS,BG,

BB,C,AW

Three-awn, purple 2 2 2 3 3 4 3 4 S 5 2 2 3 5 3 1 1 5 4 2 3 4 MB,PP,JP,MB,BS,

WS,SS,C,AW

Timothy, mountain 4 4 4 4 5 4 5 4 S 3 4 3 4 3 3 5 5 5 4 4 5 3 SA,A,WM

Timothy 5 4 4 4 5 5 5 5 S 4 4 4 4 3 5 5 5 4 3 4 4 3 SA,A,WM,MB,PP,JP,MS

Trisetum, spike 2 2 4 3 4 3 4 3 S 5 4 3 4 3 2 3 3 3 3 3 2 3 MB,JP,MS,BS,WS

Wheatgrass, bluebunch 5 5 5 4 5 4 4 4 S 5 3 4 4 5 5 5 4 4 4 3 5 5 MB,PP,JP,MS,BS,

WS,C,AW

Wheatgrass, fairway 5 5 5 5 5 5 5 5 S 5 3 3 1 5 4 5 3 5 5 3 4 3 MB,PP,JP,MS,BS,WS,SS,

 crested BG,BB,C,AW

Wheatgrass, 5 5 5 5 5 5 5 5 V 5 4 3 1 5 5 5 4 5 5 4 4 5 A,MB,PP,JP,MS,BS,WS,

 intermediate SS,R,C,AW

Wheatgrass, pubescent 5 5 5 5 5 5 5 5 V 4 3 5 1 5 5 5 3 5 5 2 2 5 MB,PP,JP,MS,BS,WS,

SS,BB,R,C,AW

Wheatgrass, Siberian 3 5 5 5 5 4 4 4 S 5 2 2 2 5 3 5 3 5 4 2 2 4 BS,WS,SS,BG,BB,

 crested IS,C,AW

Wheatgrass, slender 4 5 5 5 4 5 4 4 S 4 4 3 4 3 4 5 4 4 4 4 3 5 SA,A,MB,PP,JP,MS

Wheatgrass, Snake River 5 5 5 5 5 5 4 4 S 5 3 4 4 5 5 5 4 4 4 2 2 5 MB,PP,JP,MS,BS,

WS,C,AW

Wheatgrass, 5 5 5 5 5 5 5 5 S 5 2 3 1 5 4 5 3 5 5 3 2 4 MB,PP,JP,MS,BS,WS,

 standard crested SS,BG,BB,C,AW

Wheatgrass, tall 5 5 5 5 5 5 5 4 S 4 3 3 2 4 5 3 2 4 4 3 5 4 MB,PP,JP,MS,BS,WS,

BG,IS,R,C,AW

Wheatgrass, thickspike 4 5 5 5 4 4 4 3 V 5 3 3 2 5 3 4 2 5 5 4 3 5 MB,PP,JP,MS,BS

Wheatgrass, western 3 5 5 5 3 3 3 5 V 5 3 3 3 5 3 4 3 5 5 4 3 5 MP,PP,JP,MS,BS,WS,

SS,BG,R,C,AW

Wildrye, Altai 5 5 5 4 2 2 3 5 V 4 4 3 3 4 5 4 3 5 5 4 4 3 MB,PP,JP,BG,IS,R

Wildrye, blue 4 5 5 4 5 5 4 3 S 4 4 3 3 4 5 4 4 2 4 3 3 2 A,MB

Wildrye, Canada 4 3 2 4 4 3 4 4 S 4 4 3 4 4 4 4 4 3 4 4 3 2 A,MB,WM,IS,R

Wildrye, alkali 3 4 5 4 4 4 4 4 V 4 4 2 3 4 4 4 3 4 5 2 3 2 WM,JP,BS,WS,SS,BG,IS

Wildrye, Great Basin 5 4 5 4 4 3 3 3 S 5 4 2 4 5 5 4 3 2 4 2 4 5 WM,MB,MS,BG,IS,

R,C,AW

Wildrye, mammoth 1 3 5 4 3 3 3 2 V 4 4 2 4 5 5 2 2 4 4 2 4 4 MB,JP,MS,BS

Wildrye, Russian 4 4 5 5 4 3 3 5 S 5 4 4 4 5 4 5 5 5 4 2 2 4 MS,JP,MS,BS,WS,SS,

BG,BB,IS,C,AW

Wildrye, Salina 2 4 5 4 1 2 2 4 V 4 3 2 2 5 2 1 1 5 5 2 3 2 MB,PP,JP,MS,BS,WS,

SS,BG,IS,C,AW

a

Vegetative types to which the species is adapted: A = Aspen-conifer; AW = Annual weeds; BB = Blackbrush; BG = Black greasewood; BS = Basin

big sagebrush; C = Cheatgrass; IS = Inland saltgrass; JP = Juniper-pinyon; MB = Mountain brush; MS = Mountain big sagebrush; PP = Ponderosa

pine; R = Riparian; SA = Subalpine; SS = Shadscale saltbush; WM = Wet and semiwet meadows; WS = Wyoming big sagebrush. b Key to ratings: 1 = Poor – difficult; 2 = Fair; 3 = Medium; 4= Good; 5 = Excellent – easy; A = Annual - reproduction from seed; S = Reproduction

from seed; V = Reproduction vegetative and from seed .

Table 2 (Con.) Species

Seed production

Ease of seed harvesting

Ease of seed cleaning

Ease of seeding

Germination

Seedling vigor

Growth rate

Final establishment

Method of spread

Spring growth

Summer growth

Fall growth

Compatibility with other species

Longevity

Forage production

Spring palatability

Summer palatability

Grazing tolerance

Soil stabilization

Shade tolerance

Flood tolerance

Ecotypic variability

Vegetative typesa
310 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses Table 3—Scientific names, synonyms, and common names of grass species. ND = Species not described in reference. Scientific name Common name Barkworth and Dewey (1985), Hitchcock (1951) Dewey (1984) Arnow (1987) USDA NRCS (2002) Plummer and others (1977) USDA NRCS (2002) Agropyron cristatum Agropyron cristatum Agropyron cristatum Agropyron cristatum Wheatgrass, fairway crested Wheatgrass, crested Agropyron dasystachyum Elymus lanceolatus Elymus lanceolatus Elymus lanceolatus Wheatgrass, thickspike Wheatgrass, streambank Agropyron desertorum Agropyron desertorum Agropyron desertorum Agropyron desertorum Wheatgrass, standard crested (desert) Wheatgrass, desert Agropyron elongatum Thinopyrum elongatum Elymus elongatus Thinopyrum ponticum Wheatgrass, tall Wheatgrass, rush Agropyron inerme Pseudoroegneria spicata Elymus spicatus Pseudoroegreria spicata ssp. inermis Wheatgrass, beardless bluebunch Wheatgrass, beardless Agropyron intermedium Thinopyrum intermedium Elymus hispidus Thinopyrum intermedium Wheatgrass, intermediate Wheatgrass, intermediate Agropyron junceum Thinopyrum junceiforme ND Thinopyrum junceiforme Wheatgrass, rushleaf Wheatgrass, Russian Agropyron repens Elytrigia repens Elymus repens Elymus repens Quackgrass Quackgrass Agropyron riparium Elymus lanceolatus Elymus lanceolatus Elymus lanceolatus ssp. lanceolatus Wheatgrass, steambank Wheatgrass, steambank Agropyron scribneri Elymus scribneri Elymus scribneri Elymus scribneri Wheatgrass, Scribner Wheatgrass, spreading Agropyron sibiricum Agropyron fragile Agropyron cristatum Agropyron fragile Wheatgrass, Siberian Wheatgrass, Siberian Agropyron smithii Pascopyrum smithii Elymus smithii Pascopyrum smithii Wheatgrass, western Wheatgrass, western Agropyron spicatum Pseudoroegneria spicata Elymus spicatus Pseudoroegneria spicata ssp. spicata Wheatgrass, bluebunch Wheatgrass, bluebunch Agropyron subsecundum Elymus trachycaulus Elymus trachycaulus Elymus trachycaulus ssp. subsecundus Wheatgrass, bearded Wheatgrass, slender Agropyron trachycaulum Elymus trachycaulus Elymus trachycaulus Elymus trachycaulus ssp. trachycaulus Wheatgrass, slender Wheatgrass, slender Agropyron trichophorum Thinopyrum intermedium Elymus hispidus Thinopyrum intermedium Wheatgrass, pubescent Wheatgrass, intermediate Agrostis alba ND Agrostis stolonifera Agrostis gigantea Bentgrass, redtop Redtop Agrostis stolonifera ND Agrostis stolonifera Agrostis stolonifera Bentgrass, carpet Bentgrass, creeping Alopecurus arundinaceus ND ND Alopecurus arundinaceus Foxtail, reed Foxtail, creeping meadow Alopecurus pratensis ND Alopecurus pratensis Alopecurus pratensis Foxtail, meadow Foxtail, meadow Aristida longiseta ND Aristida purpurea Aristida purpurea var. longiseta Threeawn, red Threeawn, fendler Arrhenatherum elatius ND Arrhenatherum elatius Arrhenatherum elatius Oatgrass, tall Oatgrass, tall Bouteloua curtipendula ND Bouteloua curtipendula Bouteloua curtipendula Grama, sideoats Grama, sideoats Bouteloua gracilis ND Bouteloua gracilis Bouteloua gracilis Grama, blue Grama, blue Bromus anomalus ND Bromus anomalus Bromus anomalus Brome, nodding Brome, nodding Bromus biebersteinii ND Bromus biebersteinii Bromus biebersteinii Brome, Regar (meadow) Brome, meadow Bromus carinatus ND Bromus carinatus Bromus carinatus Brome, mountain Brome, California Bromus erectus ND ND Bromus erectus Brome, meadow Brome, erect Bromus inermis ND Bromus inermis Bromus inermis Brome, smooth Brome, smooth Bromus marginatus ND Bromus carinatus Bromus marginatus Brome, big mountain Brome, mountain Bromus pumpellianus ND Bromus inermis Bromus inermis ssp. pumpellianus Brome, Pumpelly Brome, Pumpelly Bromus rubens ND Bromus rubens Bromus rubens Brome, foxtail Brome, red Bromus tectorum ND Bromus tectorum Bromus tectorum Brome, cheatgrass Cheatgrass Calamagrostis canadensis ND Calamagrostis canadensis Calamagrostis canadensis Reedgrass, bluejoint Bluejoint Calamagrostis epigeios ND ND Calamagrostis epigeios Reedgrass, chee Reedgrass, chee Dactylis glomerata ND Dactylis glomerata Dactylis glomerata Orchardgrass Orchardgrass Deschampsia caespitosa ND Deschampsia cespitosa Deschampsia caespitosa Hair-grass, tufted Hairgrass, tufted Distichlis spicata ND Distichlis spicata Distichlis spicata Saltgrass, inland Saltgrass, inland Elymus angustus Leymus angustus ND Leymus angustus x Wildrye, Altai Wildrye, Altai Elymus aristatus ND ND Elyleymus aristatus Wildrye, purple Wildrye, purple

(Elymus elymoides x Leymus triticoides)

Elymus canadensis Elymus canadensis Elymus canadensis Elymus canadensis Wildrye, Canada Wildrye, Canada

Elymus cinereus Leymus cinereus Elymus cinereus Leymus cinereus Wildrye, Great Basin Wildrye, basin

Elymus flavescens Leymus flavescens ND Leymus flavescens Wildrye, yellow Wildrye, yellow

Elymus giganteus Leymus racemosus ND Leymus racemosus Wildrye, mammoth Wildrye, mammoth

Elymus glaucus Leymus glaucus Elymus glaucus Elymus glaucus Wildrye, blue Wildrye, blue

Elymus junceus Psathyrostachys juncea Elymus junceus Psathyrostachys juncea Wildrye, Russian Wildrye, Russian

Elymus sabulosus ND ND Leymus racemosus ssp. sabulosa Wildrye, sabulosa Wildrye, mammoth

Elymus salinus Leymus salinus Elymus salinus Leymus salinus ssp. salinus Wildrye, salina Wildrye, saline

(con.)
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Chapter 18 Grasses Elymus simplex Leymus simplex Elymus simplex Leymus simplex Wildrye, low creeping Wildrye, alkali Elymus triticoides Leymus triticoides Elymus triticoides Leymus triticoides Wildrye, creeping Wildrye, beardless Festuca arizonica ND Festuca ovina var. Festuca arizonica Fescue, Arizona Fescue, Arizona arizonica Festuca arundinacea ND Festuca arundinacea Lolium arundinaceum Fescue, reed (Alta, tall) Fescue, tall Festuca elatior ND Festuca pratensis Lolium pratense Fescue, meadow Ryegrass, meadow Festuca idahoensis ND Festuca ovina var. Festuca idahoensis Fescue, Idaho Fescue, Idaho ingrata Festuca ovina var. ND Festuca ovina var. Festuca trachyphylla Fescue, hard sheep Fescue, hard duriuscula duriuscula Festuca ovina var. ovina ND Festuca ovina var. ovina Festuca ovina Fescue, sheep Fescue, sheep Festuca thurberi ND Festuca thurberi Festuca thurberi Fescue, Thurber Fescue, Thurber Hesperochloa kingii ND Leucopoa kingii Leucopoa kingii Fescue, spike Fescue, spike Hilaria jamesii ND Hilaria jamesii Pleuraphis jamesii Galleta Galleta, James Hordeum brachyantherum Hordeum brachyantherum Hordeum brachyantherum Hordeum brachyantherum Barley, meadow Barley, meadow Hordeum bulbosum ND ND Hordeum bulbosum Barley, bulbous Barley, bulbous Hordeum jubatum Hordeum jubatum Horedum jubatum Hordeum jubatum Barley, foxtail Barley, foxtail Hordeum vulgare ND Horedum vulgare Hordeum vulgare Barley, beardless Barley, common Koeleria cristata ND Koeleria macrantha Koeleria macrantha Junegrass, prarie Junegrass, prarie Lolium multiflorum ND Lolium perenne Lolium perenne ssp. multiflorum Ryegrass, Italian Ryegrass, Italian Lolium perenne ND Lolium perenne Lolium perenne Ryegrass, perennial Ryegrass, perennial Melica bulbosa ND Melica bulbosa Melica bulbosa Oniongrass Oniongrass Oryzopsis hymenoides ND Stipa hymenoides Achnatherum hymenoides Ricegrass, Indian Ricegrass, Indian Phalaris arundinacea ND Phalaris arundinacea Phalaris arundinacea Canarygrass, reed Canarygrass, reed Phleum alpinum ND Phleum alpinum Phleum alpinum Timothy, alpine Timothy, alpine Phleum pratense ND Phleum pratense Phleum pratense Timothy Timothy Poa ampla ND Poa secunda Poa secunda Bluegrass, big Bluegrass, Sandberg Poa bulbosa ND Poa bulbosa Poa bulbosa Bluegrass, bulbous Bluegrass, bulbous Poa canbyi ND Poa secunda Poa secunda Bluegrass, Canby Bluegrass, Sandberg Poa compressa ND Poa compressa Poa compressa Bluegrass, Canada Bluegrass, Canada Poa cusickii ND Poa fendleriana Poa cusickii Bluegrass, Cusick Bluegrass, Cusick Poa fendleriana ND Poa fendleriana Poa fendleriana Bluegrass, mutton Muttongrass Poa longiligula ND Poa fendleriana Poa fendleriana ssp. longiligula Bluegrass, longtongue Muttongrass Poa nevadensis ND Poa secunda Poa secunda Bluegrass, Nevada Bluegrass, Sandberg Poa pratensis ND Poa pratensis Poa pratensis Bluegrass, Kentucky Bluegrass, Kentucky Poa reflexa ND Poa reflexa Poa reflexa Bluegrass, nodding Bluebrass, nodding Poa scabrella ND Poa secunda Poa secunda Bluegrass, pine Bluegrass, Sandberg Poa secunda ND Poa secunda Poa secunda Bluegrass, Sandberg Bluegrass, Sandberg Secale montanum ND Secale montanum ND Rye, mountain ND Sitanion hystrix Elymus elymoides Elymus elymoides Elymus elymoides Squirreltail, bottlebrush Squirreltail Sporobolus airoides ND Sporobolus airoides Sporobolus airoides Sacaton, alkali Sacaton, alkali Sporobolus cryptandrus ND Sporobolus cryptandrus Sporobolus cryptandrus Dropseed, sand Dropseed, sand Stipa columbiana ND Stipa nelsonii Achnatherum nelsonii Needlegrass, subalpine Needlegrass, Columbia Stipa comata ND Stipa comata Hesperostipa comata Needle-and-thread Needle-and-thread Stipa lettermanii ND Stipa lettermanii Achnatherum lettermanii Needlegrass, Letterman Needlegrass, Letterman Stipa thurberiana ND Stipa thurberiana Achnatherum thurberianum Needlegrass, Thurber Needlegrass, Thurber Stipa viridula ND Stipa viridula Nassela viridula Needlegrass, green Neddlegrass, green Trisetum spicatum ND Trisetum spicatum Trisetum spicatum Trisetum, spike Tristetum, spike Table 3 (Con.) Scientific name Common name Barkworth and Dewey (1985), Hitchcock (1951) Dewey (1984) Arnow (1987) USDA NRCS (2002) Plummer and others (1977) USDA NRCS (2002)
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Chapter 18 Grasses

and about 0.04 inch (l mm) long. The spike is 0.8 to 4

inches (2 to 10 cm) long and 0.2 to 0.9 inch (5 to 23 mm)

wide. Spikelets are several times longer than the

internodes. Each produces 3 to 10 florets, but the

upper florets are sterile. Glumes and lemmas are

tapered to a short awn about 0.12 inch (3 mm) long.

Arnow (1987), Brown (1979), Cronquist and others

(1977), Hitchcock and others (1969), Gould (l951),

Looman (1982), and Powell (1994) provide detailed

descriptions of the species.

Although fairway crested wheatgrass resembles standard crested wheatgrass and Siberian wheatgrass,

some differences can usually be observed (fig. 1). The

major taxonomic feature used to differentiate the

three grasses is the spike shape. Fairway crested

wheatgrass spikes are shorter, wider at the base, and

more distinctly tapered at the tip than those of standard crested wheatgrass. Spikes of Siberian wheatgrass are relatively long and narrow compared to

those of the other two wheatgrasses. Fairway crested

wheatgrass spikelets and awns are divergent and

arranged in a “comblike” manner, while those of standard crested wheatgrass and Siberian wheatgrass

grow at a strong upward angle. Compared to fairway

crested wheatgrass, the stems of standard crested

wheatgrass are more erect and more or less uniform in

height, its leaves are mostly basal, and the upper leaf

blades are generally without hairs.

Ecological Relationships and Distribution

Problems relative to the proper identification and

use of the crested wheatgrasses arose, in part, from

the failure of earlier workers to identify them correctly

and to recognize their differing ecological requirements. In addition, although the name “crested wheatgrass” refers to the comblike spike of fairway crested

wheatgrass, this common name has been applied to all

three taxa. Dillman (1946) suggested that some of the

confusion resulted from a 1910 report he prepared in

which both fairway crested wheatgrass and standard

crested wheatgrass were referred to as A. cristatum

(Dillman 1910). This was compounded in 1935 when

Hitchcock (1935) used a drawing of A. desertorum to

illustrate A. cristatum in the “Manual of the Grasses

of the United States.”

To address these problems, Weintraub (1953) applied the following common names: crested wheatgrass = A. cristatum; desert wheatgrass = A. desertorum; and Siberian wheatgrass = A. sibiricum. Beetle

(1961) recommended acceptance of this usage, but the

confusion continued. Dewey (1986) later suggested that

the most appropriate names for the three groups in

North America are fairway crested wheatgrass (A. cristatum), standard or desert crested wheatgrass (A. desertorum),and Siberian crested wheatgrass (A. sibiricum)

(fig. 1). Differences between Eurasian and North

American taxonomic treatments further compound

this situation and have yet to be resolved.

The crested wheatgrasses constitute a polyploid

complex (Dewey 1986; Taylor and McCoy 1973). Diploid (2n = 14) fairway crested wheatgrass is native to

the Steppe region of Eastern Russia and Southwestern Siberia (Asay 1995a,b; Dewey 1966, 1969, 1973;

Dewey and Asay 1982; Konstantinov 1920). Dewey

and Asay (1982) identified different morphological

growth forms of diploid introductions, but found that

they hybridized readily. Tetraploids (2n = 28) include

standard and Siberian wheatgrass and are widely

distributed from central Europe to the Middle East,

Siberia, Mongolia, and China (Asay and Dewey 1979;

Dewey 1986; Knowles 1955; Tsvelev 1976). The hexaploids (2n = 42) are less common and more scattered,

occurring in Turkey, Iran, and Kazakhstan (Asay and

others 1990; Dewey and Asay 1975). All possible crosses

have been made among the three ploidy levels (Asay

and Dewey 1979; Asay and Knowles 1985a,b; Dewey

1971, 1974), but breeding has generally been confined

to selections within ploidy levels, particularly the

diploid and tetraploid levels (Asay 1986).

Crested wheatgrass was first introduced in the

United States by the U.S. Department of Agriculture

in 1889 (Dillman 1946; Reitz and others 1936). Materials were obtained from the Valuiki Experiment Station, Samara Government, on the Volga River, some

150 miles (240 km) north of St. Petersburg, Russia.

Figure 1—Spikelets of fairway crested wheatgrass

(center) are divergent and comblike, while those of

standard crested wheatgrass (left) and Siberian crested

wheatgrass (right) grow upward (RMRS photo).
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Chapter 18 Grasses

Additional seed was obtained from the Valuiki Station

in 1906. This shipment included six seed lots of 4 to 5

pounds (1.8 to 2.3 kg) each. One lot, S.P.I. 19536, was

labeled A. cristatumand five lots, S.P.I. 19537 to19541,

A. desertorum. Plantings and research with crested

wheatgrass began in 1908 at the Belle Fourche Station, Newell, SD, and in 1915 at the Northern Great

Plains Field Station in Mandan, ND (Dillman 1946).

Studies initiated in Canada greatly advanced the

development and use of crested wheatgrass (Kirk

1928). Kirk and others (1934) reported that seed

obtained from the U.S. Department of Agriculture was

planted in University of Saskatchewan experimental

plots. Additional introductions were obtained from

Russia, Siberia, and Mandan, ND. In 1927, seed obtained from Montana was planted at Manyberries,

Alberta. The University of Saskatchewan developed

two strains. One, identified as ‘Fairway’ (S-10), ultimately gained widespread use (Kirk and others 1934).

Fairway was initially distributed by the Field Husbandry Department, University of Saskatchewan, in

1927, and licensed in 1932 as a variety (Kirk 1932;

Knowles 1956). Dillman (1946) concluded that the

Fairway variety originated from the same accession

S.P.I. 19536, A. cristatum.

Scientists working in Montana recognized that the

erect bunch-type growth forms of A. desertorum differed from fairway-type crested wheatgrass. The Montana Station was the first to apply the name “standard

crested wheatgrass” for certified seed increased from

S.P.I. 19537, an A. desertorum accession (Dillman

1946). The selection was used as a standard of comparison for assessment of other selections (Dewey

1986). The name “standard crested wheatgrass” slowly

evolved to distinguish any nonfairway-type crested

wheatgrass. Standard is not a specific cultivar, but is

now used to identify all standard-type A. desertorum.

Seed of standard wheatgrass was first produced commercially by a grower in Montana shortly after 1920,

and was first listed in 1927 by a dealer in North

Dakota (Westover and others 1932).

Plant Culture

Good seed crops of fairway crested wheatgrass are

regularly produced on irrigated farmlands in Western

Canada, the Pacific Northwest, the Intermountain

area, and the Great Plains (Hafenrichter and others

1949; Knowles 1956; Reitz and others 1936; Westover

and Rogler 1947; Westover and others 1932). In these

areas, dryland farms, pastures, and rangeland

plantings are also managed for seed production. Seed

is harvested from these plantings only during years

with adequate moisture, but large quantities of seed

may be harvested in good precipitation years. Yields

from irrigated fields average 300 to 600 pounds per

acre (325 to 675 kg per ha), while yields from dryland

stands produce 250 to 300 pounds per acre (275 to 325

kg per ha) (Westover and others 1932). Less than 100

pounds per acre (110 kg per ha) are normally harvested from rangeland stands in the Intermountain

area. Row spacings of 36 to 42 inches (91 to 107 cm) are

recommended for cultivated fields. This equates to 5 to

8 pounds of seed per acre (5.5 to 9.5 kg per ha).

Planting at 3 to 4 pounds per acre (3.4 to 4.5 kg per ha)

provides satisfactory, but slower developing, stands

(Westover and Rogler 1947). Direct combining provides seed lots with purities of 50 to 80 percent; this

may be increased to 95 percent purity by screening and

fanning. Germination is often near 90 percent. Seed

can usually be stored in a warehouse for several years

without loss of viability.

Fairway crested wheatgrass is one of the most reliable plants to establish from drill seedings on adapted

range and wildland sites. It generally develops fairly

uniform stands, even on highly variable sites. Maynard

and Gates (1963) reported that establishment success

of crested wheatgrass could be attributed to the tolerance of the seeds to extreme fluctuations in soil moisture. Wetting and drying cycles can increase germination. Plummer (1943) reported that the success of

crested wheatgrass stems from its ability to produce a

greater total root length in the seedling stage than

other grass species.

Drill seeding rangeland sites with 2 to 16 pounds of

fairway crested wheatgrass seed per acre (2.2 to 17.6

kg per ha) was initially advised for sites receiving

more than 12 inches (310 mm) of precipitation, and 5

to 6 pounds per acre (5.5 to 6.6 kg per ha) for sites

receiving less than 12 inches (310 mm) of moisture

(Thomas and others 1945). More recently, seeding

recommendations have been reduced to 2 to 4 pounds

per acre (2.2 to 4.4 kg per ha) for either drilling or

broadcast seeding. When fairway crested wheatgrass

is seeded alone in areas invaded by weeds, the rate

should be increased 2 to 3 times. Seeding in dense

annual weeds, however, is not advisable. Springfield

(1965) reported that crested wheatgrass stands will

reach an equilibrium within 5 to 8 years, regardless of

the amount of seed sown or the drill row spacing used.

He found similar herbage yields on sites seeded at 2, 4,

and 6 pounds seed per acre (2.2, 4.4, and 6.6 kg per ha)

at 5-, 12-, and 18-inch (13-, 30-, 46-cm) row spacings in

the fifth, sixth, and eighth years after seeding.

Fairway crested wheatgrass seed should be drilled

or otherwise planted at depths of 0.5 to 1.0 inch (1.3 to

2.5 cm). Planting at greater depths in heavy-textured

soils is not advised. Compared with many other rangeland grasses, fairway crested wheatgrass establishes

better with limited seedbed preparation. McGinnies

(1962), however, found that packing to create a firm

seedbed improved planting success. Cultipacking before seeding provides a firm seedbed, permitting furrow openers to function more effectively. Broadcast
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Chapter 18 Grasses

seeding without some means of seed coverage is not

recommended. Direct drilling or broadcast seeding

and covering the seed following burning or removal of

competition is generally successful. Stevens (1987b)

reported that fairway crested wheatgrass plants establish extremely well following aerial seeding and

chaining on pinyon-juniper and big sagebrush disturbances.

Fall seedings are recommended, but spring plantings

on prepared seedbeds may be successful if adequate

spring precipitation is received. Frischknecht (1951)

found that the total emergence from crested wheatgrass and other grass seedings was greater from early

fall than from late fall or spring plantings, and that

cold temperatures accounted for most grass seedling

mortality. Buman and Abernethy (1988) found that

mountain rye, crested wheatgrass, and cheatgrass

have similar germination temperature requirements,

which accounts for the ability of fairway crested

wheatgrass seedlings to compete favorably with

cheatgrass. Asay and Johnson (1983) found that genetic differences among populations of crested wheatgrass accounted for over 50 percent of the total phenotypic variation in traits affecting seeding

establishment. Seed weight was closely correlated to

seeding establishment.

Established stands of fairway crested wheatgrass

may produce seed crops annually, depending on precipitation, but plants do not spread rapidly to adjacent

areas. Seedlings of this species quickly occupy openings created in fairway crested wheatgrass stands.

When seeded in mixtures with other grasses, fairway

crested wheatgrass often increases in density at the

expense of the other species. Walker and others (1995),

reporting on long-term studies of seedings in pinyonjuniper sites in central Utah, found that fairway crested

wheatgrass density increased at a higher rate than

native bunchgrasses during periods of normal precipitation, but native bunchgrasses increased in density

and ground cover at a greater rate during years of low

precipitation. They found that over a 23-year period,

native grasses were not able to compete with introduced species including fairway crested wheatgrass.

Broadcast seeding fairway crested wheatgrass at low

rates of 1 to 2 pounds per acre (1.1 to 2.2 kg per ha) in

pinyon-juniper and big sagebrush disturbances has

been employed to encourage the recovery of native

species. Although native herbs and shrubs increase

initially in some situations, slow but persistent increase of fairway crested wheatgrass followed, and

within 25 to 40 years it gained dominance.

Uses and Management

Fairway crested wheatgrass has been extensively

used in pasture and rangeland plantings in the United

States and Canada. In Western Canada, it is used far

more than standard crested wheatgrass (Westover

and Rogler 1947). In the United States, the Civilian

Conservation Corps used both fairway and standard

crested wheatgrass extensively for conservation

plantings in the 1930s. Use increased in 1945 following a special Congressional Appropriation for research

to improve rangelands (Reynolds and Springfield 1953).

Fairway crested wheatgrass is well adapted to the

Northern and Central Great Plains and the drier

communities of the Intermountain region. Fairway

and standard crested wheatgrass are seeded to stabilize marginal farmlands and deteriorated rangelands

in the Western United States. Fairway crested wheatgrass has been used in mixtures with other grasses

and alfalfa for hay production. It has also been planted

alone in lawns, yards, and on roadways where supplemental irrigation cannot be provided (Knowles 1956).

Fairway crested wheatgrass cultivars are best

adapted to ponderosa pine, mountain brush, pinyonjuniper, and benchland sites occupied by big sagebrush. They are also adapted to open, exposed slopes

with intermixed aspen and spruce-fir communities.

Reitz and others (1936) reported that fairway crested

wheatgrass persists on dry, open areas at elevations

up to 6,800 ft (2,070 m). Plantings in central Utah have

established well on open slopes at elevations near

8,000 ft (2,440 m).

On adapted sites, fairway crested wheatgrass becomes dormant during dry periods, but it will resume

growth in fall or spring when adequate moisture

becomes available. It tolerates a range of soil textures

and pH levels, and withstands saline conditions, but it

will not survive flooding for more than a few days.

Fairway crested wheatgrass grows better in moderate

shade than either standard crested wheatgrass or

Siberian wheatgrass, but it does not persist in dense

shade. It can establish even with some grazing during

seedling establishment (Johnson 1986), although grazing of new seedings is not advised. Mature plants can

persist with heavy and repeated grazing. Resistance

to grazing abuse is a primary factor that has promoted

seedings of this grass.

Fairway crested wheatgrass is generally less well

adapted to low-elevation shrubland communities occupied by Wyoming big sagebrush and salt desert

species than are other crested wheatgrasses. Fairway

crested wheatgrass plantings have established and

persisted in arid regions for a number of years, but

some of these have been lost during droughts. The

species is suited to sagebrush communities that receive more than 10 inches (250 mm) of precipitation,

but it should not be planted in communities receiving

less than this amount. In the Southwest and areas

with long warm summers, it will not persist unless

annual precipitation exceeds 15 inches (380 mm)

(Reynolds and Springfield 1953).
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Chapter 18 Grasses

Thousands of acres of abandoned farmland and

overgrazed rangelands have been seeded to fairway

crested wheatgrass to provide spring-summer forage

for livestock, soil stabilization, and weed control

(Barnes and others 1952; Bridges 1942; Christ 1934;

Hull 1974; Knowles 1956; Westover and Rogler 1947).

In some situations, fairway crested wheatgrass has

been seeded as a replacement for native grasses (Stark

and others 1946). Many of these areas were disked or

burned to remove existing competitive vegetation prior

to seeding (Malechek 1986). In addition, extensive

areas dominated by big sagebrush have been plowed,

disked, burned, or sprayed to remove the shrubs, then

seeded with this species and other introduced perennials to enhance livestock grazing (Plummer and others

1955; Robertson and others 1966). Fairway crested

wheatgrass seedings have prevented the recovery of

big sagebrush where vigorous stands of the grass have

developed and the sagebrush seed source has been

eliminated. Natural recovery of sagebrush, however,

has occurred in other areas.

If seeded in mixtures, fairway crested wheatgrass

can be drill seeded with other introduced grasses and

broadleaf herbs that produce vigorous seedlings. It is

capable of persisting with competitive sod-forming

perennials, including intermediate wheatgrass and

smooth brome. Although these two sodforming grasses

may prevent the spread of crested wheatgrass, a

bunchgrass, they do not eliminate it. Fairway crested

wheatgrass is often planted in mixtures with alfalfa in

mountain brush, ponderosa pine, and other upland

communities to provide summer forage for game and

livestock. Once established, both species persist if

properly managed. Production of fairway crested

wheatgrass may be enhanced by the presence of alfalfa. Fairway crested wheatgrass is also frequently

seeded with alfalfa and late-maturing grasses to maintain herbage quality and diversity (Mayland 1986;

USDA Forest Service 1937).

Fairway crested wheatgrass can be broadcast seeded

with most revegetation species, including those that

are slower to develop if appropriate planting designs

are adopted. Drill seeding fairway crested wheatgrass

with slower developing native shrubs and herbs is not

recommended unless provisions are made to seed the

species in separate rows or strips that are adequately

spaced to minimize competition (Hubbard 1957). Planting bitterbrush in rows less than 2 ft (0.6 m) from

fairway crested wheatgrass rows, for example, was

found to significantly decrease vigor and growth of the

shrubs (Hubbard and others 1962).

Fairway crested wheatgrass can be successfully

interseeded in cheatgrass using conventional, deep

furrow, rangeland drills that create openings about 4

to 6 inches (10 to 15 cm) wide. This reduces competition and permits establishment of the perennial grass.

Areas with dense populations of annual weeds should

be avoided. Fairway crested wheatgrass plantings

have also been important in the control and elimination of halogeton and Russian thistle (Mathews 1986;

Mayland 1986). The species has been particularly

useful in replacing cheatgrass and slowing the spread

of wildfires (Monsen and Kitchen 1994). Few other

perennial grasses establish as well as crested wheatgrasses amid weedy competition.

Crested wheatgrass begins growth in early spring

and provides considerable herbage during this period

(Malechek 1986). Nutritive value is good for energy

and fair for protein (Dittberner and Olson 1983).

Crude protein, calcium, phosphorus, and ash decrease

as plants mature in late May and June (Mayland

1986). The degree of use is rated as good for livestock

and fair for wildlife (Dittberner and Olson 1983).

Fairway crested wheatgrass has stabilized eroding

areas, reduced runoff, and provided a stable vegetative cover on many disturbed sites. It performs well as

a pioneer species capable of stabilizing exposed substrata, and it does not require heavy fertilizer applications to maintain an acceptable cover. It is frequently

used in big sagebrush communities to control weed

invasions and stabilize watersheds, roadways, and

mine disturbances. It furnishes soil protection on sites

where high-intensity storms are common.

In many cases, however, fairway crested wheatgrass seedings have adversely impacted wildlife habitat. Conversion of native shrub and herb communities

to solid stands of this grass have reduced cover and

forage for big game and nongame animals (Marlette

and Anderson 1984). Fairway crested wheatgrass

seedings provide important supplements to deer diets

in spring and fall. Elk, pronghorn antelope, and bighorn sheep also use it (Urness l986). These seedings,

however, do not afford the diversity and quality of

forage provided by native communities. A primary

concern in seeding fairway crested wheatgrass is the

resulting monotypic stands that often develop, reducing species diversity and habitat quality (DePuit 1986;

Marlette and Anderson 1986). Development of technology for replacing fairway crested wheatgrass with

more diverse native species to improve habitat characteristics is becoming a management objective in many

areas.

Single species plantings of fairway crested wheatgrass normally persist as a monoculture, although

invasion of a few other species may occur after 15 to 20

years. Marlette and Anderson (1984) found that invading species seldom comprise more than 10 percent

of the total plant density. Stands of fairway crested

wheatgrass have persisted for over 50 years in the

Great Plains (Mayland 1986), and for over 30 years in

the Intermountain West (Marlette and Anderson 1984;

Plummer and others 1968). This species does not
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Chapter 18 Grasses

invade or migrate into disturbances of early successional communities occupied by annual weeds, and it

has made only minor migrations into climax grasslands (Allred 1940).

Varieties and Ecotypes

Since its introduction in 1906, the crested wheatgrass complex has been studied intensively. Cultivars

have been developed for a broad range of planting sites

and conditions (Asay 1986; Dillman 1946). Selection

and breeding programs are currently underway to

select strains that are more drought tolerant and

productive, and that remain green longer in summer.

Rhizomatous forms of crested wheatgrass are being

developed to provide low maintenance turf and roadway plantings. Breeding has been primarily within

the diploid or tetraploid levels, but hybrids between

standard crested wheatgrass and induced tetraploid

fairway crested wheatgrass developed by Agricultural

Research Service scientists in Logan, UT, are highly

competitive on semiarid rangeland sites.

Scientists at the Agricultural Research Service, Crops

Research Laboratory in Logan, UT, developed ‘CD-II’

crested wheatgrass from the original ‘Hycrest’ hybrid.

Selections were made from Hycrest foundation seed

fields based on vegetative vigor, absence of purple

leaves in spring, freedom from disease and insects,

and leafiness. From these clonal lines, a second evaluation was conducted for the same vegetative characteristics and for individual seed weight and emergence

of polycross seed lots from deep seedings.

CD-II is leafier than Hycrest and produces more

forage at low spring temperatures. Stand establishment and adaptation to arid conditions are comparable to Hycrest. Seed production is excellent, ranging

from 600 to 800 pounds per acre (660 to 940 kg per ha).

CD-II is recommended for semiarid sites at elevations

up to 7,150 ft (2,345 m) receiving from 8 to 18 inches

(200 to 450 mm) of annual precipitation (Asay and

others 1997).

‘Douglas’ crested wheatgrass, developed by the Agricultural Research Service in Logan, UT, and released in 1994, is the first hexaploid (2n = 42) cultivar

of crested wheatgrass released in North America (Asay

and others 1995a) (fig. 2). It was named in honor of

Dr. Douglas R. Dewey, who initiated research on the

perennial Triticeae at the Logan laboratory. Douglas

crested wheatgrass was derived from hybrids between

a broad-leaved hexaploid accession from the former

Soviet Union and hexaploid lines from Iran and Turkey (Asay and others 1996). Douglas was selected for

its leafiness, retention of green leaves into summer,

palatability to wildlife and livestock, and high in vitro

digestibility. It produces larger seeds than diploid or

tetraploid cultivars. Douglas should be fall seeded on

sites receiving at least 12 to 15 inches (300 to 376 mm)

of annual precipitation. Its seedling vigor and ability

to establish from a 3-inch (7.6-cm) planting depth were

greater than for ‘Nordan’, Fairway, or ‘Ephraim’, and

equaled Hycrest (Asay and others 1996).

Douglas demonstrates promise for seeding in areas

dominated by cheatgrass that are prone to frequent

burning. It can he used to replace cheatgrass, and it

can be planted in strips as a fuel or fire barrier.

Douglas is less drought resistant than standard or

Hycrest crested wheatgrass, but it remains green

longer in summer, and plants are less flammable

during this period. Fires that ignite in cheatgrass may

burn into the firebreak provided by Douglas, but their

advance is slowed.

The Forest Service, Utah Division of Wildlife Resources, Natural Resource Conservation Service, and

the Agriculture Experiment Stations of Utah, Arizona, and Idaho released Ephraim crested wheatgrass

in 1983. It was developed from an introduction from

Ankara, Turkey, and planted by A. Perry Plummer at

an arid range site in central Utah in the early 1940s

where it survived for over 40 years. Seed and plants

from this site were tested at additional locations

throughout the Intermountain area (Plummer and

others 1968).

Ephraim was the first tetraploid cultivar of crested

wheatgrass released in the United States. It is a

persistent, sodforming cultivar that produces as much

herbage as Fairway. It is drought tolerant, and can be

seeded in areas receiving as little as 10 inches (250

mm) of annual precipitation. Ephraim has persisted

when planted in warm, dry areas of southern Utah and

northern Arizona. It is also well adapted to pinyonjuniper and big sagebrush communities of the Intermountain region. Plants establishing in moister areas

Figure 2—Mule deer preferred ‘Douglas’ fairway

crested wheatgrass to other cultivars of crested wheatgrass in a fall palatability study (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 317

Chapter 18 Grasses

produce rhizomes within 2 to 3 years (Stevens and

Monsen 1985).

Ephraim is recommended for seeding in pinyonjuniper, ponderosa pine, mountain brush, and big

sagebrush communities. It provides excellent ground

cover and protection on disturbed soils. It is highly

competitive with annual weeds, and can be used with

Nordan and other cultivars of standard crested wheatgrass to reduce or control cheatgrass, medusahead,

and other annual weeds. Seedlings of Ephraim are not

as vigorous as those of Hycrest, but once established,

they are extremely persistent in the face of drought,

cold winters, and heavy grazing. Ephraim furnishes

early spring herbage, and greens up in fall if late

summer or fall moisture is received. It is not shade

tolerant, but it does persist as an understory in big

sagebrush and other shrub communities.

Ephraim is a competitive cultivar. Established stands

have not allowed the entry of other perennials, even

when subjected to heavy and prolonged grazing.

Ephraim has been successfully seeded on mine disturbances in northern Nevada. It is also widely used to

plant roadways and other disturbances in the Intermountain region. It is well adapted to infertile soils

and moderately dry conditions.

Fairway crested wheatgrass was the first wheatgrass cultivar developed in North America. Released

in 1927, it was selected from P.I. 19536, an introduction received from Western Siberia and developed by

Agriculture Canada (Knowles 1956). It is shorter in

stature, leafier, more procumbent and winter hardy,

but less drought tolerant than most standard crested

wheatgrass cultivars. Although initially developed for

open parkways and turf plantings, Fairway has been

widely seeded throughout the Intermountain region,

and has proven better adapted to upland sites than

standard crested wheatgrass. It is widely used in the

mountain brush, ponderosa pine, pinyon-juniper, and

mountain big sagebrush zones. Fairway is extremely

long lived, and has been planted to provide forage,

produce ground cover, and prevent the invasion and

spread of weeds. Fairway seedlings are vigorous and

capable of establishing amid some competition, but

they are suppressed by dense cheatgrass. Fairway

reseeds itself well, as an abundance of seeds are

generally produced, even under semiarid conditions.

Fairway is well suited to a variety of site conditions

and is recommended for seeding in mixtures with sodforming grasses to provide erosion control. It produces

tillers and short rhizomes, and provides excellent

ground cover and high-quality forage. Herbage production of Fairway is usually less than that of standard crested wheatgrass, but Fairway produces a

greater proportion of leaves. With age, standard crested

wheatgrass develops an abundance of stems that become less palatable to grazing animals.

In 1984 the Agricultural Research Service in cooperation with the Utah Agricultural Experiment Station and the Natural Resources Conservation Service

released Hycrest crested wheatgrass, the first interspecific hybrid of crested wheatgrass. Hycrest was

developed by crossing an induced tetraploid of A.

cristatum with natural tetraploids of A. desertorum.

Reciprocal crosses were made to ensure that the cytoplasm of both species would be represented (Asay and

others l985). Open pollinated progenies were then

established at a nursery site in northern Utah. Eighteen clones were selected from these trials and planted

in isolated crossing blocks to provide the first generation of the synthetic strain (Asay and others 1985).

Seedlings of Hycrest are more competitive than

those of other cultivars. Hycrest competes well with

cheatgrass, halogeton, and other weeds. Seeded Hycrest

stands establish in competition with annual weeds;

consequently, this cultivar has been useful in seeding

sites where other weed-control measures cannot be

fully implemented. In plantings made across the Intermountain region, Hycrest has established better

stands and produced more forage during the first 2

years than either Nordan or Fairway (Alderson and

Sharp 1994). Examination of plantings indicates that

seedling vigor, stand establishment, and root growth

of Hycrest generally exceed those of other crested

wheatgrass cultivars. Hycrest has proven adapted to

big sagebrush and pinyon-juniper sites, but it is not as

drought tolerant as selections of standard or Siberian

crested wheatgrass.

A diploid cultivar, ‘Parkway’, was developed from

Fairway through several generations of recurrent selection for vigor, forage, and seed production (Alderson

and Sharp 1994). Parkway was released in

Saskatchewan in 1969 for use in hay and pasture

production. Its yields exceed those of Fairway, as the

plants are slightly taller, generally 19 to 30 inches (48

to 76 cm) in height. This cultivar has not been widely

planted on range or wildland sites in the Intermountain region, but it can be used on many of the same

sites as Fairway. It is not recommended for turf

plantings.

‘Ruff’ crested wheatgrass is a diploid cultivar jointly

released by the Agricultural Research Service and the

Nebraska Agricultural Experiment Station in 1974.

The parental material was derived from Fairway (Asay

1986). Ruff has a spreading bunchgrass habit and

short, leafy culms (Alderson and Sharp 1994). It begins growth in early spring and is valued as a shortseason pasture plant in drier areas of the Great Plains.

In the Central Great Plains, its forage production

equals that of Fairway (Alderson and Sharp 1994). Its

seeds are relatively small, but its seedlings establish

well from fall plantings. Ruff competes well with

annual and perennial weeds and can be seeded as a
318 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

conservation planting to protect soil and control weed

invasion in Intermountain areas where Fairway is

adapted. Ruff plantings become somewhat sodforming

due to the broad bunching habit and seeding within

the stand.

Agropyron dasystachyum

Thickspike Wheatgrass __________

Synonyms

Agropyron albicans

A. elmeri

A. griffthsii

A. riparium

Elymus lanceolatus

Description

Barkworth and Dewey (1985) transferred several

species of Agropyron into the genus Elymus, and

combined the former taxa A. dasystachyum, A.

dasystachyum var. elmeri, and A. dasystachyum var.

riparium with E. lanceolatus (Scribn. & Sm.) Gould.

Arnow (1987) also combined several Agropyron species with E. lanceolatus. Barkworth and others (1983)

and Barkworth and Dewey (1985) described four subspecies of thickspike wheatgrass:

E. lanceolatus ssp. albicans (Scribn. & Sm.) Barkw.

& D. R. Dewey

E. l. ssp. lanceolatus

E. l. ssp. psammophilus (Gillett & Senn) A. Love

E. l. ssp. yukonensis (Scribn & Merr.) Love

Taxa now included in treatments of E. lanceolatus

are similar in floral parts and growth form, thus

simplifying identification. These taxa, however, occupy quite different ecological sites. Montana wheatgrass (formerly A. albicans) and streambank wheatgrass (formerly A. riparium), for example, differ greatly

in areas of adaptation and require different management strategies.

Thickspike wheatgrass is a native, cool-season perennial (Barkworth and Dewey 1985; Hitchcock and

others 1969; Looman 1982; USDA Forest Service 1914;

Welsh and others 1987). The root system and long,

rapidly developing rhizomes form a sod that varies

with location and subspecies (USDA Forest Service

l937; Wambolt 1981) (fig. 3). Plants may develop a

dense and more or less continuous sod, or they may

occur as open, irregular patches intermixed with other

species. Stems are erect, 16 to 32 inches (40 to 80 cm)

tall, and most often bluish green. Leaf sheaths have

thin, overlapping edges that are smooth to slightly

roughened. Blades are 0.04 to 0.12 inch (1 to 3 mm)

wide, and 2 to 8 inches (5 to 20 cm) long. They are

usually stiff and ascending with slightly rolled edges.

The upper surface of the blade is furrowed, while the

lower surface is smooth. The upper surface and sometimes the base of the lower surface are roughened.

Auricles are about 0.08 inch (2 mm) long. The ligule is

membranous and up to 0.04 inch (1 mm) long with

ragged edges and a rounded apex. Inflorescences are

2.4 to 4.8 inches (6 to 12 cm) long. Spikelets are 0.4 to

0.6 inch (1 to1.5 cm) long, each with 3 to 12 loosely to

closely overlapping florets. Tips of the glumes are

subequal and firm, slightly roughened, and often with

small hairs. Glume tips are acute or extended into

short awns. Lemmas are 0.2 to 0.4 inch (6 to 9 mm)

long, convex on the back, firm, and five to seven

nerved. They are sparsely to densely hairy, and generally more hairy than the glumes. Lemma tips are

awnless or extended into tiny awns. The palea is about

as long as the lemma and obtuse at the tip.

Ecological Relationships and Distribution

Thickspike wheatgrass is widely distributed, occurring in Alaska, southward through the Western United

States and Northern Mexico, eastward across Canada

and much of the Great Plains, south to Kansas, and as

far east as West Virginia (Arnow 1987; Ross and

Figure 3—Thickspike wheatgrass is a widely distributed native, sod-forming wheatgrass (photo courtesy

of Kevin Jensen, USDA ARS, Logan, UT).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 319

Chapter 18 Grasses

Hunter 1976). It grows on sites ranging in elevation

from 4,000 to 10,900 ft (1,220 to 3,300 m) that receive

8 to 20 inches (200 to 500 mm) of annual precipitation.

Different ecotypes are adapted to soils that are well

drained and sandy, sandy loams, loams, or heavy

textured. It occurs on soils that are alkaline or weakly

saline.

In the Western United States, thickspike wheatgrass is common in sagebrush, pinyon-juniper, mountain brush, aspen, moist open parks, and dry meadows

(Wasser 1982). It occurs in desert and chaparral communities of the Southwest and in ponderosa pine and

western spruce forests of the Northwest (Arnow 1987;

Bernard and Brown 1977; Hitchcock and others l969;

Sours l983). It is often regarded as a disturbance

species, but it is also a major component of grama

grass/needlegrass, Idaho fescue, bluebunch wheatgrass, and mountain brush/aspen communities.

Thickspike wheatgrass is tolerant of moderate shade

provided by associated shrubs, but it is not adapted to

dense overstory cover. It recovers quickly following

fires, and often flourishes in sagebrush and juniper

communities following burning (Wright and Bailey

1982). It has persisted as a dominant species on

burned sites for over 30 years (Harniss and Murray

1973).

Plant Culture

Thickspike wheatgrass initiates growth in early

spring (Dittberner and Olsen 1983). New shoots emerge

as early as March 1; flowering occurs from May to July.

Seeds mature in late July and remain on the plant

until mid-August (Blaisdell 1958). Dispersal of mature seeds normally occurs over a 2- to 3-week period.

Seed from wildland stands is collected using reel-type

harvesters mounted on four-wheel-drive vehicles or by

hand stripping. Seed fields are harvested with conventional combines. Little debris is collected with the

seeds, as the culms extend above the leaves and the

seeds shatter readily. Purity of freshly harvested seed

lots usually exceeds 80 to 90 percent with about 85

percent germination (Wasser 1982). Most seed used in

wildland plantings is from named varieties grown

under cultivation. Few wildland stands are large

enough to support extensive harvesting. Seeds are

slightly larger than those of crested wheatgrass, averaging about 154,000 per pound (339,500 per kg)

(Wheeler and Hill 1957).

Seed germination is enhanced by a prechill of about

30 days, but some nonstratified seed will germinate in

about 20 days under laboratory conditions. Compared

to other wheatgrasses, germination and seedling development of thickspike wheatgrass is slow, but satisfactory stands generally develop over time. The seeding rate is sometimes increased to compensate for this

problem.

Fall seeding is recommended. Seeds should be planted

about 0.25 to 0.75 inch (0.6 to 1.9 cm) deep by drilling

or by broadcasting followed by harrowing. Thick stands

often develop when thickspike wheatgrass is seeded in

drill rows on well-prepared seedbeds. When broadcast

seeded with other species, thickspike wheatgrass tends

to form irregular patches that add diversity to wildlife

habitat. Development of new stands is aided by rhizome development. Seedlings of thickspike wheatgrass varieties compete well with annual weeds. The

species should not be seeded directly with slower

developing plants, but it can be seeded in alternate

rows or on adjacent sites.

Uses and Management

Thickspike wheatgrass is often seeded alone on

disturbances in big sagebrush and mountain brush

communities to stabilize soil and control erosion. It is

particularly well adapted to arid environments and

does well on disturbed, slightly alkaline soils. It is

seeded on sites such as mined areas, roadside disturbances, recreation areas, drainageways, and other

sites that receive little maintenance and where a low

sodforming grass is desired. Thickspike wheatgrass is

also commonly used to control the spread of weeds,

particularly annual grasses. It is often seeded for this

purpose in waste areas and nonirrigated portions of

cultivated fields.

Thickspike wheatgrass is widely used in range and

wildlife plantings. Herbage production is less than for

many other grasses, but it does provide excellent

spring-fall and early summer forage. It remains green

later into the summer than cheatgrass. Consequently,

it can be seeded to extend the period of grazing in areas

infested with this and other annuals. If moisture

becomes available in fall, plants will green up and

provide an important source of forage. Thickspike

wheatgrass recovers quickly following burning, providing herbage and ground cover. It also reduces the

spread of cheatgrass and other annual weeds as burned

areas recover.

Populations of thickspike wheatgrass may have restricted ranges of adaptation. Consequently, attempts

should be made to collect and seed local ecotypes. The

two released cultivars, ‘Critana’ and ‘Sodar’, may not

be adapted to all sites within the species range. Upland sites, including dry meadows and areas intermixed with aspen and conifer forests should be planted

with ecotypes adapted to these areas.

Varieties and Ecotypes

‘Bannock’ thickspike wheatgrass, a composite of six

collections from Oregon, Idaho, and Washington, was

released in 1995 by the Natural Resource Conservation Service Plant Materials Center at Aberdeen, ID.
320 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Bannock is a vigorous, leafy, long-lived, sodproducing

variety. Its rhizome growth averages about 4 inches

(10 cm) per year. It establishes quickly and develops a

protective sod on dry sites. It is a more effective

sodproducer than Sodar when seeded in the 8- to 10-inch

(200- to 250-mm) precipitation zone. Plants are 18 to

24 inches (45 to 60 cm) tall, and may grow to 40 inches

(100 cm) in height when irrigated. Leaves are abundant

and are produced 8 to 12 inches (20 to 30 cm) up the stem.

Bannock is recommended for seeding in mixtures on

arid rangeland sites and for conservation plantings to

provide soil protection. It is useful for revegetating

mined lands and other disturbances, and its highly

competitive sod reduces the spread of cheatgrass and

other annual weeds (USDA Natural Resourcess Conservation Service 2001). Inflorescences are large and

the florets are awnless. Seed yields range from 250 to

400 pounds per acre (275 to 440 kg per ha).

The Natural Resource Conservation Service Plant

Materials Center at Bridger, MT, and the Montana

and Wyoming Agriculture Experiment Stations cooperatively released Critana thickspike wheatgrass in

1971. D. E. Ryerson collected the original material

from a roadcut near Havre, MT, in 1960. Critana

provides a low-stature ground cover and a dense sod

that requires little maintenance. This cultivar exhibits excellent seedling vigor and establishes well, even

in areas receiving low rainfall. Plants are strongly

rhizomatous and produce low to moderate amounts of

herbage. Leaves are small, fine, and palatable. Plants

are tolerant of grazing and produce useful spring and

early summer forage. Critana is best adapted to the

Pacific Northwest, the Intermountain West, and the

Western Great Plains. It is commonly seeded in Montana, Wyoming, and the western Dakotas on rangelands and mine sites receiving 10 to 16 inches (250 to

400 mm) of annual precipitation (Thornburg 1982).

Critana can be used to stabilize infertile disturbances,

provide soil protection in areas where some traffic can

be expected, and control weeds. It has been successfully seeded and maintained along roadways, airport

runways, dry drainageways, recreational areas, and

mine disturbances (Alderson and Sharp 1994).

‘Schwendimar’ thickspike wheatgrass was released

in 1994 by the Natural Resources Conservation Service

Plant Materials Center at Pullman, WA. It was named

for John Schwendiman who collected seed from its site

of origin, a sandy, windblown area on the banks of the

Columbia River east of the Dallas, OR, in 1934.

Schwendimar is a rhizomatous, cool-season perennial

with bluish, glaucous culms. It develops a dense and

fibrous but shallow root system. Some roots extend

more than 2 ft (0.6 m) from the plant. Seeds are

relatively large. Seedlings establish well from spring or

fall plantings. They are no more vigorous than those of

other thickspike wheatgrass cultivars, but stands

develop quickly. Schwendimar is recommended for

areas with neutral, coarse-textured soils that receive 8

to 20 inches (200 to 500 mm) of annual precipitation.

The variety can be used to quickly stabilize well-drained,

droughty sites. It can also be seeded in mixtures to

provide mid to late summer forage for wildlife and

livestock. Plantings have persisted on a wide range of

sites in the Columbia River Basin, but the variety has

not seen extensive use outside this area (USDA Soil

Conservation Service 1994).

Sodar was released as a streambank wheatgrass

variety, but it is now considered a thickspike wheatgrass. The Idaho and Washington Agricultural Experiment Stations cooperatively released it in 1954.

Sodar was developed by several generations of mass

selection from material collected near Canyon City,

Grant County, OR, and grown at the Natural Resource

Conservation Service Plant Materials Center, Aberdeen, ID (Alderson and Sharp 1994).

Sodar is a rhizomatous, drought-resistant perennial, 20 to 30 inches (5 to 7.6 dm) tall that is used

primarily for erosion control and conservation

plantings. It is more drought tolerant and widely

adapted to big sagebrush communities than Critana.

In the Intermountain region, Sodar is seeded at elevations below 3,200 ft (975 m) on sites that receive 10 to

12 inches (250 to 300 mm) of annual precipitation.

Sodar receives use on roadways, dry drainageways,

destabilized watersheds, and mine disturbances. Sodar

seedlings are vigorous and compete well with annual

weeds; established stands restrict weed spread.

Sodar is not commonly seeded to provide livestock

forage because of its low herbage production. Plants

begin growth as early as standard crested wheatgrass

and provide good spring forage. They remain green

somewhat later in summer than standard crested

wheatgrass. The presence of green leaves into the dry

season can reduce the spread of fire. Sodar is more

drought tolerant than most populations of bluebunch

wheatgrass. Fall regrowth is generally limited. Sodar

provides diversity and ground cover in dry communities where few other species are adapted. It can be

established easily by drilling or by broadcast seeding

and harrowing.

Agropyron desertorum

Standard Crested Wheatgrass or

Desert Crested Wheatgrass _______

Description

Standard crested wheatgrass is an introduced, coolseason, generally nonrhizomatous, perennial bunchgrass that is similar to fairway crested wheatgrass in

appearance (figs. 1 and 4). Dewey (l986) described
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 321

Chapter 18 Grasses

characteristics that can sometimes be used to differentiate the two species. The spikelets of standard

crested wheatgrass are subcylindrical and oblong to

linear, diverging from the rachis at angles of 30 to 45

degrees. The glumes of standard crested wheatgrass are appressed to the lemmas, and the lemmas

support a short, straight awn that is 0.1 inch (3 mm)

or less in length. Fairway crested wheatgrass has

broad, pectinate spikes and spikelets that diverge

from the rachis at much greater angles (45 to 90

degrees), and its glumes are not appressed to the

lemmas, thus the spike has a bristly appearance. Its

lemmas are short with straight awns that may reach

0.2 inch (5 mm) in length.

Arnow (1987) combined A. cristatum, A. desertorum,

and A. fragile (= A. sibiricum) into a single species, A.

cristatum, based on their morphological similarities.

She also provided a key for separating standard and

fairway crested wheatgrasses in the field using many

of the same floral characteristics described by Dewey

(1986). This key described the culms of fairway crested

wheatgrass as often less than 1.3 ft (4 dm) tall and

typically variable in height with both basal and cauline

leaves. Culms of standard crested wheatgrass were

described as often more than 1.3 ft (4 dm) tall and

typically more or less equal in height with strictly

basal leaves.

Spikes of standard crested wheatgrass are 6 inches

(15 cm) long and 0.2 to 0.9 inch (5 to 23 mm) wide.

Spikelets are 0.2 to 0.6 inch (5 to 15 mm) long, excluding the awns, sessile, and spaced 0.04 to 0.12 inch (1 to

3 mm) apart along the rachis. Each spikelet produces

3 to 10 florets, but the upper florets are sterile. Glumes

are nearly equal, about 0.28 inch (7 mm) long, and

linear to lanceolate with the tip often tapering to a

short awn. Glumes and paleas are often rough along

the keel. The lemma is 0.16 to 0.33 inch (4 to 8.5 mm)

long and sometimes short pubescent. Leaf blades are

2 to 8 inches (5 to 20 cm) long and 0.08 to 0.2 inch (2 to

5 mm) wide. Most are flat or slightly bent along the

midrib. Plants reproduce by seeds and tillers (Arnow

1987; Hitchcock 1950).

Ecological Relationships and Distribution

Standard crested wheatgrass is native to European

Russia, the Caucasus Mountains, Western Siberia,

and Central Asia (Weintraub 1953). It has been widely

planted throughout North America in dryland and

pasture seedings. In the Western United States it is

extensively seeded in salt desert shrublands and in big

sagebrush, pinyon-juniper, and mountain big sagebrush communities (Johnson 1986). Standard crested

wheatgrass is typically less tolerant of shade, highelevation conditions, and salty soils than fairway

crested wheatgrass (Plummer and others 1958).

Standard crested wheatgrass is well adapted to the

Northern Great Plains (Westover and others 1932)

and the prairies of Western Canada (Hubbard 1949).

Sarvis (1941) reported that crested wheatgrasses,

including standard crested wheatgrass, progressed

from a promising species in 1917 to the leading hay

and pasture species in the Northern Great Plains.

Lorenz (1986) reported that it had replaced smooth

brome as the most important species planted in the

Northern Great Plains of the United States and in

Canada. It exceeds smooth brome in drought tolerance, forage production, seed production, ease of establishment, and preference by livestock as hay or

pasture.

Figure 4—Standard crested wheatgrass is a Eurasian introduction that has been widely seeded in

semiarid areas of the Western United States

(RMRS photo).
322 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Standard crested wheatgrass has been seeded widely

and has proven well adapted in almost all big sagebrush communities in the West. It is recognized for its

ability to establish and persist in disturbances on

semiarid lands. It is adapted to the high desert of

eastern Oregon and similar areas throughout the

Pacific Northwest (Jackman and others 1936).

Leckenby and Toweill (1983) reported that it performed well in juniper/big sagebrush-antelope bitterbrush communities in Oregon. It is well adapted to the

cold desert regions of Utah and Nevada. Standard

crested wheatgrass and Russian wildrye have persisted better than any other species tested in seedings

on winterfat/shadscale sites in south-central Utah.

These two introduced grasses demonstrate the ability

to persist for over 60 years without significant loss of

vigor or stand density. Standard crested wheatgrass is

also used in the warm desert regions of the Southwest,

particularly pinyon-juniper and big sagebrush communities of northern New Mexico and Arizona

(Reynolds and Springfield 1953) and the mid and

lower elevation plant communities of the Four Corners area.

Although it is widely planted, standard crested

wheatgrass it is not suited for some situations where

it has been seeded (Hull 1963b; Johnson 1986). Standard crested wheatgrass is adapted to areas receiving

10 to 20 inches (250 to 500 mm) of annual precipitation. Stands often establish and persist on sites receiving less than 10 inches (250 mm) of annual precipitation, but favorable conditions must exist at the time of

seeding and seedling emergence to assure establishment. Seedings have persisted for many years in very

dry situations, but eventually succumb during periods

of extreme drought. Standard crested wheatgrass is

better adapted to dry sites than varieties of fairway

crested wheatgrass, but it is less salt tolerant (Maas

and Hoffman 1977). Dewey (1960) observed differences in salt tolerance among collections of both species, but concluded Fairway crested wheatgrass generally produced greater yields when grown under salt

stress than did standard crested wheatgrass.

Plant Culture

Standard crested wheatgrass generally produces

good seed yields when grown under cultivation on

irrigated or nonirrigated sites. It is highly competitive

and can normally be grown without serious weed

problems. Seed yields vary among years, but some

seed is normally produced. Considerable seed is harvested from dry farms or wildland stands during years

of above average precipitation. Stable markets have

developed for this species, and seed prices are generally low.

Seed production fields can be established rather

quickly, normally within 2 years. Seed yield on dry

lands averages about 250 to 300 pounds per acre

(280 to 336 kg per ha) (Mayland 1986). For cultivated

fields in the Great Plains, Intermountain region, and

the Southwest, yields vary between 100 and 600

pounds per acre (112 to 672 kg per ha) (Reynolds and

Springfield 1953; Westover and others 1932). Wide

row spacings, 24 to 36 inches (61 to 91 cm), increase

yields (Windle and others 1966). Knowles and Kilcher

(1983) reported maximum yields resulted when stands

were planted at 12 to 18 inches (30 to 46 cm) row

spacings and fertilized with about 40 pounds of nitrogen per acre (45 kg per ha). Narrow spacings of less

than 18 inches (46 cm) are not advisable. McGinnies

(1971) found that during periods of below average

rainfall, plantings with row spacings of 18 to 24 inches

(46 to 60 cm) failed to produce harvestable crops.

Seed production continues for many years on fields

that are kept free of weeds. Seeds shatter when mature, and they must be harvested before being dislodged by wind. Some debris can be removed during

the combining process. Once seeds are harvested, they

are easily threshed and cleaned using fanning mills.

Seed quality standards are normally 95 percent for

purity and 90 to 95 percent for germination. Seeds can

be stored for 2 to 3 years or longer without appreciable

loss in germination or seedling vigor.

Seeds germinate rapidly when placed in cool, moist

environments. Love and Hanson (1932) found that

seeds of crested wheatgrass placed in incubation at 25

∞C (77 ∞F) produce a coleorhiza and several root hairs

within 24 to 48 hours. Both the coleoptile and primary

root were well developed within 72 to 96 hours. This

explains, in part, why crested wheatgrass plants establish so well under arid conditions. Seedlings are

capable of emerging and establishing within a short

period before soil surfaces dry in spring. Plummer

(1943) and Plummer and others (1958) concluded that

the widespread success of crested wheatgrass establishment is due to the greater root length attained by

developing seedlings at low temperatures when compared to many natives.

Seedlings can be established from both fall and

spring plantings, but fall seedings are recommended.

Seeds are drill seeded or broadcast and covered using

a chain or harrow. Standard crested wheatgrass is

often seeded on rough soil surfaces without any attempt to cover the seed. Although this practice is not

advised, satisfactory stands are sometimes achieved.

Planting at depths of about 0.5 to 0.75 inch (1.3 to 1.9

cm) enhances seedling emergence. Deep-furrow drilling has been recognized as a satisfactory means of

planting rangeland sites (McGinnies 1959), especially

where weeds are a factor. Evans and others (1970)

concluded that deep furrows provide a favorable seedbed by reducing temperature extremes and moisture

loss.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 323

Chapter 18 Grasses

Hull and Klomp (1967) reported that drilling produces 10 times more seedlings than broadcasting and

that drilled stands reach maturity much sooner. This

is important if a single species is desired, but it

produces more competition with other species, including recovering natives. Seeding closely spaced furrows

at high rates is advisable for weed control. Broadcast

seeding followed by chaining or harrowing to cover

the soil can produce successful plantings. Broadcast plantings are not as prone to frost damage as

drill seedings that promote more uniform seedling

emergence.

Mixed plantings of standard crested wheatgrass

and less competitive, slower developing species can be

established by interseeding the species in alternate

rows if adequate row spacing is provided (McKell

1986). Hubbard and others (1962), for example, found

that antelope bitterbrush must be seeded at least 2 ft

(0.6 m) from crested wheatgrass to assure initial

seedling survival. Walker and others (1995) found

that mixed seedings of standard crested wheatgrass

and other introduced and native species on pinyonjuniper sites declined in diversity over a 30-year period. Crested wheatgrass increased at the expense of

native plants during average or above average precipitation years. Monsen and Shaw (1983c) reported that

crested wheatgrass, seeded as an understory with

antelope bitterbrush, restricted shrub seedling recruitment over a 40-year period. Standard crested

wheatgrass reduces natural reestablishment of big

sagebrush and associated species in areas where annual precipitation is near the lower limit for the native

species.

Recommended seeding rates vary considerably

among regions. Rates of 8 to 12 pounds per acre (9 to

13.5 kg per ha) were commonly employed in early

plantings. Drill seeding rows 6 to 12 inches (15 to 30

cm) apart at this rate provides competitive cover that

will prevent weed invasion. For other plantings we

recommend rates of 1 to 2 pounds per acre (1.1 to 2.2

kg per ha) to permit initial establishment of other

seeded species and recovery of existing native herbs

and shrubs. Attempting to manage for natural recovery by seeding standard crested wheatgrass with native species is, however, a questionable practice, as full

stands of the wheatgrass may develop over time.

Standard crested wheatgrass does not spread into

adjacent areas, but seedling recruitment does occur

within the planting. Few seedlings of other species

establish in dense wheatgrass stands because the

existing crested wheatgrass plants and emerging seedlings are quite competitive (Marlette and Anderson

1986). As large disturbances are seeded to standard

crested wheatgrass, potential native seed banks are

depleted over time, and recruitment of new plants is

limited to the edges of the seedings. Areas seeded to

this perennial wheatgrass are not likely to revert to a

native composition.

Uses and Management

Standard crested wheatgrass has been used for a

number of purposes. Westover and Rogler (1947) concluded that this grass has considerable value as a

supplement to native range. It is perhaps the most

widely seeded grass in the Intermountain region,

because it is adapted to semiarid sites where native

communities have been seriously altered, and where

few other species establish. It can be grazed 2 to 3

weeks earlier than most native ranges, providing

highly succulent forage for lactating livestock in early

spring (Harris and others 1968). Considerable regrowth becomes available if fall precipitation occurs.

The ease with which this species establishes and its

early spring growth favor its use to provide soil protection and control weeds such as halogeton and cheatgrass

(Sharp 1986; Stoddart and others 1975). It is particularly important for seeding semiarid and arid sites

(Lorenz 1986). Standard crested wheatgrass gained

early recognition for its use in restoring degraded

ranges during the drought of the 1930s (Lorenz 1986;

Sarvis 1941). It remains an important species for

seeding abandoned and degraded rangelands and is

considered one of the most important species for weed

and fire control in big sagebrush and salt desert

communities.

Standard crested wheatgrass has also been used to

convert big sagebrush and native bunchgrass ranges

to crested wheatgrass monocultures. Extensive sagebrush acreages have been cleared and seeded with

standard crested wheatgrass to provide a more palatable species for livestock forage (Frischknecht 1963).

Some managers have promoted such conversions of

native bunchgrass ranges because standard crested

wheatgrass is more tolerant of grazing. Standard

crested wheatgrass sometimes provides spring forage

2 or 3 weeks earlier than some native grass and brome

grass pastures (Williams and Post 1941), but not as

early as Sandberg bluegrass or muttongrass. Compared to bluebunch wheatgrass, standard crested

wheatgrass can more rapidly activate buds, initiate

new tillers, and better allocate limited carbon reserves

to support regrowth (Caldwell and Richards 1986b).

Forage quality of standard crested wheatgrass is

high during May and June, but declines rapidly as

flowering begins (Malechek 1986). For this reason it is

often seeded with perennial broadleaf herbs, including alfalfa, to enhance and sustain high-quality herbage later in the growing season. Mixed grass-legume

seedings also substantially increase yields (Johnson

and Nichols 1969; Schuman and others1982) and are

recommended for sites exceeding 14 inches (350 mm)

of annual precipitation.
324 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Mixed seedings must be carefully managed to ensure that all species become established and persist.

Competition from the grass coupled with selective

grazing of legumes can restrict natural reseeding of

the legumes. Rosenstock and Stevens (1989) found no

recruitment of alfalfa in mixed crested wheatgrass

seedings on pinyon-juniper sites in central Utah and

attributed this to competition from established grasses.

Mixed seedings of standard crested wheatgrass and

forage kochia have been planted to provide midsummer and winter grazing not provided by the grass

alone (Monsen and others 1990). Cattle graze these

seedings during periods when the grass is dry and not

grazed.

Standard crested wheatgrass has been widely used

to seed drastic disturbances including mined lands,

roadways, and degraded watersheds. It establishes

and persists well on such sites and provides weed

control and soil protection (DePuit 1986). It can be

damaged by wildfires, but it normally recovers unless

the burn occurs under extreme conditions.

The decision to use standard crested wheatgrass in

wildlife improvement projects must be carefully considered. Because it is a competitive species that frequently gains dominance (Johnson and Nichols 1969;

Schuman and others 1982; Stitt 1958), important

habitat can eventually be lost. Increases in crested

wheatgrass can also occur over periods of 10 to 40

years or more without livestock grazing (Monsen and

Anderson 1993). Introduced grasses, including both

standard and fairway crested wheatgrass, have been

used to seed degraded big sagebrush, salt desert shrub,

and pinyon-juniper communities. In some situations,

these species are used to replace natives. Although

both grasses furnish early spring herbage, they are not

the species most preferred by mule deer (Austin and

others 1994) or pronghorn antelope (Urness 1986;

Yoakum 1980). Memmott (1995) reported that the

nutritive qualities of various native shrubs exceeded

those of crested wheatgrass in nearly all months.

Seeding standard crested wheatgrass and other highly

competitive introduced grasses has reduced the productivity of sage-grouse habitat. These grasses have

greatly decreased the density and distribution of forbs

and sagebrush in seeded communities and inhibited

their natural establishment and successful reproduction. Yoakum (1983) concluded that broadleaf herbs

and woody species are more important to pronghorn

antelope than are seeded grasses.

Elk herds have benefited immensely from large

revegetation projects in Utah. Animals have been

attracted to these areas for winter grazing, and they

use grass-herb seedings at all seasons. Sites seeded to

standard crested wheatgrass, alfalfa, small burnet,

and other grasses have successfully attracted animals

in early spring, thus reducing damage to agricultural

fields. To be fully successful, however, revegetation

efforts must allow for the recovery of native species,

and standard crested wheatgrass prevents this from

occurring. Restoration of diverse native plant communities is essential to maintain ecologically sound and

properly functioning systems. Converting extensive

areas to a single introduced species is not advised.

Standard crested wheatgrass is a persistent perennial that can withstand annual grazing. Reynolds and

Springfield (1953) recommend that use not exceed 55

percent in any year. Many stands seeded between

1932 and 1934 in southeastern Idaho have been grazed

each year, and persist after 60 years (Hull and Klomp

1966). Standard crested wheatgrass has been used as

a hay crop in dryland pastures (Stitt 1958), but more

productive and better quality herbage is produced

from mixed plantings. This species can be used as

winter forage if supplements are provided.

Varieties and Ecotypes

Nordan, released in 1953, is the most widely used

standard crested wheatgrass variety in the United

States. It was selected from material received from the

U.S.S.R. and planted at Dickerson, ND, by the Agricultural Research Service, Northern Great Plains

Research Center. Nordan was selected for its large

seeds, reduced awns, and good seedling vigor (Rogler

1954; Wolfe and Morrison 1957). It is a uniform, erect,

and productive cultivar with excellent establishment

attributes. Plants are palatable when green, but palatability is reduced as the dry stems attain maturity.

Nordan generally produces high amounts of seed each

year, even under arid conditions. This cultivar is

widely adapted to sagebrush steppe communities, but

it is also seeded in salt desert shrublands. It is one of

the main cultivars planted in large restoration projects

on semiarid lands.

‘Summit’ was developed and released in 1953 by

Agriculture Canada at Saskatoon, Saskatchewan

(Hanson 1959). This cultivar is similar to standard

crested wheatgrass varieties grown in the United

States. It was developed from materials obtained from

the Western Siberian Experiment Station. Fairwaytype plants were removed during the first year of

increase (Alderson and Sharp 1994). Summit provides

abundant forage, but problems associated with seed

processing have limited its popularity and use (Asay

and Knowles 1985b).

The Agricultural Research Service in Logan, UT, is

currently investigating other selections of crested

wheatgrass. Attention is being given to the development of a turf-type, more decumbent, and rhizomatous cultivar. In addition, studies are being conducted

to combine leafiness, retention of forage quality, larger

seed size, and vegetative vigor with the establishment

traits and early growth of Hycrest.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 325

Chapter 18 Grasses

Agropyron elongatum

Tall Wheatgrass _________________

Synonyms

Agropyron elongatum ssp. ruthenicum

Elytrigia ponticum

Elymus elongatus ssp. ponticus

Lophopyrum ponticum

Description

Tall wheatgrass is an introduced, perennial, coolseason, bunchgrass (fig. 5). It is a tall, very coarse,

productive, and late-maturing species. Culms are

densely tufted, erect, leafy, and 16 to 42 inches (4 to 10

dm) tall. Leaves are flat or involute, 0.08 to 0.3 inch (2

to 8 mm) wide, and glabrous or with the upper surface

scabrous or pubescent with short stiff hairs. Spikes

are generally erect and 3 to 16 inches (8 to 40 cm) long

with a continuous rachis. Internodes are mostly 0.3 to

0.8 inch (7 to 20 mm) long. Spikelets are solitary at

each node, compressed at maturity, and 0.5 to 1.4

inches (13 to 35 mm) long. Each produces five to eight

flowers. Glumes are typically 0.2 to 0.4 inch (6 to 11

mm) long, thick, hardened at maturity, conspicuously

five to seven nerved, and rounded on the back. Lemmas are 0.4 to 0.5 inch (9 to 13 mm) long, thick,

hardened, oblong to lanceolate, and truncate to acute

at the apex (Arnow 1987).

Ecological Relationships and Distribution

Tall wheatgrass is a native of southern Europe and

Asia Minor, where it occurs on saline meadows and

seashores (Beetle 1955). Selections from Russia and

Turkey were introduced in the United States in 1909,

but discarded as being too aggressive. The species

received favorable ratings in Athens, GA, in 1920, but

it was ignored for another decade (Weintraub 1953).

By 1930 newer introductions and studies prompted

increased use of tall wheatgrass in the Western United

States. Plants discovered growing in salt marshes

were subsequently found well adapted to saline or

alkaline soils of the Intermountain region. Tall wheatgrass is adapted to subirrigated and irrigated saline

soils and poorly drained alkali soils at elevations to

6,560 ft (2,000 m) (Hafenrichter and others 1968).

Seeding tall wheatgrass has increased forage production and quality on greasewood and saltgrass sites

receiving 8 to 12 inches (200 to 300 mm) of annual

precipitation. Tall wheatgrass is also adapted to sagebrush and mountain brush rangelands, but it has not

performed well on acid soils of the Idaho Batholith. It

survives extremely cold winter temperatures, even

when soils are bare of snow. It is widely used in the

Intermountain region, the Great Plains, and the Northwestern United States, and parts of South America.

Plant Culture

Seed of tall wheatgrass is produced in cultivated

fields at mid- and low-elevation locations characterized by long growing seasons (Hafenrichter and others

1968). Seed fields should have deep, well-drained

soils. Recommended row spacing is 36 to 42 inches (90

to 106 cm). Fields must be cultivated to control weeds

and maintain proper spacing. Plants are leafy and

quite tall, sometimes exceeding 6.5 ft (2.0 m) in height,

but they must be irrigated and fertilized with nitrogen

at 60 to 80 pounds per acre (67 to 89 kg per ha) to

maximize yields. Seeds ripen in late summer, particularly when grown under irrigation. Foliage and dry

seeds can be harvested with a combine when dry. If

plants are too green to harvest directly, stands can

first be cut, windrowed, and allowed to dry before

combining. Yields vary from 300 to 500 pounds per

acre (333 to 555 kg per ha) (Hafenrichter and others

1968). If properly managed, seed fields will remain

productive for many years. Field-harvested materials

are easily cleaned, as the seeds are large, smooth, and

easily detached from the spikelet. Seed lots with over

90 percent purity and germination are commonly sold.

Late fall or early winter seeding is recommended for

establishing tall wheatgrass in upland rehabilitation

plantings. Meadows and wetlands that do not dry out

in early spring may be seeded in late spring and

summer. Sites requiring control of excess salinity or

weedy competition may also be seeded in these seasons. Surface salts can be flushed from the seedbed by

spring and early summer irrigation; mechanical weed

control treatments may be continued until midsummer. Spring and summer seeded sites usually require

Figure 5—Tall wheatgrass was introduced from

southern Europe and Central Asia and is adapted to

saline and alkaline soils (RMRS photo).
326 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

irrigation before and after planting to assure seed

germination and seedling establishment.

Seeds must be incorporated into the soil at 0.5- to

1.0-inch (1.2- to 2.5-cm) depths; deep furrow drilling is

recommended. If surface salts are a problem for small

seedlings, the deep furrows provide ridges where evaporation can occur, leaving salts on the ridge surfaces

and reducing the salt content in the bottom of the

furrows where the seeds are planted.

Tall wheatgrass has large seeds that can be planted

with most equipment. Seeds germinate rapidly.

Germinants are vigorous and develop quickly, producing seedlings that compete well with weeds. Tall

wheatgrass is often seeded alone, but it can be successfully established in mixed plantings.

Uses and Management

Tall wheatgrass is adapted to valley bottoms that

may be saline or alkaline, and to foothill communities.

It is the most robust and erect bunchgrass currently

under use for range and pasture plantings. On salty

soils it has established better and produced higher

yields than most other forage species tested. When

seeded on low-value saltgrass sites it produces as

much as 7 tons of forage per acre (15.5 metric tons per

ha) (Richards and Hawk 1945) and 600 pounds of seed

per acre (660 kg per ha). It will grow on sites with high

water tables, but it will not survive flooding for extended periods.

Dewey (1960) found that tall wheatgrass was superior to other wheatgrass species in its ability to grow

on salt affected soils. Salinity levels of 6,000 to 18,000

ppm increase yields of this species, resulting, in part,

through reduction in competition with other plants.

Forsberg (1953) found tall wheatgrass and slender

wheatgrass were more resistant to saline conditions

than other plants tested. These two species persisted

in soils with conductivity readings as high as 15.13

mmhos per cm. Roundy and others (1983) found that

tall wheatgrass and Great Basin wildrye cultivars

were well suited to degraded saline and alkaline rangelands in arid regions of the West. Both species survived soil osmotic potentials as low as –3.5 MPa but

grew very little below –1 MPa. Robertson (1955) reported that roots of tall wheatgrass penetrated to

depths of more than 12 ft (3.7 m) in saline or alkaline

soils.

Tall wheatgrass is less drought tolerant than standard crested wheatgrass, but it is considerably more

drought tolerant than smooth brome. It does well on

rangelands that receive more than 12 inches (300 mm)

of annual precipitation. It provides excellent spring

forage, as it is about 2 weeks earlier than most native

vegetation. It is the latest maturing grass adapted to

the continental climates of the West (Hafenrichter

and others 1968). Some leaves remain green most of

the summer. As plants mature, the leaves become

coarse and less palatable, but even in winter large

amounts are consumed by livestock and game animals. Heinrichs and Carson (1956) found that the

protein content of tall wheatgrass was lower and the

crude fiber content higher at all seasons than for other

grasses used in dryland revegetation. Cook and others

(1967) also reported lower levels of total protein for tall

wheatgrass compared to selected species planted in

central Utah. Total digestible nutrients and digestible

protein for tall wheatgrass, however, are higher in the

early leaf stages than for crested wheatgrass

(Hafenrichter and others 1968).

Tall wheatgrass can be seeded in irrigated and

nonirrigated pastures. On rangelands, plants may

reach heights of more than 3 ft (1 m), while irrigated

plants often attain heights of nearly 6 ft (1.8 m).

Although tall and rank, tall wheatgrass produces

abundant forage. It is often used for hay and provides

good silage when cut between the heading and softdough stages. On rangelands it responds well to fall

rains and greens up as moisture becomes available.

With irrigation, plants provide green forage throughout the summer. It does not recover as well from

clipping as other pasture plants, but it can be grazed

during all seasons. Tall wheatgrass is often seeded

alone and grazed during midsummer. Sheep as well as

cattle use this species. Young lambs and ewes forage

on tall wheatgrass from early spring to mid to late

summer. Horses and dry cattle are often wintered on

tall wheatgrass pastures.

Tall wheatgrass is generally seeded with other species on salt desert shrublands and in sagebrush and

pinyon-juniper communities. Its production may equal

or exceed that of crested wheatgrass under these

conditions. It generally develops a dense stand when

seeded on seasonally dry meadows and lowlands, but

it forms more open stands on foothill benchlands. Once

established, plants persist well, but spread from natural seeding is quite slow.

Tall wheatgrass is useful for planting disturbed wastelands where soil salinity prevents the establishment or

growth of other species. It is planted along waterways

and on roadway disturbances, abandoned croplands,

and sites where grazing has eliminated native species.

It is useful for controlling annual weeds, particularly

chenopods. Tall wheatgrass is a useful cover crop and

can be used to provide nesting cover, winter protection,

and escape cover for upland game birds. It is commonly

used to protect the soil and provide habitat for wildlife

on farmlands where it is adapted.

Varieties and Ecotypes

‘Alkar’ tall wheatgrass was released in 1951 by the

National Resources Conservation Service Plant Materials Center, in Pullman, WA. It was developed by
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 327

Chapter 18 Grasses

mass selection from spaced plantings of materials

introduced from the U.S.S.R. in 1932 (Hanson 1959).

Alkar is a tall, very late maturing bunchgrass with

large, coarse, blue-green leaves. This cultivar produces large seeds, and its seedlings are vigorous and

fast growing. Alkar is tolerant of wet, alkaline soils

and can be seeded on semiarid rangelands at elevations between 290 and 5,500 ft (90 to1,830 m). Because

of its palatability, it receives use as a pasture plant on

subirrigated and irrigated saline and alkaline soils

(Alderson and Sharp 1994).

‘Jose’ was released in 1965 by the National Resources Conservation Service Plant Materials Center

at Los Lunas, NM. It was selected from material

received from Eurasia. Jose was developed as a pasture and hay crop. It is a uniform, medium-tall bunchgrass with green leaves. It is shorter and produces less

seed and herbage than other tall wheatgrass cultivars.

It is drought tolerant and very salt tolerant (Alderson

and Sharp 1994).

‘Largo’ tall wheatgrass was cooperatively released

in 1961 by the National Resources Conservation Service Plant Materials Center in Los Lunas, NM, the

Utah Agricultural Experiment Station in Logan, UT,

the Agricultural Research Service, Logan, UT, and the

New Mexico Agricultural Experiment Station, Las

Cruces, NM. It was developed as a forage species for

seedings in Colorado, Utah, Arizona, and New Mexico,

but Jose has largely replaced it.

‘Orbit’ was selected at the Agriculture Canada Research Station, Swift Current, Saskatchewan, in 1966.

Alkar was selected for its winter hardiness, seed

yields, and forage production. This cultivar is superior

to Alkar in winter hardiness, but similar in seed and

forage yields. It can withstand spring flooding for 3 to

4 weeks (Alderson and Sharp 1994).

‘Platte’ was selected at the Nebraska Agriculture

Experiment Station in Lincoln, NE, and released in

1972 by the Agricultural Research Service and the

University of Nebraska, Lincoln. It is a winter-hardy

bunchgrass that spreads from short rhizomes and

produces long, narrow leaves and rather large seeds.

It is primarily used to seed saline and alkaline soils in

the Great Plains (Alderson and Sharp 1994).

Agropyron intermedium

Intermediate Wheatgrass _________

Synonyms

Agropyron trichophorum

Elymus hispidus

Elytrigia intermedia

Thinopyrum intermedium

Thinopyrum intermedium ssp. barbulatum

Description

Intermediate wheatgrass is an introduced, coolseason, sodforming perennial (fig. 6). Stems are bluish

green, robust, erect, and 16 to 48 inches (40 to 120 cm)

tall. Leaf sheaths are typically ciliate, at least along

one margin. Blades are 4 to 16 inches (10 to 40 cm)

long, 0.2 to 0.4 inch (5 to 10 mm) wide, dark green to

bluish green, and flat or involute. Blade surfaces are

distinctly veined and glabrous to pubescent with short,

stiff hairs. The ligule is generally flat, 0.4 to 0.8 inch

(1 to 2 mm) long, membranous, and smooth to ragged

along the edges. Auricles are well developed. Spikes

are erect, 4 to 8 inches (10 to 20 cm) long, and slender.

Internodes of the rachis are 0.24 to 0.80 inch (6 to 20

mm) long near the center of the spike. Spikelets are

solitary at each node, compressed at maturity, and

overlap only slightly, if at all. They are 0.35 to 0.8 inch

(9 to 20 mm) long, 0.8 to 0.12 inch (2 to 3 mm) wide, and

three to eight flowered. Glumes are thick and hardened, oblong to lanceolate, conspicuously three to

seven nerved, and rounded to slightly keeled on the

back with a blunt or pointed tip. Spikelets are awnless or awn tipped (Arnow 1987; Hitchcock 1950;

Stubbendieck and others 1992).

Pubescent wheatgrass was previously considered a

separate species from intermediate wheatgrass (Nevski

1934), but they differ only in the presence or absence

of a pubescence on the spikes, seeds, and leaves.

Figure 6—Intermediate wheatgrass, a sodforming

introduction, is seeded to provide hay, forage, and

stabilization of disturbed sites (RMRS photo).
328 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Dewey (1977) concluded that these are not sufficient

differences for separation of species within this tribe,

particularly as the taxa do not differ in greatly in

geographical distribution, ecological adaptation, mode

of reproduction, or cytological characteristics. Considerable variability, however, does occur within each

form, and cultivars of each have been developed and

released.

Ecological Relationships and Distribution

Intermediate wheatgrass and its pubescent form,

pubescent wheatgrass, are distributed from southern

Europe through the Middle East to Western Pakistan

(Bor 1970). Both forms of the species were introduced

into the United States from Central Asia in 1907 (Asay

and Knowles 1985b). Additional introductions were

received in the 1920s, but the value of the plant was

not recognized until the 1930s (Weintraub 1953).

Intermediate wheatgrass is adapted to areas in the

Northwest, the Intermountain West, and the Plains

States. Both do well as understory species on welldrained dry sites in open stands of ponderosa pine and

antelope bitterbrush. At low and midelevations, intermediate wheatgrass is adapted to open parks and

extremely hot, well-drained sites on exposed slopes.

Intermediate types are more shade tolerant than the

pubescent types and they are better adapted to cooler

and wetter situations. They perform better on upland

benches occupied by Wyoming big sagebrush and in

mountain big sagebrush, herblands, mountain brush,

aspen, and fir communities receiving annual precipitation of 14 inches (350 mm) or more. Pubescent forms

are more tolerant of sites with long, warm summers

than are intermediate forms. They are better adapted

to blackbrush, southern desert shrub, and some Wyoming big sagebrush communities receiving less than

12 inches (300 mm) of annual precipitation. Populations of intermediate wheatgrass seeded at low elevations will eventually shift to support individuals with

pubescent characteristics. The reverse will occur if

pubescent forms are planted in upland communities.

Intermediate wheatgrass is less drought tolerant than

standard crested wheatgrass, but it is equally cold

tolerant. Pubescent forms withstand high summer

temperatures of the blackbrush and southern desert

shrub types better than standard or fairway crested

wheatgrass.

Intermediate wheatgrass is adapted to soil types

ranging from heavy-textured with low pH to coarsetextured granite materials with neutral or slightly

acidic pH. Both the intermediate and pubescent forms

are suited to droughty infertile soils, but the pubescent forms are better adapted to saline and alkaline

soils.

Plant Culture

Intermediate wheatgrass seed is produced in

nonirrigated and irrigated fields. Yields range from

250 to 500 pounds per acre (280 to 560 kg per ha)

(Hafenrichter and others 1968). Good yields can be

produced with limited fertilization and irrigation.

Plants develop a competitive sod rather quickly, and

renovation of seed production fields with mechanical

tillage is required to maintain acceptable seed production. Intermediate wheatgrass produces seedstalks

with large seedheads. Seeds ripen uniformly and remain on the plant, but can be easily separated from the

inflorescence during cleaning. Immature florets can

be removed using fanning mills and other separators.

Seeds can be cleaned to provide high purity and

germination levels. Seed viability is retained for a

number of years when seeds are stored under warehouse conditions.

Seeds should be planted at depths of 0.5 to 1.5 inches

(1.3 to 3.8 cm) using conventional drills. They germinate quickly when seedbed conditions are favorable

and produce vigorous seedlings. Small seedlings and

young plants are able to compete favorably with most

other species. Unstable soils and weedy sites are often

drill seeded with intermediate wheatgrass to provide

permanent stands and uniform, competitive cover in 3

to 5 years. Seeding at rates below 2 pounds per acre

(2.2 kg per ha) is recommended when planting mixtures for range and wildlife habitat improvement.

Rates from 8 to 12 pounds per acre (9 to 13 kg per ha)

are suggested for planting highly erodible roadways

and watershed disturbances. Intermediate wheatgrass

is one of the most successful grasses to establish from

broadcast seedings on sites inaccessible to drill seeders. Patches or clumps normally develop when sites

are broadcast seeded followed by chaining or harrowing; these can coalesce over time.

Intermediate wheatgrass is often seeded in mixtures with other herbs to provide forage and cover, but

it is highly competitive and can suppress and replace

many other species. Monsen and Anderson (1993)

inventoried a 50-year old species trial in central Idaho

and found that intermediate wheatgrass and sheep

fescue were the two species best able to persist and

displace other introduced and native grasses seeded in

adjacent plots.

Uses and Management

Prior to about 1950, intermediate wheatgrass was

not planted as frequently as the crested wheatgrasses

and smooth brome in the Great Plains and Western

regions of the United States. In the Intermountain

area, it has gained importance and is now widely

planted. It has been included in seed mixtures for big
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 329

Chapter 18 Grasses

sagebrush, pinyon-juniper, mountain brush, ponderosa pine, aspen, spruce-fir, and open park herblands.

Because it is a sodformer that forms a dense root

mass (Hafenrichter and others 1968), intermediate

wheatgrass is used to improve ground cover and to

stabilize erodible soils and watershed disturbances. It

is adapted to infertile soils associated with roadway

disturbances, mined lands, waterways, and other unstable sites. It has become one of the most important

soil conservation species currently seeded. Most collections exhibit good seedling vigor, establish quickly,

and spread rapidly, even during the first year. Plants

remain green and provide effective ground cover

throughout the growing season. Hafenrichter and others (1968) reported that intermediate wheatgrass produces better ground cover than smooth brome in spring,

fall, and winter because it does not freeze as easily. It

is a highly persistent species that retains vigor and

density and performs better than orchardgrass and

timothy when planted in mixtures on infertile soils.

Intermediate wheatgrass plants spread vegetatively,

forming a dense, competitive sod. They are capable of

increasing areas of occupation even when grown on

harsh sites. Cultivars of intermediate wheatgrass

usually produce a more aggressive and rapidly developing sod than do pubescent wheatgrass cultivars.

When grown at low elevations and in more arid situations, intermediate wheatgrass forms patches that

may be widely spaced. Although pubescent wheatgrass cultivars respond similarly, they are more likely

to form a continuous stand on such sites. When grown

at mid and upper elevations, intermediate wheatgrass

develops a very complete and solid ground cover. Few

interspaces remain, and the plants are very competitive. Under these conditions, intermediate wheatgrass spreads rapidly and can dominate and displace

pubescent wheatgrass.

Intermediate wheatgrass is commonly seeded in

irrigated and dryland pastures. It is suitable for seeding with alfalfa for hay production or rotational grazing. Plants are leafy, but not rank. They are generally

in the early stage of flowering when alfalfa is ready to

cut. At this stage, both species provide high-quality

herbage. Annual clipping does not weaken stands of

this grass, and plants are capable of recovery and

regrowth from early spring and summer clippings.

Pastures and even dryland seedings can be maintained with considerable grazing.

Intermediate wheatgrass matures 2 to 3 weeks later

in summer than crested wheatgrass, and it provides

much better herbage in mid and late summer (Cook

and others 1956). It remains green in summer and fall

in mountain brush communities and at higher elevations. Both pubescent and intermediate wheatgrass

are palatable, but investigators have reported seasonal differences (Cook and others 1956; Hafenrichter

and others 1968; Hyder and Sneva 1963). Austin and

others (1994) found that ‘Luna’ pubescent wheatgrass

received greater preference than other commonly

seeded grasses when grazed in spring and fall by deer.

Differences in palatability between cultivars of the

two grasses are due in part to differences in their

phenological development. Some intermediate wheatgrass cultivars mature later in the season and are

more selectively used as pubescent growth forms become dormant (Cook and others 1956). In general,

intermediate wheatgrass cultivars are more palatable

to livestock for a longer grazing period than are pubescent wheatgrass cultivars.

Intermediate wheatgrass has been seeded in oak

brush and mountain brush thickets. It is tolerant of

some shade, and provides considerable herbage as an

understory species. It begins growth earlier than the

shrubs, and has been seeded to suppress shrub growth

and provide more open communities. Intermediate

wheatgrass withstands burning and can be used to

seed woody sites that tend to burn frequently. Once

established, it is highly persistent and capable of

withstanding underground foraging by rodents.

Although intermediate wheatgrass provides useful

herbaceous cover in aspen, mountain brush, pinyonjuniper, and sagebrush communities, this species can

slowly replace native herbs and shrubs. Monsen and

others (1996) found that extensive seedings in aspen

and associated open parks in central Utah have replaced important broadleaf herbs, big sagebrush, antelope bitterbrush, and other associated species. Complete conversion may require as much as 20 to 30

years. This grass should not be seeded in areas where

native species should be retained. Problems created by

seeding intermediate wheatgrass, particularly in aspen and open park herblands, are quite serious. Removal of wheatgrass from such stands is difficult, and

reestablishment of diverse communities is often not

practical. This plant can restore ground cover and

compete effectively with weeds, but it should not be

planted in areas where retention or reestablishment

of native species is desired. It is particularly restrictive to the establishment and survival of shrub seedlings. It is widely grown for pasture and hay crops from

Nebraska to Manitoba, Canada, the Pacific Northwest, and to a lesser extent the Intermountain West.

Varieties and Ecotypes

Cultivars of pubescent and intermediate wheatgrass forms differ in growth and utility; individual

cultivars should be carefully selected. Varieties of both

forms will grow on the same sites and frequently

hybridize, but there are some major differences between them. The first promising cultivar of intermediate wheatgrass was developed through selection from
330 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

introduction ‘PI 98568’, received from Maikop, U.S.S.R.,

in 1932 (Hanson 1965). The South Dakota Agricultural Experiment Station later released this accession

as ‘Ree’. Additional cultivars developed from this accession include ‘Chief’, ‘Greenar’, ‘Oahe’, and ‘Slate’.

Some early cultivars including ‘Amur’ have been replaced by other, better-adapted cultivars. Only the

most used cultivars are described here.

Intermediate Wheatgrass Varieties

Chief was released by Agriculture Canada in 1961

as a selection from the cultivar Ree. It was developed

by mass selection of plants with high seed yields and

good seed quality. It is primarily used in grass and

alfalfa pastures and for hay production in the Canadian Prairie Provinces. Forage and seed yields are

slightly higher than those of Ree (Alderson and Sharp

1994). This cultivar has not been widely used in the

Intermountain region.

‘Clarke’ was released in 1980 by Agriculture Canada

as a 20-clone synthetic. It was developed through eight

cycles of recurrent selection to combine winter hardiness, drought tolerance, seedling vigor, resistance to

aphid-virus infection, seed quality, and forage and

seed production. It has been useful for hay and pasture

production in dryland or irrigated fields in the Canadian Prairie region and the Northern Great Plains. It

is extremely drought tolerant and winter hardy, and

produces excellent seed yields (Alderson and Sharp

1994). Clarke is primarily used in irrigated pastures

in the Western United States.

Greenar is a leafy and highly productive variety that

was developed at the Natural Resources Conservation

Service Plant Materials Center in Pullman, WA, from

PI 98568, a collection of the Westover-Enlow expedition to the U.S.S.R. It was released in 1945 as P-2327

and renamed Greenar in 1956 (Alderson and Sharp

1994). This release is tall, leafy, productive, medium to

late maturing, and mildly sodforming (Hafenrichter

and others 1968). Plants are broadleaved and green to

dark green. Greenar is widely seeded on rangelands,

pasturelands, and hayfields in the West. It is used in

conservation plantings because it produces considerable ground cover and develops an extensive and

persistent sod. Greenar establishes well and is adapted

to harsh, dry, exposed sites. It is not shade tolerant.

Plants are disease resistant and begin growth in early

spring. They retain green herbage late into the season.

Greenar is well adapted to big sagebrush communities

as well as higher elevation sites in aspen and mountain herblands. It has persisted well on such sites,

even when grazed heavily by livestock. It remains

green and palatable through the summer and fall

months.

The South Dakota Agricultural Experiment Station

released Oahe intermediate wheatgrass in 1961. It

was selected from a Russian introduction, PI98568,

that also provided material for the Greenar and Ree

releases. Oahe was developed from two cycles of selection for high seed production, forage production, and

rust resistance. It was developed for hay, pasture, and

conservation plantings in the Northern Great Plains

(Alderson and Sharp 1994). Oahe demonstrates some

of the same vegetative traits as Greenar, but it is more

drought tolerant. Both cultivars establish quickly and

develop a dense sod. Oahe does not provide as much

late summer and fall herbage as Greenar. Both cultivars are widely adapted for conservation and pasture

plantings in the Intermountain region.

‘Reliant’ was selected at the Agricultural Research

Service Northern Great Plains Research Laboratory

in Mandan, ND, and cooperatively released by the

Agricultural Research Service, Natural Resources

Conservation Service, and the North Dakota Agricultural Experiment Station in Fargo, ND. It was selected for its vigor and resistance to leaf-spotting

disease. Reliant is adapted to hay and pasture uses in

the Northern Great Plains of the United States and

Canada. It has not been extensively planted in the

West, but appears useful in dryland pastures seeded

with alfalfa.

‘Rush’ intermediate wheatgrass was released in

1994 by the Natural Resources Conservation Service

Plant Materials Center, in Aberdeen, ID. It was originally received as A. junceum. Rush develops an extensive sod that provides useful ground cover. It is recommended for conservation plantings to control erosion

on roadways, mine disturbances, and dry waterways

(Alderson and Sharp 1994). Rush establishes well

from direct seedings and spreads quickly. It is quite

competitive and can restrict the occurrence of other

species. It has an upright growth habit and provides

considerable spring, summer, and fall forage.

The Nebraska Agricultural Experiment Station and

Agricultural Research Service cooperatively released

Slate intermediate wheatgrass in 1969. It was developed by blending equal quantities of seed from two

unrelated strains, ‘Nebraska 50’ and a derivative of

Amur (Alderson and Sharp 1994).

Slate is an upright variety with large, broad, lightgreen or blue-green leaves that produces a strong,

spreading sod. It is primarily seeded in mixtures with

other grasses and alfalfa in irrigated or nonirrigated

pastures and hayfields in the Central Plains. Although it has not been seeded extensively in the West,

it is a useful cool-season forage or hay crop. It is not

tolerant of saline or alkaline soils.

The Idaho and Washington Agricultural Experiment Stations in Aberdeen, ID, and Pullman, WA, and
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 331

Chapter 18 Grasses

the Natural Resources Conservation Service Plant

Materials Centers in Aberdeen, ID, and Pullman, WA,

released ‘Tegmar’ intermediate wheatgrass in 1968. It

was developed from accession ‘PI 109219’ collected

near Bolu, Turkey, in 1934 (Alderson and Sharp 1994).

Plants are erect and rather small statured. They are

about one-third to one-half the height of the most

robust forms of intermediate wheatgrass. Leaves are

light green to blue green with some hairs along the

margins. Tegmar is one of the most commonly used

cultivars in the Western States for stabilizing dry

waterways, highly disturbed watersheds, and disturbed rangeland sites. Plants establish quickly and

develop a dense, vigorous, rapidly spreading, and

persistent sod that can reduce erosion and withstand

some burial. Tegmar is adapted to warm and lowelevation sites. It has been seeded in lower mountain

brush, big sagebrush, and salt desert communities.

Plants furnish useful spring forage, but other cultivars may produce more succulent herbage in mid and

late summer. Tegmar is frequently seeded alone on

disturbed sites, particularly on areas with diverse

topographical conditions and infertile soils.

Tegmar is not commonly seeded as a hay or pasture

plant on irrigated sites because it does not provide late

summer herbage. It responds poorly to clipping, and it

is not as productive as other cultivars. It grows well in

open stands with big sagebrush and other shrubs, but

it is not shade tolerant. Once stands are established, it

persists well, even when heavily grazed.

Pubescent Wheatgrass Varieties

‘Greenleaf’ was developed at the Agriculture Canada

Research Station in Lethbridge, Alberta, by screening

open-pollinated progenies for seedling vigor and tendency to creep. Greenleaf is a winter-hardy forage

cultivar intended for pasture and hay production. It

produces bright-green pubescent foliage, and it is

fairly tolerant of saline soils and droughty sites.

Greenleaf is used primarily for short-duration pasture

forage in the Canadian Prairie region and the Northern Great Plains. It is similar to ‘Mandan 759,’ but it

produces more vigorous seedlings. Greenleaf is superior to ‘Topar’ in forage production, seedling vigor, and

winter hardiness (Alderson and Sharp 1994).

The New Mexico Agricultural Experiment Station

and Natural Resources Conservation Service Plant

Materials Center at Los Lunas, NM, released Luna

pubescent wheatgrass in 1963. It was developed from

materials collected by the Westover-Enlow expedition

to the former U.S.S.R. and Turkey in 1934. Luna’s

primary use has been for irrigated pastures and revegetation of range and watershed disturbances (Alderson

and Sharp 1994). It has become one of the most

commonly seeded cultivars in the West.

Luna is a tall, highly productive cultivar that is

widely seeded to furnish spring, summer, and fall

herbage. Basal leaves are moderately pubescent, but

the seedheads are usually glabrous. Its ability to

remain green throughout the summer contributes to

its popularity. It is adapted to open parks, aspen,

spruce-fir, mountain brush, and juniper-pinyon woodlands. It is not shade tolerant, but it grows well on

open, exposed slopes, particularly in areas with warm

summer temperatures. Because of its drought tolerance, Luna is adapted to big sagebrush and southern

desert shrublands. It requires 10 to 12 inches (250 to

300 mm) of annual precipitation, but it has been found

to persist in areas where rainfall is considerably lower.

Luna seedlings establish quickly and are quite vigorous. Plantings reach maturity in 1 to 2 years, and

provide some control of spread by annual weeds. Luna

is commonly seeded as a dual-purpose species to control erosion and provide forage. Luna is planted on

burned sites, roadways, and unstable watersheds to

provide initial ground cover and supply forage or

habitat. It provides a productive cover during periods

of drought and heavy grazing. Luna is competitive and

can exclude native species.

Topar pubescent wheatgrass was developed from ‘PI

ID7330’ and cooperatively released in 1953 by the

Washington, Idaho, Oregon, and California Agricultural Experiment Stations, and by the Natural Resources Conservation Service Plant Materials Centers

at Aberdeen, ID; Pleasanton, CA; and Pullman, WA.

Topar is typical of all pubescent and most intermediate cultivars in being a late-maturing, vigorous,

sodforming selection, adapted to droughty sites

(Alderson and Sharp 1994). It is seeded in aspen,

mountain brush, pinyon-juniper, big sagebrush, and

salt desert shrublands. Seedlings establish quickly

and are very competitive. These characteristics contribute to use of this cultivar on disturbed, weedy,

infertile sites. Plantings can be used to control weeds

and furnish late-season herbage. Topar is also planted

to control erosion and stabilize watersheds, roadways,

and waterways. It has been seeded in mixtures to revegetate rangeland disturbances. Topar requires between

10 and 12 inches (250 to 300 mm) of annual rainfall. It is

sufficiently drought tolerant for seeding spring and fall

foothill ranges in the Intermountain area.

Topar is commonly seeded with Tegmar intermediate wheatgrass on well-drained, droughty exposures,

where native vegetation has been lost as a result of

grazing, wildfires, or other disturbances. Both grasses

are capable of establishing on soils that dry quickly

and may be subjected to prolonged periods of summer

drought and high temperatures. Both have been used

to colonize and stabilize unstable soils. Young seedlings are able to establish on erosive surfaces, where

they stabilize the soil and suppress invasive species.
332 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Agropyron sibiricum

Siberian Crested Wheatgrass _____

Synonyms

Agropyron fragile

Description

Siberian wheatgrass is an introduced perennial

bunchgrass that is similar in appearance to both

standard and fairway crested wheatgrass, but it has a

much narrower spike (fig. 1). It is usually smaller in

stature and develops narrower bunches with less foliage. Its leaves are finer and more decumbent. The

spike is narrow, cylindrical, and usually about 2.3 to

4.7 inches (6 to 12 cm) long. The spikelets are somewhat spreading and develop directly from the rachis.

Glumes and lemmas are awnless or with awns 0.04 to

0.08 inch (1 to 2 mm) long (Hitchcock 1950).

Siberian wheatgrass is part of the crested wheatgrass complex that consists of a series of diploids,

tetraploids, and hexaploids (Dewey 1974). Diploids

are represented primarily by the Fairway cultivar.

The most commonly used tetraploids are standard

crested wheatgrass and Siberian crested wheatgrass

cultivars. Selection and hybridization have been accomplished within ploidy levels, but are primarily

confined to the diploid and tetraploid populations, as

hexaploid populations are very rare (Asay 1986).

Taxonomic changes have confused the identification

of Siberian crested wheatgrass (Dewey 1986). Jones

(1960) placed all crested wheatgrasses with narrow or

cylindrical spikes in A. sibiricum. Dewey (1986) recommended acceptance of Tsvelev’s (1976) treatment

of the wheatgrasses, which identifies three species of

crested wheatgrass—A cristatum, A. desertorum, and

A. fragile. Tsvelev’s decision to combine A. sibericium

with A. fragile resulted in the acceptance of A. fragile,

as it is the older of the two names. Some authors

(Arnow 1987) have recently grouped all crested wheatgrasses into one species, A. cristatum.

Ecological Relationships and Distribution

Siberian crested wheatgrass occurs in eastern Russia, the Caucasus, Western Siberia, and Central Asia

(Weintraub 1953). In Central Asia it is distributed

from the Caspian Sea to Lake Balkhash (Dewey 1986).

In the United States, the first introduction was received in about 1906, but the plant was not successfully grown until the 1920s (Weintraub 1953). Dewey

(1986) reported that about 50 accessions entered the

United States between 1906 and 1940. This exceeds

the number of A. desertorum introductions during this

period, but published references to it are infrequent.

Most accessions were apparently grouped within the

broad crested wheatgrass category.

Siberian crested wheatgrass is more tolerant of

drought and alkali soils than is standard crested

wheatgrass. Consequently, it performs better at lower

elevations in big sagebrush and salt desert shrublands.

In the Intermountain area it is adapted to upper

benchlands supporting mountain brush and pinyonjuniper communities and to the extensive Wyoming

big sagebrush and shadscale communities

Siberian crested wheatgrass is ecologically better

adapted to arid shrublands of the Intermountain area

than standard or fairway crested wheatgrass. It grows

in soils ranging from well-drained and sandy to heavytextured clay loams. Young plants are able to establish

on soils that dry quickly in spring and remain dry

throughout the summer. This plant demonstrates

tolerance to consecutive years of drought when planted

in the lower regions of the big sagebrush zone. It is

capable of surviving the hot, dry summers of warm

desert communities. It is not shade tolerant and generally grows better in regions receiving less than 12 to

14 inches (300 to 350 mm) of annual precipitation.

Plant Culture

Annual seed production of Siberian crested wheatgrass is lower than that of fairway or standard crested

wheatgrass. However, good seed crops can be produced on both irrigated and nonirrigated fields. Yields

averaging 200 pounds per acre (225 kg per ha) for

dryland plantings and 400 pounds per acre (450 kg per

ha) for irrigated sites are common. Seed can be harvested with combines and cleaned with air-screen

separators. Cleaned seed lots normally are of high

viability and purity. Seeds are slightly smaller than

those of other wheatgrasses.

Cleaned seed lots are usually free of large debris,

and the seeds can be metered through most seeding

equipment. Seeds should be planted between 0.5 to

0.75 inch (1.3 to 1.9 cm) deep to support germination

and establishment. Young and Evans (1986a) found

that germination characteristics of Siberian wheatgrass were similar to those of fairway and standard

crested wheatgrass. Hyder and Sneva (1963) reported

that morphological and phenological development of

Siberian and crested wheatgrass were very similar.

However, when grown under rangeland conditions,

seed maturation of Siberian crested wheatgrass is

often 7 to 10 days later than for standard crested

wheatgrass.

Seedlings of Siberian crested wheatgrass cultivars

are less vigorous than those of most fairway or standard crested wheatgrass cultivars, but established

plants are persistent. Seedling establishment strategies differ among the crested wheatgrasses. Siberian
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 333

Chapter 18 Grasses

crested wheatgrass seedlings do not grow as quickly or

become as large during the first growing season, as the

plants adjust to drought by becoming dormant earlier

in the season. Young plants may remain small for 1 to

2 years if growing conditions are unfavorable. Plants,

however, generally develop satisfactory stands, even

during periods of drought and in areas where few other

revegetation species are adapted. Plummer and others (1968) ranked seed germination, initial establishment, growth rate, and persistence of this species as

excellent.

Siberian wheatgrass can be established with limited

seedbed preparation. Seeds must be covered to assure

acceptable establishment; stands can be planted by

drilling or broadcast seeding followed by chaining or

harrowing. Seeding directly into weedy sites should be

avoided, but established plants do compete favorably

with annual weeds. Seedlings are quite hardy, and few

succumb to frost.

Uses and Management

Siberian crested wheatgrass is less widely planted

than other crested wheatgrasses, primarily due to its

lower productivity. Most rangeland and pasture

plantings are conducted to supply forage for grazing

animals, and other, more productive, species are usually planted. Siberian wheatgrass is more drought

resistant than most introduced grasses, and it is

better able to establish and provide a ground cover in

drier habitats. It provides useful, more consistent

spring and fall herbage on drier sites than other

crested wheatgrasses. It is also a formidable competitor with annual weeds.

Siberian wheatgrass can be seeded at low rates with

bottlebrush squirreltail and Sandberg bluegrass to

stabilize Wyoming big sagebrush and salt desert communities. Seedlings develop slowly, allowing establishment of big sagebrush and other shrub seedlings.

Siberian wheatgrass is adapted to soil disturbances; it

is seeded on roadway and mine disturbances in arid

regions. It is often seeded along dry streambeds where

soil stabilization is required.

Leaves of Siberian crested wheatgrass are usually

finer and remain green later into the summer than

those of other crested wheatgrasses. Siberian crested

wheatgrass is seeded with Russian wildrye to provide

later spring and summer pastures. It is also seeded on

disturbances in pinyon-juniper and big sagebrush

communities to restore forage for big game and domestic livestock. Although it is somewhat less competitive

than either standard or fairway crested wheatgrass, it

is not compatible with native herbs and shrubs, and

should not be seeded in areas where recovery of native

communities is desired.

Varieties and Ecotypes

The Idaho Agricultural Experiment Station, Moscow, and the Natural Resources Conservation Service

Plant Materials Centers at Aberdeen, ID, and Pullman, WA, cooperatively released the cultivar ‘P-27’ in

1953 (fig. 7). It was developed from material designated ‘PI 108434’, obtained in 1934 from Kazakhstan,

U.S.S.R., and from the Institute of Plant Industry,

Leningrad, U.S.S.R., by the Westover-Enlow expedition (Hanson 1959). It was evaluated in field and

nursery plantings beginning in 1935. Individual clones

were selected in 1949 and served as a basis for further

increase (Alderson and Sharp 1994). P-27 is primarily

used in range seedings. Plants are leafy and fine

stemmed. They are characterized by good seedling

vigor, high seed yields, and drought tolerance. This

cultivar has been the primary accession available for

large seedings.

Figure 7—‘P-27’, a leafy and finestemmed cultivar of Siberian wheatgrass,

was released in 1953 and has been widely

used (photo courtesy of Loren St. John,

USDA Natural Resources Conservation

Service, Aberdeen Plant Materials Center, Aberdeen, ID).
334 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

‘Vavilov’ Siberian wheatgrass was released in 1994

by the Agricultural Research Service, the Utah Agricultural Experiment Station, and the Natural Resources Conservation Service in Aberdeen, ID. It was

named to acknowledge the contribution of the N. I.

Vavilov Research Institute of Plant Industry, St. Petersburg, Russia, to the range grass plant breeding

program of the Agricultural Research Service. Vavilov

was derived from accessions obtained from the Vavilov

Research Institute; the Stavropol Botanical Garden,

Stavropoal, Russia; the Eskisehir Plant Breeding Station, Eskisehir, Turkey; and selections from the cultivar P-27 (Asay and others 1995b). The parental accessions were selected for their retention of green color

and vegetative vigor in late summer. Breeding populations were also screened for seedling vigor, seed

production, plant type, and response to drought, disease, and insects. Vavilov spikes intergrade from the

long narrow forms typical of Siberian crested wheatgrass to shorter, wider spikes more typical of standard

crested wheatgrass (Asay 1995a,b). A tetraploid,

Vavilov is fully interfertile with standard crested

wheatgrass and the Hycrest cultivar.

Vavilov is a particularly important cultivar because

of its seedling vigor. It is comparable to Hycrest and

consistently better than P-27 in terms of seedling

emergence and vigor. Vavilov is also more productive

than P-27 (Asay and others 1995b). It is recommended

for seeding semiarid sites at elevations below 6,800 ft

(2,100 m) that receive 8 to 17 inches (200 to 450 mm)

of annual precipitation.

Agropyron smithii

Western Wheatgrass or

Bluestem Wheatgrass____________

Synonyms

Elytrigia smithii

Elymus smithii

Pascopyrum smithii

Description

Western wheatgrass is a native, rhizomatous, longlived, cool-season perennial that produces an open and

highly uniform sod (fig. 8). Stems are erect and 12 to 24

inches (30 to 60 cm) tall. Leaves are typically blue

green, but the entire plant may have a grayish-blue

cast that varies among populations. Leaf sheaths are

normally glaucous and rough. Leaf blades are about 4

to 10 inches (10 to 25 cm) long and mostly 0.04 to 0.08

inch (2 to 4 mm) wide. They are usually firm, stiff, and

taper to a sharp point. The upper surface of the leaf is

rough and distinctly ridged or grooved along the nerves

while the underside is usually smooth. Green leaves

are flat to involute; when dry they remain stiff and

erect, but the edges curl toward the midrib. Auricles

are often well developed. Ligules are about 0.04 inch (1

mm) long. Spikes are stiff, erect, and about 2.8 to 6

inches (7 to 15 cm) long with one or two spikelets per

node. Spikelets are to 0.4 to 0.8 inch (1 to 2 cm) long and

overlap each other by about one-half their length.

Glumes are firm, unequal, linear-lanceolate to lanceolate, and gradually tapering from the base to a

short awn. The first glume is 0.25 to 0.45 inch (6 to 12

mm) long and the second 0.3 to 0.6 inch (7 to 15 mm)

long. Lemmas are 0.04 inch (1 mm) long and firm with

obscure nerves and a pointed tip or short awn (Arnow

1987; Asay 1995a,b; Hitchcock 1950; Sarvis 1941;

Stubbendieck and others 1992).

Ecological Relationships and Distribution

Western wheatgrass is morphologically similar to

thickspike wheatgrass, but it differs in having asymmetrical glumes (Cronquist and others 1977). Western wheatgrass has recently been placed in a separate

genus and renamed Pascopyrum smithii (Rydb.) Love

(Dewey 1984; Love 1980). P. smithiiis the sole member

of this genus. Dewey (1975) considers its putative

parents to be thickspike wheatgrass and beardless

Figure 8—A comparison of growth forms of western wheatgrass demonstrates some of the variation in leafiness in this species (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 335

Chapter 18 Grasses

wheatgrass. Its adaptability to heavy saline and alkaline soils and its forage qualities reflect the input of

these parental species (Dewey 1984).

Western wheatgrass occurs across the western twothirds of Canada and the United States, from Alaska

and British Columbia to Quebec, and southward

through the Western and Central United States (Arnow

1987; Asay 1996) east of the Cascade Mountains to

New Mexico (Hanson 1972), the Central and Northern

Great Plains, and the Texas Panhandle (Hanson 1972;

Hitchcock and others 1969). It is the only octoploid

wheatgrass native to North America (Asay 1995a,b).

Ecotypes of western wheatgrass varying in stature,

color, and sodding traits occur in a variety of plant

communities. On the Central Great Plains, it is abundant in lowlands, and may occur as nearly pure stands

aligning watercourses (Sarvis 1941). Throughout this

region it grows with blue grama, sideoats grama, alkali

sacaton, buffalo grass, and needlegrasses. In the Western United States it is often the dominant grass in salt

desert, big sagebrush, mountain brush, and pinyonjuniper communities. In Utah it rarely occurs on slopes

above 6,900 ft (2,130 m) (Arnow 1987), but it grows at

higher elevations in Wyoming and Colorado (Dittberner

and Olsen 1983). Western wheatgrass often grows in

association with bluebunch wheatgrass and needleand-thread on upland benches and with bottlebrush

squirreltail and Sandberg bluegrass in Wyoming big

sagebrush and salt desert communities.

Beetle (1955) ranked this species as the most alkali

tolerant of all North American wheatgrasses. It is

commonly found on heavy clay soils with moderate

alkalinity in salt desert shrublands. It is common in

cold desert uplands of northern Utah, western Wyoming, and eastern Idaho where salty outcrops occur

with Wyoming big sagebrush, winterfat, and various

saltbush species. It is also common in dry valley

bottoms, swales, and dry drainages where periodic

flooding occurs (Rogler 1973). Although western wheatgrass exists on fine-textured soils, it grows on welldrained bottomlands, open plains, and benchlands of

lower mountain and foothill ranges in the Intermountain region (Stewart and others 1939). It occurs in

areas receiving 10 to 20 inches (250 to 500 mm) of

annual precipitation, but performs well in the 10- to

14-inch (250- to 360-mm) zone. It is sometimes an

important understory species in ponderosa pine and

pinyon-juniper, but it is not shade tolerant and is

usually restricted to openings between trees.

Plant Culture

Seed production of native western wheatgrass stands

in the Northern Great Plains is generally greater than

from stands in more arid regions. Released cultivars

have improved seed production capabilities compared

to wildland collections. Yields ranging between 150

and 300 pounds per acre (168 to 336 kg per ha) are

expected from cultivated fields with irrigation (USDA

Natural Resources Conservation Service 2002).

Yields from dryland fields are half those of irrigated

fields. Fields require regular tillage to maintain

proper spacing and prevent overcrowding by sodding. Plants flower in June, and seeds ripen in August

or September (Wambolt 1981), later than for many

other grasses. Seed stalks are erect and project above

the leaves, facilitating combine harvesting. Seeds are

large, firm, and easily separated from the spikelet

using standard cleaning equipment.

Western wheatgrass seeds, particularly those from

wildland collections, do not germinate readily or uniformly, resulting in erratic establishment (Beetle 1955;

Ferguson and Frischknecht 1985; Monsen and

McArthur 1985). The seeds often exhibit low or delayed germination at warm temperatures between 66

to 73 ∞F (19 to 23 ∞C) (Brown and Hallman 1984;

Plummer 1976), but alternating temperatures of 59 to

80 ∞F (15 to 30 ∞C) tend to promote germination.

Erratic germination decreases survival in rapidly

drying seedbeds. Although slow- or late-developing

grass seedlings may succumb to competition with

weeds, western wheatgrass seedlings are extremely

drought tolerant. When seeded in mixtures with more

aggressive species, poor and spotty stands may develop. Increasing the seeding rate usually does not

improve plant density. However, when seeded in weedfree, but harsh environments typical of mine disturbances, this grass establishes satisfactorily. Seed germination and establishment characteristics of recently

developed cultivars are much improved over previously used wildland collections.

Established plants are vigorous and drought resistant. Plants spread vegetatively from rhizomes, forming open, irregular patches, even when seeded alone.

Western wheatgrass can be seeded with slower developing native species, including shrubs.

Uses and Management

Western wheatgrass is one of the most important

native forage grasses of the Northern Great Plains

(Sarvis 1941) and the Intermountain West (Stewart

and others 1939; Vallentine 1961). It has not been

planted as extensively on range and wildland sites as

crested wheatgrass and intermediate wheatgrass because its seedling establishment is less reliable and its

herbage yields are lower. Western wheatgrass begins

growth relatively early in spring, and plants mature

late in the season. Standing plants cure well, remaining palatable and nutritious. Energy values are rated

good (Dittberner and Olson 1983). Livestock and game

graze it during most seasons. Because it provides good

winter grazing, it is often managed to provide late fall

and winter pastures (Sarvis 1941).
336 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Western wheatgrass has been reduced or eliminated

from many Western rangelands by continuous and

prolonged grazing. Close and repeated clipping can

reduce yields and weaken stands (Wasser 1982). However, plants have demonstrated the ability to recover

if protected from grazing. Plants spread quickly from

rhizomes and are able to repopulate abandoned lands

(Rogler 1973; Weaver and Albertson 1956). Considerable recovery has been noted in Wyoming big sagebrush and pinyon-juniper communities protected from

grazing for 1 to 20 years. Plants have increased during

periods of drought (White and others 1978) and replaced annual weeds. Sarvis (1941) reported that

western wheatgrass recovers so quickly in

noncultivated fields that it has been called “go-back”

grass. Rate of recovery is based on plant vigor, density,

and the presence of other species.

Western wheatgrass is particularly important for

restoring semiarid communities because it is one of

the few species adapted to these sites and it can be

established successfully. It grows in association with

other native species on sites that have been seriously

altered and are occupied by annual weeds. It can be

seeded with bluebunch wheatgrass, slender wheatgrass, bottlebrush squirreltail, Sandberg bluegrass,

and related broadleaf herbs to reestablish native communities and promote secondary succession.

Western wheatgrass has been used to revegetate

mine disturbances. Although establishment from direct seeding is slow, plants spread by rhizome development and stabilize exposed soils (Thornburg 1982).

This grass is particularly useful for controlling erosion

of sandy soils (Scheetz and others 1981) and for providing protective ground cover throughout the entire

year. It establishes quickly on barren mine wastes

that are free of competitive weeds. Established sod is

very persistent, but not restrictive to seedling invasion by some native herbs and shrubs.

Western wheatgrass is also commonly planted on

saline seeps associated with mine disturbances (Rogler

1973). It grows well on loam and clay loam soils and

fine-textured mine wastes, and it persists on soils with

moderate alkalinity (Reitz and Morris 1939; Wambolt

1981), overflow sites, and subirrigated wetlands with

poor drainage. It also withstands short periods of

flooding.

Western wheatgrass recovers from wildfires and

prescribed burning through rhizome proliferation

(Wasser 1982; White and Currie 1983). Aboveground

stems may be burned, but little heat is transferred to

the root meristematic tissue (Gartner and others 1978).

Varieties and Ecotypes

Considerable variability exists among populations

of western wheatgrass, and use of local or adapted

materials is recommended. However, sufficient seed

cannot be harvested from wildland stands to plant

large projects. Consequently, for most seedings, the

best adapted of the currently available cultivars must

be selected.

The Natural Resources Conservation Service in Los

Lunas, NM; the Colorado Agricultural Experiment

Station in Fort Collins; and the New Mexico State

Highway Department in Santa Fe, cooperatively released ‘Arriba’ western wheatgrass in 1973. It was

developed from a collection obtained in 1957 from a

site near Flagler, CO, at an elevation of 4,970 ft (1,530

m) that receives annual precipitation of nearly 16

inches (400 mm). Seed production was increased by

selection through five generations (Alderson and Sharp

1994). Seeds of Arriba germinate rapidly and uniformly, providing vigorous seedlings. Seed production

of this cultivar exceeds that of other tested accessions.

Plants establish quickly and spread rhizomatously.

Herbage yields are good, but less than those of the

variety ‘Barton’ when grown in pastures. Arriba has

performed well on arid mine sites. It has also done well

when seeded on semiarid shrublands. It produces

dark green foliage that is sought by grazing animals.

Young plantings establish quite well, even on infertile

sites. Arriba and ‘Rosana’ are the two most widely

seeded cultivars of western wheatgrass in the Intermountain region.

Barton western wheatgrass was cooperatively released in 1970 by the Natural Resources Conservation

Service Plant Materials Center at Manhattan, KS; the

Kansas Agricultural Experiment Station, and the

Agricultural Research Service. The original seed was

collected from native grassland in Barton County, KS.

It is used in the Midwest to stabilize disturbed soils. It

is the most productive cultivar currently available

(Alderson and Sharp 1994), and it is also strongly

rhizomatous. It has not been extensively seeded in the

Intermountain area.

The Nebraska Agriculture Experiment Station and

the Agricultural Research Service cooperatively released ‘Flintlock’ in 1975. It is a broad-based cultivar

genetically, derived from materials collected in central and southwestern Nebraska and northwestern

Kansas. Flintlock produces aggressive rhizomes. It is

commonly used for conservation plantings, dryland

hay production, and early-season pastures in the Central Great Plains (Alderson and Sharp 1994).

‘Rodan’ was developed from materials collected in

the Missouri River bottoms near Mandan, ND. The

Agricultural Research Service, the Natural Resources

Conservation Service, and the North Dakota Agriculture Experiment Station released it in 1983. Rodan is

primarily adapted to the Great Plains of the United

States and the prairies of Canada. This cultivar was

selected for its seed production and drought tolerance.

It is used primarily for revegetation of disturbed

lands. It is also planted for grazing and hay production
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 337

Chapter 18 Grasses

because it yields well on dry sites (Alderson and Sharp

1994).

The Natural Resources Conservation Service Plant

Materials Center at Bridger, MT, and the Montana

Agricultural Experiment Station released ‘Rosana’

western wheatgrass in 1972. It was selected from

materials harvested in native meadows near Forsyth,

MT, and developed for seeding irrigated hayfields and

pastures. It has also been successfully planted on

rangelands and mine disturbances (Alderson and Sharp

1994). Seed is readily available. Rosana is one of the

most widely used cultivars in the Intermountain region. Its range of adaptability, however, has not been

adequately defined.

Rosana has been used to control weeds and provide

ground cover because it produces a closed sod soon

after establishment. Seedlings are vigorous and establish well, particularly when seeded in mixtures or

planted on harsh sites. Moderate to good amounts of

light blue-green herbage are produced annually.

‘Walsh’ was developed from southern Alberta and

southwestern Saskatchewan collections and released

in 1982 by Agriculture Canada. It is a dual-purpose

cultivar, adapted for pastures and hay production. It

is also extensively seeded to revegetate disturbed

lands in Canada. Walsh is tolerant of saline soils,

drought, and periodic flooding (Alderson and Sharp

1994).

Agropyron spicatum

Bluebunch Wheatgrass __________

Synonyms

Agropyron inerme

Agropyron spicatum var. inerme

Elymus spicatus

Elytrigia spicata

Pseudoroegneria spicata

Description

Bluebunch wheatgrass (fig. 9) is a perennial, coolseason, native bunchgrass (Arnow 1987; Hitchcock

1950; Stubbendieck and others 1992). Plants are rarely

rhizomatous. Stems are erect and 24 to 40 inches (60

to 100 cm) tall. Sheaths are glabrous or puberulent.

Blades are flat or involute, 0.04 to 0.08 inch (1 to 2 mm)

wide, and 2 to 10 inches (5 to 25 cm) long. The upper

surface normally has small hairs, but the underside is

smooth. Auricles are well developed. Ligules are membranous and about 0.04 inch (1 mm) long. Spikes are

typically erect, slender, and 2.4 to 8 inches (6 to 20 cm)

long with a continuous rachis. Spikelets are rarely

more than one per node, five to eight flowered, and

mostly 0.4 to 0.8 inch (1 to 2 cm) long. Those near

midlength on the spike rarely exceed the internodes by

more than one-third their length. Glumes are mostly

unequal, somewhat oblanceolate, three to seven nerved,

0.18 to 0.47 inch (4.5 to 12 mm) long, and obtuse to

acute or rarely awn tipped. Lemmas are 0.28 to 0.51

inch (7 to 13 mm) long, acute or terminating in a widely

divergent awn.

Both diploid (2n = 14) and autotetraploid (4n = 28)

forms occur in the West, but tetraploids are primarily

limited to eastern Washington, northwestern Idaho

(Hartung 1946), and Canada. Dewey (1966) concluded

that bluebunch wheatgrass has been involved in the

phylogeny of several other grasses. Beardless wheatgrass has been designated a separate species by some

authors, but Heller (1900), Beetle (1961), and Cronquist

and others (1977) combined it with bluebunch wheatgrass. Beardless wheatgrass lacks the distinctive divergent awn, a recessive trait of bluebunch wheatgrass (Asay 1995a,b). Stebbins and Fung (1953)

presented cytogenetic evidence demonstrating that

both varieties are completely interfertile. Both hybridize freely and produce fertile progeny. Daubenmire

(1960) concluded that the cespitose growth forms of

Figure 9—Bluebunch wheatgrass is one of the

most important native bunchgrasses (photo courtesy of Kevin Jensen, USDA ARS, Logan, UT).
338 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

bluebunch wheatgrass are adapted to arid grasslands

and semideserts, while rhizomatous forms require

more mesic sites.

The cultivar ‘Secar’ was originally identified and

released in 1980 as a bluebunch wheatgrass. Asay

(1995a,b) later referred to it as “thickspike wheatgrass” because it is morphologically similar to

bluebunch wheatgrass, but genomically aligned with

thickspike. It is now considered a separate taxon,

Snake River wheatgrass (Agropyron spicatum ssp.

laceolata), originally designated Elymus lanceolatus

ssp. wawawaiensis (Carlson 1986), and more recently

Elymus wawawaiensis (Natural Resource Conservation Service 2002). Snake River wheatgrass, which

includes Secar, is most prevalent along the lower

Snake River drainage of central Idaho (Asay 1996).

Since Secar was initially considered a variety of

bluebunch wheatgrass and as it is ecologically and

morphologically aligned with this species, we have

treated it as a bluebunch wheatgrass in this text.

Ecological Relationships and Distribution

Bluebunch wheatgrass occurs from Alaska to

Saskatchewan and south to California, Arizona, New

Mexico, and Texas (Arnow 1987). On the Palouse

Prairie it frequently occurs in pure stands, but it is

most often codominant with Idaho fescue and Sandberg

bluegrass (Miller and others 1986). It is often a dominant species on grasslands of the Pacific Northwest. In

areas of the Intermountain area receiving 7 to 30

inches (150 to 760 mm) of annual moisture it grows in

association with big sagebrush and antelope bitterbrush (Stoddart 1941; Wright and Bailey 1982).

Bluebunch wheatgrass generally occurs on sagebrush

flats, low foothills, and benchlands (Pickford 1932). It

is not highly shade tolerant, but it does occur in

pinyon-juniper, mountain brush, aspen, ponderosa

pine, and spruce-fir forests. It forms distinctive and

important associations with ponderosa pine at mid

and upper elevations in Oregon, Washington, Montana, and Idaho, but it is equally important in more

arid grass or shrub communities on dry, well-drained

slopes.

Bluebunch wheatgrass occurs in some unique plant

communities. It exists in restricted areas with low

sagebrush, alkali sagebrush, and black sagebrush in

Nevada (Zamora and Tueller 1973); pygmy and stiff

sagebrush in Idaho, throughout the extensive steppes

of Washington (Daubenmire 1970); and in the sagebrush-grass steppes of Colorado (Francis 1983) and

southern Idaho (Hironaka and others 1983).

Bluebunch wheatgrass occurs on soils that vary in

texture, depth, and parent materials. It is often found

on coarse, well-drained mountain slopes (Tisdale and

Hironaka 1981). It is not tolerant of alkaline or saline

soils, but is does grow on dry, calcareous soils.

Plummer and others (1968) reported that this grass

exhibits considerable variation. Some beardless and

bearded forms have similar areas of adaptation, but

some exist in differing sites. Plummer and others

(1968) concluded that sod-forming strains are superior to bunch types for seeding game ranges with

variable topography in Utah, because they are adapted

to a wider range of moist to dry habitats. Daubenmire

(1960) noted that caespitose individuals prosper in

arid grasslands in eastern Washington and northern

Idaho, and that rhizomatous types dominate in more

mesic grasslands. Dewey (1970) suggested that the

presence of rhizomes may result from introgression

with thickspike wheatgrass. A greater chance for

introgression would occur at upper elevations where

thickspike wheatgrass is more common. Rhizomatous

types are not always restricted to more mesic conditions, and moderately spreading growth forms are

sometimes found growing in drier situations. Some

rather extensive rhizomatous populations occur in

pinyon-juniper, antelope bitterbrush, and big sagebrush communities. Since this species occurs over

such a wide range of sites, and exhibits considerable

difference in vegetative characteristics, selection and

use of specifically adapted ecotypes is recommended.

Plant Culture

Seeds of bluebunch wheatgrass cultivars are commonly produced under cultivation. Wildland stands

normally produce seed crops most years (Stewart and

others 1939), and plants are capable of producing some

seeds even during periods of drought.

Plants grown under cultivation are usually planted

in rows spaced 24 to 36 inches (60 to 90 cm) apart.

Satisfactory crops can be grown under irrigation or if

sufficient moisture is available, under dryland conditions. If irrigation is required, application is made

prior to pollination or the late boot stage. In fields that

dry rapidly, irrigation soon after harvest may be

required to ensure fall regrowth and tillering. Under

cultivation, bluebunch wheatgrass fields normally

produce 300 to 385 pounds of seed per acre (336 to 430

kg per ha) (Hafenrichter and others 1968). Seeds ripen

early, generally by mid-July. Moderate amounts of

fertilizer are required to maintain productivity.

Bluebunch wheatgrass develops an extensive root

system, and properly spaced plantings are normally

somewhat competitive with weeds. Seeds usually ripen

uniformly, but mature seeds shatter quickly. Fields

can be swathed when seed is in the soft-dough stage

and then combined; however, direct combining is most

practical. Bluebunch wheatgrass seed is easily cleaned,

but awns must be removed from some cultivars and

wildland collections. Infertile florets can be removed

during cleaning. Seed viability from wildland collections may be less than 60 percent.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 339

Chapter 18 Grasses

Bluebunch wheatgrass is best sown in fall, but it will

establish from early spring plantings if moisture conditions are favorable. Seeds can easily be planted with

most drills or other seeders. Seeds are of moderate size

and should be planted at depths of 0.5 to 1.5 inches (1.3

to 3.8 cm). Seedlings are not as vigorous or competitive

as those of cheatgrass (Harris 1967), but they establish and persist well when planted in weed-free seedbeds. Bluebunch wheatgrass seeds germinate fairly

uniformly, seedlings grow rapidly, and mature stands

will develop in 2 to 3 years. Young plants develop

extensive root systems, and after about 2 years they

provide considerable competition.

Dewitt (1969) compared the germination rates of 51

accessions of bluebunch wheatgrass in an attempt to

select sources capable of germinating at cooler temperatures and competing more successfully with

cheatgrass seedlings. Although differences among collections were noted, they were not great enough to be

of practical importance. Improvement of germination

rate and emergence success through selection of larger

seeds from different populations appears possible

(Kitchen and Monsen 1994). As additional ecotypes

are selected for wildland seedings, emphasis on development of seedling vigor and establishment traits

should be emphasized.

Bluebunch wheatgrass has often been replaced by

crested wheatgrass in range and wildlife seeding

projects. However, bluebunch wheatgrass develops

strong stands, and it can be seeded as a major component of native mixtures. Established seedings of

bluebunch wheatgrass are very persistent

(Hafenrichter and others 1968; McGinnies and others

1983; Plummer and others 1955). Native stands eliminated by grazing have been reestablished by direct

seeding. Monsen and Anderson (1993) found that

bluebunch wheatgrass plantings established in southcentral Idaho in 1927 persisted through two wildfires

and prevented invasion of introduced perennial grasses

during a 64 year period.

Bluebunch wheatgrass stands can be restored with

protective management if some residual plants remain. Considerable increase in plant density was

noted on a number of sites in the Intermountain West

between 1989 and 1994, during a period of extended

drought. Although bluebunch wheatgrass seedlings

are not able to compete with annual weeds as successfully as crested wheatgrass seedlings (Harris 1967),

they are quite hardy. Bluebunch wheatgrass can be

seeded in areas where it once dominated to reduce

weed encroachment and provide a permanent cover.

When seeded in disturbances where residual native

species still occur, established bluebunch wheatgrass

usually allows for natural spread and encroachment of

other native species.

Seedings of bluebunch wheatgrass and other adapted

native herbs are required to restore native shrub and

herb associations. Antelope bitterbrush, Stansbury

cliffrose, and mountain big sagebrush have been eliminated from many game ranges through excessive grazing by wild ungulates. In many cases, cheatgrass and

other annual weeds have invaded and become the

dominant vegetation. Reestablishment of native shrubs

by interseeding and transplanting shrubs in clearings

within annual grass stands has been successful, but

without reestablishment of the native understory, the

seeded shrubs are unable to reproduce naturally.

Planting success is dependent, in part, on adaptability of the seeded ecotype. Nearly all seedings using

bluebunch wheatgrass are conducted using seed of

three available cultivars: ‘Goldar’, Secar, and

‘Whitmar’. Although these are useful and widely used

cultivars, each has specific areas of adaptation. Goldar

and Whitmar are better adapted to uplands and most

moist sites, while Secar is better adapted to the lower

ranges, particularly in the Snake River drainage.

None are universally adapted throughout the entire

range of this grass. In addition, none of these cultivars

are well adapted to Great Basin sites. As additional

releases or seed sources become available, planting

success is likely to improve.

Uses and Management

Bluebunch wheatgrass is one of the most important,

productive, and palatable native grasses in big sagebrush communities. It is a major species in sagebrushgrass, pinyon-juniper, and ponderosa pine communities. It is valuable because of its widespread distribution

and because it provides excellent forage in spring,

summer, and fall (Vallentine 1961). It is also used in

winter at certain locations. Bluebunch wheatgrass

begins growth in early spring as soil temperatures

approach 42 ∞F (17 ∞C) and air temperatures reach 58

to 77 ∞F (24 to 32 ∞C) (Anderson 1991). Although

differences in palatability occur among ecotypes

(Willms and others 1980), this grass is palatable to

most classes of domestic stock and big game, and

supplies excellent herbage even after growth has

ceased. It cures well and in many locations it is used as

winter forage by wildlife (Peek and others 1979).

Reestablishment of bluebunch wheatgrass is desirable

over a broad range of sites. This species has been reduced

from extensive areas by poorly regulated grazing

(Anderson 1991; Blaisdell and Pechanec 1949; Forsling

and Dayton 1931; Rickard and others 1975). Loss of

bluebunch wheatgrass and related species has resulted

in rapid invasion by annual weeds, particularly cheatgrass

(Christensen 1963; Cottam and Evans 1945; Daubenmire

1940; Pickford 1932). Christensen (1963) reported that

mechanical disturbances and frequent fire also reduced
340 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

the occurrence of the species. This change in species

composition has diminished forage production and species diversity, and increased fire frequency. As a result of

overgrazing by domestic animals, cheatgrass and other

annual exotic weeds have displaced bluebunch wheatgrass communities across extensive areas of the West

(Christensen 1963; Cottam and Evans 1945; Daubenmire

1940; Pickford 1932). Christensen (1963) reported that

mechanical disturbances and frequent fires have also

reduced the occurrence of this species.

Reestablishment of bluebunch wheatgrass and associated herbs has been hindered by a lack of adapted

seeds and by the competitive ability of cheatgrass and

other weeds. Elimination and control of weeds is

essential if bluebunch wheatgrass communities are to

be restored. Although bluebunch wheatgrass seedlings and those of many other perennial grasses are

not as competitive as cheatgrass and other annual

weeds, reestablishment of bluebunch wheatgrass communities by careful management and artificial seeding are the most practical solutions to weed invasion

and spread.

Bluebunch wheatgrass plants are particularly sensitive to season and intensity of grazing. Most plants

succumb from repeated moderate to heavy clipping in

late spring at the height of the growing season

(Daubenmire 1940). Hyder (1972) found that species

such as bluebunch wheatgrass produce culms late in

the growing season when the apical meristems are

elevated as internode elongation proceeds. These meristems are susceptible to removal by grazing, and

further growth must originate from axillary buds at

the base of the plant. Defoliation during spring, when

vegetative growth is proceeding rapidly and the apical

meristem is elevated, is most detrimental (Stoddart

1946b). Miller and others (1986) concluded that plants

can withstand use early in the growing season if

conditions permit adequate growth after grazing.

Rickards and others (1975) found that 50 percent

use for 2 consecutive years on previously protected

sites resulted in a 26 percent decrease in the basal area

of bluebunch wheatgrass. Heady (1950) concluded

that clipping once to a 6-inch (15-cm) height at flowering was too severe to permit plants to maintain themselves. Wilson and others (1966) found bluebunch

wheatgrass may be eliminated by grazing to a 1-inch

(2.54-cm) stubble height for 3 consecutive years during the boot stage. Mueggler (1975) found that

bluebunch wheatgrass plants growing with competition may require 8 years to approach normal vigor

following late spring clipping.

Bluebunch wheatgrass plants do not curtail root

growth following defoliation (Richards 1984). This

reduces the availability of carbon reserves for shoot

development (Caldwell 1984; Richards 1984). Crested

wheatgrass has often been planted as a substitute for

bluebunch wheatgrass because it has a much greater

capacity to allocate limited carbon resources to foliage

regrowth following clipping or grazing (Caldwell and

others 1981; Caldwell and Richards 1986b; Richards

and Caldwell 1985).

Many bluebunch wheatgrass communities have been

converted to crested wheatgrass and intermediate

wheatgrass seedings because these species withstand

more intensive late spring and summer grazing. Most

wildland sites are not managed as intensive pastures,

and costs for site conversion usually do not justify this

effort. Plummer and others (1955) found that bluebunch

wheatgrass, including beardless bluebunch wheatgrass, is more productive than crested wheatgrass on

poor sites. These investigators also found that on sites

receiving more than 10 inches (250 mm) of annual

precipitation, yields of bluebunch wheatgrass are comparable to those of crested wheatgrass, but fluctuate

less than those of introduced wheatgrasses. Bluebunch

wheatgrass is now widely used to restore native communities. Plantings in the big sagebrush, pinyonjuniper, and ponderosa pine types are successfully

seeded with this species when planted alone or in

mixtures.

Bluebunch wheatgrass is an important species for

wildlife. It provides useful forage, and along with

woody shrubs furnishes cover and protection. Bodurtha

and others (1989) found that mule deer preferred

bluebunch wheatgrass communities that approximated

the composition of climax communities during both

spring and summer grazing in Oregon. Deer and elk

selectively graze Utah pinyon-juniper sites that are

seeded with bluebunch wheatgrass.

Bluebunch wheatgrass furnishes ground cover and

soil protection for steep slopes in watersheds where

erosion is a serious problem. It is particularly important in semiarid environments where loss of ground

cover creates major watershed problems and disturbances are difficult to restore. It provides excellent

cover in ponderosa pine and pinyon-juniper woodlands where seasonal storms can generate considerable runoff.

Bluebunch wheatgrass is fire tolerant and regenerates vegetatively following burning. Only small

amounts of litter accumulate near the base, limiting

fuel buildup and potential fire damage to root crowns

(Antos and others 1983). Leaves and stems are coarse

and burn quickly, transferring little heat to the soil

(Wright 1985). Although bluebunch wheatgrass usually survives fires, postburn production, ground cover,

tillering, and seed production vary among burns. Crown

cover can be reduced over 52 percent following

wildfires (Conrad and Poulton 1966), and foliage

production normally decreases for 2 to 3 years

(Daubenmire 1975b; Muggler and Blaisdell 1958).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 341

Chapter 18 Grasses

Varieties and Ecotypes

Only three cultivars of bluebunch wheatgrass are

currently available. Additional populations or ecotypes

are needed. Individual ecotypes differ in adaptability,

and local collections should be seeded where available

releases are not adapted.

Anatone bluebunch wheatgrass is a selected

germplasm originating from Asotin County, WA, at an

elevation of 3,200 to 3,600 ft (975 to 1,097 m). It

exhibits good seedling vigor and is recommended for

use on bluebunch wheatgrass sites receiving at least

10 inches (25 cm) of annual precipitation (Monsen and

others 2003).

Goldar bluebunch wheatgrass was initially selected

by the Natural Resource Conservation Service Plant

Materials Center at Pullman, WA, but it was moved to

the Plant Materials Center in Aberdeen, ID, where it

was released in 1989. Seed was originally collected

from an open ponderosa pine community on the

Umatilla National Forest near Asotin, WA, at an

elevation of 1,000 to 1,550 ft (310 to 475 m) in 1934.

Goldar was developed by mass selection from spaced

plantings (Alderson and Sharp 1994).

Goldar is a diploid form of bluebunch wheatgrass, an

ecotype of A. s. ssp. spicata (USDA Soil Conservation

Service 1989). This is a vigorous cultivar, representative of material found in northeastern Oregon and

southern Washington. It is a leafy, productive, and

densely tufted bunchgrass that produces some tillers

and short rhizomes. It is relatively drought tolerant

when compared with Whitmar. Goldar produces seeds

with the typical divergent awns of the species; these

must be removed prior to seeding.

Goldar is recommended for seeding rangeland sites

to improve native diversity and to provide wildlife

habitat and forage for grazing animals. It is adapted to

sites receiving 12 to 14 inches (300 to 360 mm) of

annual precipitation. In the Intermountain West, it

has done well on pinyon-juniper and upper benchlands

occupied by mountain big sagebrush. It has survived

when seeded in disturbed Wyoming big sagebrush

communities receiving at least 12 inches (300 mm) of

annual precipitation. It is not as drought tolerant as

Secar, but it is much more palatable and remains

green later in the season. It provides considerable fall

greenup, and develops a very dense and competitive

ground cover. Plants cure well and are grazed as

winter pastures. Overseeding Goldar on depleted big

sagebrush communities by aerial seeding and chaining has been very successful. This cultivar provides

useful herbage in mixed and single-species plantings.

It is particularly useful for grazing in spring and early

summer. Goldar is sensitive to grazing during the

vegetative stages of growth.

‘NewHy’ RS hybrid wheatgrass was developed by

artificially crossing quackgrass (2n = 6x = 42) and

bluebunch wheatgrass (2n = 4x = 28). The Agricultural

Research Service, Forage and Range Research Laboratory, Logan, UT, the Utah Agricultural Experiment

Station, and the Natural Resource Conservation Service released this cultivar in 1989. NewHy was derived

from two germplasms (‘RS-1’) and (‘RS-2’) released in

1980 (Asay and Dewey 1981). The initial RS hybrid

population was established in 1962. These plants were

only partially fertile, but fertility and growth characteristics were improved through an eight-generation

selection process. The primary objectives in the selection process were to combine the vigor, productivity,

salinity tolerance, and persistence of quackgrass with

the drought tolerance, caespitose growth habit, and

seed and forage quality of bluebunch wheatgrass (Asay

and others 1991). Caespitose types were emphasized,

but annual rhizome development of NewHy ranges

between 1.6 and 3.3 ft (0.5 and 1.0 m) per year.

NewHy can grow in saline soils (Currie and others

1986); its salt tolerance approaches that of tall wheatgrass. NewHy is most productive on slightly saline or

alkaline sites receiving about 17 inches (330 mm) of

annual precipitation. Seed quality varies among lots.

When seeded alone, a seeding rate 8 to 17 pounds per

acre (9 to 19 kg per ha) is sufficient. Poor seed germination can result in weak stands, but as plants develop, tillering and rhizome development in the third

and fourth year provide more complete cover. NewHy

provides high-quality forage, comparable to that of

intermediate wheatgrass. Plants begin growth in early

spring and remain green until late in the growing

season. On dryland ranges NewHy provides late spring

grazing and competes well with weeds. It tolerates

moderate grazing and recovers quickly following use,

an improvement over bluebunch wheatgrass.

P-7 bluebunch wheatgrass is a multiple-origin

polycross generating by mating 23 open-pollinated,

native-site collections and two cultivars from Washington, Oregon, Nevada, Utah, Idaho, Montana, and

British Columbia. P-7 is intended to provide increased

genetic diversity upon which natural selection may

operate within a single germplasm. It is intended for

use on semiarid to mesic sites in the Intermountain

Region. P-7 has very good seed yields. Genetically,

P-7 corresponds to the “P” metapopulation, which

includes native populations in southeastern Washington, northeastern Oregon, and western Idaho, the

region from which the great majority of the P-7

components originated (Jones 2002a).

Secar was selected at the Plant Materials Center at

Pullman, WA, from a collection obtained by J. L.

Schwendimann near Lewiston, ID, in 1938. Originally

considered a bluebunch wheatgrass, it was later identified as Snake River wheatgrass. The Agricultural

Research Service; the Idaho, Oregon, Montana, and

Wyoming Agriculture Experiment Stations; and the
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Chapter 18 Grasses

Natural Resources Conservation Service Plant Materials Center in Pullman, WS, released Secar in 1980.

Secar is a densely tufted bunchgrass with many fine

leaves and stems. Seeds are small and mature early in

the season. This cultivar is more drought tolerant and

better adapted to harsh sites than other bluebunch

wheatgrass cultivars. It is slower to establish and less

productive than the more robust cultivars.

Secar is adapted to low-elevation sites in the Pacific

Northwest receiving 8 to 12 inches (200 to 300 mm) of

annual precipitation (Alderson and Sharp 1994). Secar

is also seeded in Wyoming big sagebrush communities

on the Snake River Plain of southern Idaho, and in

northern Nevada and central Utah. It is less productive than Goldar when seeded on more upland sites,

but it produces considerable herbage under arid conditions. Secar is well adapted to sites with long, hot

summers, as it enters dormancy in late spring or early

summer. It forms closed stands when seeded in rows

spaced less than 28 to 30 inches (71 to 76 cm) apart.

Established stands, particularly in low rainfall areas,

are somewhat competitive with annual weeds. Secar

is fire tolerant and recovers well following wildfires.

Herbage is not as selectively grazed as other cultivars

or ecotypes. Secar establishes well when seeded in

mixtures with other natives.

The ‘SL-1’ germplasm was developed and released

by the Agricultural Research Service Forage and Range

Research Laboratory in Logan, UT, in 1990. It is a

hybrid derived from the diploid form of bluebunch

wheatgrass and thickspike wheatgrass. The initial F1

hybrid was made in 1965 by crossing a bluebunch

wheatgrass collection obtained in Utah with a Nevada

collection of thickspike wheatgrass (Dewey 1965). The

F1 hybrid was completely sterile, but use of colchicine

permitted development of a fertile amphiploid population. Further selection was made for improved fertility, seed production, seed size and vigor, herbage

production, leafiness, and drought tolerance (Asay

and others 1991).

Seed set of the SL-1 population approaches that of

the parental species, but seed fertility varies between

34 and 50 percent (Maughan 1988). The hybrid differs

from both parents in being leafier and producing

larger culms and leaves. Rhizome development ranges

from a caespitose growth habit to plants that spread 1 to

3.3 ft (0.3 to 1.0 m) per season (Asay and others 1991).

SL-1 exhibits usefulness in plantings on sites where

either parent occurs. The SL-1 germplasm is quite

vigorous and productive, supplying forage and remaining green late into the summer. Considerable fall

greenup occurs, extending the period of grazing. The

plant is also useful as a ground cover and could be

planted to reduce erosion in semiarid areas. Plants

establish well by direct seeding and appear capable of

controlling weeds.

Whitmar beardless bluebunch wheatgrass was originally released as beardless wheatgrass and is awnless

or nearly awnless. It was released in 1946 by the

Washington, Idaho, and Oregon Agricultural Experiment Stations, and by the Soil Conservation Service

Plant Materials Centers at Aberdeen, ID, and Pullman, WA. Whitmar was developed from material

collected from native Palouse Prairie grasslands near

Colton, WA. Original collections were growing at an

elevation of 2,780 ft (845 m) on a silt loam soil in an

area receiving 20 inches (500 mm) of annual precipitation (Alderson and Sharp 1994).

Whitmar is considered an intermediate growth form.

It is erect or semierect with moderately abundant,

medium-coarse stems and leaves. The cultivar provides good spring and fall herbage and it is a good seed

producer (Alderson and Sharp 1994). Whitmar is primarily used in the Pacific Northwest. Although initially seeded in the Intermountain area, it has been

replaced by Goldar and Secar.

Agropyron trachycaulum

Slender Wheatgrass _____________

Synonyms

Agropyron caninum

Agropyron latiglume

Agropyron pauciflorum

Agropyron subsecundum

Elymus pauciflorus

Elymus subsecundus

Elymus trachycaulus

Description

Slender wheatgrass (fig. 10) is a diverse complex of

several taxa. It is a native, short-lived, cool-season,

strongly tufted perennial that occasionally produces

rhizomes (Cronquist and others 1977; Goodrich and

Neese 1986; Hitchcock 1951; Welsh and others 1987).

Stems are green or bluish green and 20 to 60 inches

(0.5 to 1.5 m) tall. Sheaths are smooth and glabrous or

rarely hairy. Blades are ascending to lax. They are

usually narrow and flat or involute, 2 to 10 inches (5 to

25 cm) long, 0.08 to 0.16 inch (2 to 6 mm) wide, and

scabrous to pilose on the upper surface. Spikes are 1.6

to 8 inches (4 to 20 cm) long and stiffly erect. Spikelets

are overlapping, 0.35 to 0.60 inch (9 to 16 mm) long, and

three to seven flowered. Glumes are glabrous or

scaberulous, 0.25 to 0.60 inch (6 to 15 mm) long, 0.04 to

0.12 inch (1 to 3 mm) wide, and abruptly tapered to an

acute or awn-tipped apex. Lemmas are blunt or end in

a short awn. The second glume is about two-thirds the

length of the lemma.

Slender wheatgrass and four related taxa described

by Hitchcock (1950) have been combined as varieties of
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 343

Chapter 18 Grasses

Agropyron trachycaulum or placed in synonymy with

this species (Cronquist and others 1977). Arnow (1987)

followed Gould (1947) in treating the complex in

Elymus. Bearded wheatgrass is the most important

taxon included in this group. It is similar to slender

wheatgrass in areas of occurrence and growth

characteristics, but it is more productive, robust, and

longer lived (Hafenrichter and others 1968).

Ecological Relations and Distribution

Slender wheatgrass occurs in Alaska, much of

Canada, and southward through the West, Central,

and Northeastern portions of the United States, and in

Northern Mexico (Arnow 1987). It requires at least 15

to 20 inches (38 to 51 cm) of annual precipitation.

Consequently, it is usually found at higher elevations

and on deeper soils than most other wheatgrasses. In

the Midwest, it is largely limited to soils with moderate water-holding capacity (Sarvis 1941). Slender

wheatgrass normally grows in association with aspen,

spruce-fir, and mountain herblands (Mueggler and

Campbell 1986). It is not tolerant of dense shade, and

becomes quite abundant following timber cutting and

fires. It is a common species in open park herblands

(Eckert 1975; Ellison 1954) and open areas in mountain brush and ponderosa pine forests. It is less important in pinyon-juniper woodlands, but it can be quite

abundant in local areas. Slender wheatgrass occupies

some mountain big sagebrush benchlands, and it is a

common constituent in river bottoms and on alluvial

soils in stream and valley bottoms.

Slender wheatgrass can be a relatively short-lived

species, but it reseeds and spreads well by natural

seeding, exceeding most other wheatgrasses in this

characteristic. Long-term ecological studies on the

Wasatch Plateau of central Utah revealed that this

species was one of the few native grasses to persist,

although at a low density, after many years of heavy

grazing (Monsen and others 1996). As grazing was

discontinued, this species regained dominance rather

quickly in some areas, but its position fluctuated with

changes in climatic events and recovery of associated

species.

Slender wheatgrass has been able to persist in many

overgrazed mountain herblands. It persists on sites

where annual weeds, including tarweed, have become

dominant. It invades openings and spreads to occupy

harsh sites. It has also been able to persist amid

disturbances where gophers are extremely destructive. It is not a weedy invader, but it is able to persist

and spread due to its seeding abilities and short-term

growth characteristics.

Plant Culture

Slender wheatgrass produces an abundance of seeds

when grown on wildland sites or in cultivated fields.

Hafenrichter and others (1968) reported that seed

production of irrigated plantings averaged 800 pounds

per acre (896 kg per ha). Plants are self-fertile

(Fulbright and others 1982), and viable seed crops are

produced most every year. Seeds are large, detach

easily, and can be combined and cleaned without

difficulty. Seeds retained in warehouse storage lose

little viability after 5 to 10 years.

Slender wheatgrass normally requires cool

prechilling for germination. Fall seeding is recommended, but seeding high mountain ranges in early

spring or summer is successful if summer rains maintain moist seedbeds. Stewart and others (1939) recommend seeding 5 to 12 pounds of seed per acre (5.6 to

13.4 kg per ha). Seeds benefit from drill seeding and

placement of seeds in the soil at depths of 0.5 to 1.0

inch (1.3 to 2.5 cm). This species can be seeded in

mixtures with other grasses and broadleaf herbs. It is

an important native grass that can be seeded with

slower developing herbs without suppressing their

seedling development.

Slender wheatgrass can serve as an important pioneer species; its seedlings are vigorous and capable of

establishing on harsh sites. In addition, it is able to

establish and compete with weedy species, particularly cluster tarweed and knotweed. It can also be

interseeded into sites disturbed by grazing that support species such as western yarrow, Letterman

needlegrass, dandelion, and Rydberg penstemon to

improve diversity and stimulate natural recovery.

Because it is short lived, the density of slender wheatgrass in undisturbed herblands fluctuates annually,

recovering through natural seeding.

Figure 10—Slender wheatgrass grows at higher

elevations than other native bunchgrasses and

is often used for reseeding forest disturbances

(RMRS photo).
344 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Uses and Management

Slender wheatgrass was one of the first native grasses

included in early restoration plantings (Sampson 1913).

It became a widely used species for planting range and

wildland sites in the Intermountain West (Forsling

and Dayton 1931), the Northern Great Plains, and

Canada (Sarvis 1941). It was initially recognized as a

desirable forage species that is easily established and

adapted to a wide range of sites (Forsling and Dayton

1931). Slender wheatgrass was not used in cultivated

pastures until specific cultivars were developed for

that purpose. The short-lived nature of this plant

resulted in the unexpected decline of some early

seedings, but as this characteristic was better understood, slender wheatgrass became an important revegetation species.

Slender wheatgrass is a dual-purpose species.

Growth begins in midspring and continues through

summer and fall if moisture is available. It is usually

more preferred by cattle than by sheep (Vallentine

1985). Its foliage cures well and provides useful winter

herbage (Sarvis 1941). Although slender wheatgrass

produces a moderate amount of herbage, it is highly

nutritious and palatable. However, the plant can become “stemmy” as it reaches maturity. Slender wheatgrass withstands intense use, but it is not as tolerant

of grazing as western wheatgrass. Plants remain vigorous with moderate to heavy grazing. Grazing at the

time of flowering and seed set, however, can diminish

plant vigor. Slender wheatgrass is preferred by elk,

bighorn sheep, deer, and other wildlife (Hallsten and

others 1987). It provides forage and cover for wildlife.

Animals seek and graze the green herbage throughout

the summer.

Native and seeded stands of slender wheatgrass are

often managed as meadow hayfields or as pastures.

New plantings grown under cultivation remain highly

productive for 3 or 4 years; yields diminish substantially thereafter. The cultivar ‘Primar’ was developed

for pasture use and is seeded with sweetclover or red

clover in conservation and pasture plantings

(Hafenrichter and others 1968). Field production of

mixed seedings can be maintained at a higher level for

longer periods of time than if the grass is seeded alone.

Slender wheatgrass is used primarily to restore

disturbances and rehabilitate native communities. It

is widely planted to reestablish desirable cover on

mountain meadows and open park herblands. It can

be interseeded into existing cover with minimal site

preparation. Once plants become established and seed

crops develop, new plants establish from seed. Native

stands can also be managed to allow for natural

recovery. Slender wheatgrass is commonly seeded in

mixtures with other grasses and broadleaf herbs.

Slender wheatgrass can be used to quickly restore

ground cover and control erosion. This species can be

planted in forested regions following fires to provide

soil protection. New seedings establish quickly, furnishing excellent cover the first growing season. Plants

establish well on open, exposed soil. This grass is also

seeded on overgrazed ranges where soil stability is a

concern (Plummer and others 1955). Price (1938) noted

that differences occurred among ecotypes, and found

that successful plantings could be established in lowelevation communities if adapted ecotypes were

seeded. Bridges (1942) reported the success of seeding abandoned farms, dry meadows, and mountain

mesas in New Mexico, using local ecotypes of slender

wheatgrass.

Slender wheatgrass provides a useful native substitute for intermediate wheatgrass, smooth brome, and

fairway crested wheatgrass seedings in areas where it

is adapted. It can be used in mixtures with other

natives to reestablish native communities. Its wide

range of adaptation from mountain big sagebrush

communities (Beetle 1952) to subalpine types (Stewart

and others 1939) makes this species a useful plant for

restoring native understories and rehabilitating harsh

disturbances.

Varieties and Ecotypes

Five slender wheatgrass cultivars have been released. Because this species includes numerous regional ecotypes, it is important that adapted sources

are planted. Some seed can be harvested from wildland stands, but most seed must be field grown to

supply the amounts required for larger projects.

‘Adanac’ slender wheatgrass was released in 1990

by Agriculture Canada for use in hay and pasture

production in Saskatchewan. It is slightly more productive than ‘Revenue’ for hay production. It is not

commonly grown in the Western United States.

‘Primar’ slender wheatgrass was cooperately released by the Washington, Idaho, and Oregon Agricultural Experiment Stations; the National Resources

Conservation Service Plant Materials Center at Pullman, WA, and the Agricultural Research Service,

Plant Science Research Division in 1946. It was developed from materials collected in 1933 near Beebe, MT.

Primar is a vigorous, early-growing, and long-lived

selection. It is primarily seeded with sweet clover in

pasture mixtures, conservation seedings, or as a green

manure crop to improve soils (Alderson and Sharp

1994). Early spring growth exceeds that of other collections, and plants remain green throughout the

summer. Plants generally persist for fewer than 5

years on sites receiving less than 15 inches (380 mm)

of annual precipitation; they are much longer lived on

sites receiving more than 18 inches (460 mm) of annual rainfall (Schwendiman and Law 1946). Seedings

are extremely vigorous. When grown for hay, this

cultivar provides abundant yields. Plants are free of
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 345

Chapter 18 Grasses

smut, and the species can be seeded for pasture use on

a wide range of soils.

The Natural Resource Conservation Service Plant

Materials Center at Bridger, MT, and the Montana

and Wyoming Agricultural Experiment Stations released ‘Pryor’ in 1988. It was collected along a dry

drainage in a saline upland between Pryor and the

Beartooth Mountains of Montana. It produces large

seeds and vigorous seedlings. Pryor has been used

primarily for conservation and reclamation planting

in Montana and Wyoming (Alderson and Sharp 1994).

It is drought tolerant and adapted to saline soils.

Revenue slender wheatgrass was developed and

released in 1970 by Agriculture Canada, Saskatoon,

Saskatchewan, from seed of a single plant collected in

1963 near Revenue, Saskatchewan. It is recommended

for use as a pasture variety because stands can be

maintained for 3 to 5 years. This cultivar is superior to

Primar in salinity tolerance, establishment, forage

production, and seed yields. It provides considerable

leafy herbage, and plants are free of smut (Alderson

and Sharp 1994). Revenue is widely seeded in pastures and on rangelands in the Western United States.

‘San Luis’ was cooperatively released in 1984 by the

Colorado, Utah, and New Mexico Agricultural Experiment Stations; the National Resources Conservation

Service; and the Upper Colorado Environmental Plant

Center at Meeker, CO. It was developed from plants

collected in San Luis Valley, Rio Grande County, CO.

Plants establish quickly, provide effective cover, and

are relatively long lived. San Luis is recommended for

sites above 5,800 ft (1,800 m) that receive at least 14

inches (350 mm) of annual precipitation (Alderson and

Sharp 1994). It is widely seeded throughout the

Intermountain area, particularly on drier sites than

are seeded with Revenue. It is commonly seeded on

range and wildland sites, including roadway and mine

disturbances.

Agrostis stolonifera

Redtop or Carpet Bentgrass ______

Synonyms

Agrostis alba

Agrostis palustris

Agrostis gigantea

Agrostis depressa

Description

The taxonomic history and resulting synonymy of

this species is complex (Kartesz and Kartesz 1980).

Hybridization has further confused the issue

(Bradshaw 1958). Linnaeus based A. alba on a Poa,

apparently P. nemoralis (Cronquist and others 1977).

Hitchcock and others (1969) applied the name Agrostis

alba and recognized three varieties: A. a. var. alba, A.

a. var. palustris, and A. alba var. stolonifera. These

varieties had previously been treated at the species

level (Hitchcock 1950). Other more current treatments, including Arnow (1987) and Cronquist and

others (1977), combined these varieties into A.

stolonifera.

Redtop, or carpet bentgrass, is an extremely longlived rhizomatous and often-stoloniferous perennial

that sometimes forms large mats (fig. 11). The presence or absence of stolons, rhizomes, or both is reported to be habitat related. Stems are erect to decumbent and range from 0.7 to 4.9 ft (0.2 to 1.5 m) in height.

Leaf blades are flat or folded, smooth to strongly

roughened, and 0.08 to 0.8 inch (2 to 20 mm) wide.

Ligules on the upper leaves are 0.2 to 0.28 inch (5 to 7

mm) long. Panicles are oblong to elliptical, 1.6 to 15.7

inches (4 to 40 cm) long, and 0.4 to 5.9 inches (1 to 15

cm) wide with the branches eventually spreading.

Spikelets are single flowered, 0.06 to 0.1 inch (1.5 to

3.5 mm) long, and green to reddish purple with an

awnless, or rarely short-awned lemma. The common

name redtop refers to the characteristic reddish hue of

the panicle.

Ecological Relationships and Distribution

Redtop is one of the most important and widely

distributed members of its genus in the United States.

Native to Eurasia and North Africa, it was introduced

to North America prior to 1750 as a lawn, meadow, and

pasture grass. It now occurs throughout the northern

temperate region, growing in shallow water, wet meadows, and along streambanks on moderately moist

Figure 11—Redtop, a long-lived, mat-forming perennial introduced as a lawn, hay, and

pasture grass, has spread widely and now

occurs in moist areas of many Western plant

communities (photo courtesy of John Kinney,

USDA Forest Service, Rocky Mountain Research Station, Boise, ID).
346 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

pinyon-juniper, aspen, and spruce-fir communities

(Arnow 1987; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969). The species occurs

over a broad elevation range, from sea level to about

10,000 ft (3,050 m), but mostly below 8,000 ft (2,440 m)

(USDA Forest Service 1937).

Plant Culture

Plants flower from June to early September. Abundant seed crops are produced most years. Seeds are

easily harvested, but they shatter readily when mature. Seed production fields yield about 75 pounds per

acre (84 kg per ha) (De France 1953). Viability of newly

harvested seed is generally high. Seedlots are normally cleaned to at least 90 percent purity; germination is often at least 90 percent. There are 5 to 6 million

seeds per pound (5.6 to 7.2 million per kg) (De France

1953).

Redtop seeds are small and benefit from surface

seeding. Planting depth should not exceed 0.25 inch

(0.63 cm). The species establishes well by broadcast

seeding followed by light harrowing to cover the seed.

This technique works especially well when spring or

fall seeding on wet surfaces, which may crust if compacted by drill seeding. Seeding rates can be easily

adjusted to provide the desired seed density. A carrier

may be added to regulate the seeding rate and seed

distribution. Redtop is normally not seeded alone,

particularly in irrigated or dryland pastures. When

planted in mixtures, seeding rates vary between 2 and

4 pounds per acre (2.2 and 4.5 kg per ha). Rangeland

and watershed seedings normally do not require

heavier seeding rates.

Although redtop seeds germinate early, seedlings

are weak and can be suppressed by other species.

Thus, establishment is often poor. Mature stands,

however, are very persistent. Plantings established

over 50 years ago on the Wasatch Plateau have survived and flourished. Plants develop a low growth

form and spread by producing a vigorous sod. Redtop

does not spread aggressively beyond the original planting site in rangeland seedings, irrigated pastures, or

seed production fields. It does increase in density, and

spreads by extension of the root system.

Uses and Management

Redtop is recognized as a useful cultivated grass for

wetland areas. It has been used extensively with other

species in irrigated pastures (Weintraub 1953). Its

area of distribution is similar to that of Kentucky

bluegrass, but it is more tolerant of acidic, poorly

drained, and clay soils (Balasko and others 1995). It

responds well to fertilization, but it is not a highproducing species for hay production. Although other

wet meadow species may be more productive, redtop

does grow through the summer if moisture is available, and it can provide green herbage when foliage of

other species has dried. Redtop is not highly palatable,

and it does not tolerate close and repeated grazing.

However, it is useful for game and livestock.

Redtop receives considerable use in some areas. It is

seeded in wet and semiwet meadows in aspen and

mountain herblands, particularly in areas where erosion is a major concern (Stubbendieck and Jones

1996). Redtop is also well adapted to acidic soils and

has promise for treating disturbances at high elevations. It is a vigorous grower and will form good turf in

a short time, making it valuable for use as a soil binder

in reclaiming eroded drainages and stabilizing slopes,

banks, gullies, and waterways. It is also commonly

seeded in mixtures with other species, including alfalfa, to provide pasture and hay (Weintraub 1953).

Redtop and several other stoloniferous species introduced from Europe are used extensively in turf culture, especially for use on golf courses.

Redtop has been seeded to protect disturbed riparian sites and high-elevation herblands because seed of

many desirable native species is not available.

Plantings have provided excellent ground cover and

herbage, but they have maintained dominance on

planted sites. Natural recruitment of native species

has not occurred. Redtop should not be recommended

or seeded if recovery of native species is desired.

Varieties and Ecotypes

‘Streaker’ was developed in Idaho for pastures, reclamation, and turf plantings. It is presently the only

cultivar available (Jacklin and others 1989).

Alopecurus arundinaceus

Creeping Foxtail

Reed Foxtail ____________________

Synonym

Alopecurus ventricosus

Alopecurus pratensis

Meadow Foxtail _________________

Synonym

None.

Description

The genus Alopecurus consists of about 50 species

distributed throughout the Temperate and Arctic Zones

of the Northern Hemisphere (Boe and Delaney 1996).

Its name, Alopecurus, is derived form the Greek words
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 347

Chapter 18 Grasses

Alopex meaning fox, and oura meaning tail, referring

to the characteristic cylindrical panicle. The two most

common forage species in this genus are meadow

foxtail and creeping or reed foxtail. These two perennial species are similar in taxonomy, growth form, site

requirements, and uses. Consequently, they will be

discussed together.

Linnaeus named meadow foxtail A. pratensis in

1753, and the name has never been changed. Hitchcock

(1950) classified creeping foxtail as A. arundinaceus,

and this name continues in use. Creeping foxtail and

meadow foxtail are separated taxonomically by the

position and extension of the awn. The awn of creeping

foxtail arises below, or rarely above, the middle of the

lemma. It is straight or slightly bent, and measures

0.04 to 0.1 inch (1 to 3 mm) in length. The awn is either

included or exerted (Arnow 1987). The awn of meadow

foxtail is attached near the base of the lemma and is

bent or geniculate.

Creeping foxtail is a rhizomatous perennial that

ranges from 1.6 to 3.9 ft (0.5 to 1.2 m) in height (fig. 12).

Culms are solitary or in small clusters. Leaf blades are

flat, 0.3 to 0.5 inch (8 to 12 mm) wide, glabrous above

and rough below. Ligules are 0.04 to 0.20 inch (1 to 5

mm) long. Panicles are purple or dark colored, spikelike, cylindric, 1.6 to 3.9 inches (4 to 10 cm) long, and

up to 0.3 inch (8 mm) wide. Spikelets are one flowered

and laterally compressed, disarticulating below the

equal glumes. Diagnostic characters for this species

include the glume tips, dark purple spikelets, and

shortly exerted awns (Arnow 1987).

Meadow foxtail is a stout, nonrhizomatous perennial ranging from 1.3 to 2.6 ft (4 to 8 dm) in height, but

sometimes reaching 3.3 ft (1 m) (fig. 13). Roots sometimes develop from the lower culm nodes. Leaf blades

are flat, roughened on both surfaces, and 0.12 to 0.24

inch (3 to 6 mm) wide. Ligules are 0.08 to 0.14 inch (2

to 3.5 mm) long and covered with short, fine, soft hairs.

Ligule margins are irregularly serrate to entire.

Panicles are 1.4 to 3.4 inches (3.5 to 8.5 cm) long and

0.2 to 0.3 inch (5 to 7.5 mm) wide. Spikelets are ovoid

and one flowered. Glumes are subequal, often with

green nerves, and relatively large, measuring 0.12 to

0.20 inch (3 to 5.2 mm) in length. Keels of the glumes

have long, soft, unmatted hairs; those on the lateral

nerves are appressed. Lemmas are 0.12 to 0.20 inch (3

to 5 mm) long and usually glabrous, but sometimes

with hairs near the keel apex.

Ecological Relationships and Distribution

Creeping foxtail and meadow foxtail are widespread

in Europe and temperate Asia (Strelkova 1938). Creeping foxtail was introduced in the United States in 1935

(Stevens 1963), and is now naturalized in the Great

Plains, Pacific Northwest, and Intermountain regions

(Hafenrichter and others 1968; Sutherland 1986).

Figure 12—Creeping foxtail, a rhizomatous introduction, grows on wet sites over a wide

elevational range. Seeded to provide hay, forage, and erosion control, it has become weedy

along waterways (photo courtesy of Kevin

Jensen, USDA Agricultural Research Service,

Logan, UT).

Meadow foxtail was introduced in North America in

the mid-1800s (Wheeler 1950), and it has naturalized

across Canada and the northern half of the continental

United States (Hitchcock 1950; Scoggan 1978). Both

species are long lived, winter hardy, and adapted to

high-elevation meadow sites, including those with

poorly drained, acidic soils. Both have a high tolerance

to flooded and frozen soils (Stroh and others 1978) and

are capable of persisting on sites where standing

water may be present for more than a month. They are

seeded as pasture and hay crops in wet meadows and

have spread along waterways and moist drainages.

Plant Culture

Panicles mature from the apex toward the base.

Seeds near the panicle apex may shatter before seeds

near the base mature, and seeds can shatter within a
348 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

few hours after reaching maturity (Wheeler and Hill

1957). Consequently, seeds must be harvested when

the greatest number of seeds are mature, but before

they disperse. Seeds tend to darken or turn black

during maturation, a trait that aids in selecting harvest dates. Seeds can be combine harvested by raising

the cutter bar above the leaves to remove seed heads

and some stem material. Harvesting can also be accomplished using strippers that dislodge the seeds by

flailing or brushing. Seed can be harvested when

slightly green if it is allowed to dry before processing.

Seed yields are not usually as high as for other pasture

species, but yields between 268 to 400 pounds per acre

(300 to 450 kg per ha) have been reported (Hafenrichter

1968). Meadow foxtail and creeping foxtail usually

produce good-quality seeds that can be cleaned to

relatively high standards. Seed germination and purity each normally exceed 80 to 85 percent.

Spring or fall seedings can be successful in wet

meadows, as seeds germinate rapidly on moist seedbeds. Ten to 14 pounds of seed per acre (11.2 to 15.7 kg

per ha) is adequate for planting irrigated pastures. In

most rangeland situations, lower rates provide adequate stands. When seeded in mixtures, particularly

with legumes or other pasture grasses, 3 to 5 pounds

per acre (3.4 to 5.6 kg per ha) is sufficient. Seeds can

be distributed on the soil surface or planted in shallow

furrows at a depth of less than 0.5 inch (1.3 cm).

Surface-sown seed should be harrowed or covered

using a light drag. The planted unit consists of a

caryopsis enclosed in a light, fluffy, membranous covering that can bridge or flow irregularly in most

conventional seed boxes. Consequentially, adaptors

are required to move seed uniformly through drills or

other planting machinery.

Creeping foxtail has been seeded primarily on range

and wildland sites. It has established well without

irrigation from both drill and broadcast seedings.

Dispersed seeds germinate quickly, allowing plants to

spread and develop competitive stands rather quickly.

Mature plants compete well with other species, and

ultimately dominate areas where the species is adapted.

Although seedlings of creeping foxtail and meadow

foxtail appear to be weakly competitive when seeded

in cultivated pastures with other species, both plants

are aggressive natural spreaders on rangelands

(Plummer and others 1968). They spread by root

proliferation and natural seeding, particularly the

latter (Lewis 1958b).

Uses and Management

The foxtails were among the first perennial grasses

used to rehabilitate disturbances on mountain rangelands. Both species are well adapted to subalpine

grasslands, aspen forests, and mountain brushlands.

They have been widely planted in irrigated and

nonirrigated pastures in mountain valleys, wet and

semiwet meadows, and riparian zones. They have also

performed well on drier sites where supplemental

irrigation is not available. Plants are equally adapted

to meadow sites in lowlands or in subalpine sites at

elevations over 10,000 ft (2,857 m). Both foxtails are

shade tolerant, and have persisted well as understory

species in aspen forests. Regrowth following grazing

can occur beneath an aspen canopy throughout the

summer, even though these sites are shaded (Piper

1944).

Foxtails are usually planted as pasture or forage

species. They begin growth in early spring, earlier

than most other forage species, including timothy and

orchardgrass. Plants often emerge through snow cover

and begin growth when the soil is still frozen (Stroh

and others 1978). In addition, they produce an early

seed crop that is used by a variety of animals. Plants

continue growth during mild winters (Hafenrichter

and others 1949).

Geese, big game animals, and other wildlife species

are attracted to the foxtails throughout the growing

season, but particularly in spring and fall (Boe and

Delaney 1996). Plants produce considerable herbage

early in the season, and recover quickly from grazing

or clipping, as only about 20 percent of all shoots are

reproductive (Rumburg and Siemer 1976). Domestic

Figure 13—Introduced in the mid 1800s,

meadow foxtail has become naturalized on

high-elevation sites in Canada and the Northern United States (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 349

Chapter 18 Grasses

livestock are also attracted to foxtail plantings in

midsummer because new shoots are produced throughout the growing season. Upland game birds seek the

cover and seeds provided by both species.

Both foxtails are best used to support high stocking

management, but have reportedly depressed animal

gains, possibly due to the presence of an unidentified

antiquality component (Rode and Pringle 1986; Voigt

and Sharp 1995). Both species are grown for hay

production, particularly in high-elevation areas with

short, cool growing seasons where few hay or pasture

species are adapted. On such sites early summer

pastures are important, particularly hayfields that

can be cut or harvested early, but which can recover

and be intensively grazed later in the season.

Foxtails are commonly seeded with legumes to improve yields and forage quality (Boe and Delaney

1996). They have been seeded with various clovers,

birdsfoot trefoil, and alfalfa for pasture and hay production. On rangelands they are more commonly seeded

with alfalfa, although some plantings with cicer

milkvetch have also produced excellent pastures.

Creeping foxtail plants have been noted to spread as

much as 4 ft (1.2 m) over a 3-year period (Stroh and

others 1978). Consequently, they can outcompete associated species, including sedges and rushes. Plants

also form dense rooting systems that provide excellent

soil protection. Hart and others (1980) reported that

creeping foxtail suppressed Canada thistle. These

species have been used on high-elevation disturbances

to stabilize watersheds, waterways, streambanks, and

aspen disturbances. In these situations, however, they

have gained dominance and prevented the recovery of

native species.

Varieties and Ecotypes

‘Dan’ meadow foxtail was developed from material

collected in Poland (Alderson and Sharp 1994). It is

used primarily on low-elevation sites, while ‘Mountain’, developed in Ontario, Canada, is a more upland

variety that is seeded in semiwet meadows associated

with mountain brush, aspen, and conifer communities. Mountain greens up in early spring and again in

fall. Neither variety has been seeded to any extent in

the Intermountain West.

‘Garrison’ creeping foxtail is a pasture and hay grass

that is adapted to waterways, wetlands, and other

areas with high water tables. It was developed from

Western European material growing near Max,

McLean County, ND (Alderson and Sharp 1994). Garrison is seeded extensively in wet and semiwet pasture

renovations. It can be seeded directly into weak stands

of forbs and perennial grasses, including saltgrass. Its

seedlings establish slowly, but established plants are

highly productive and spread aggressively. Garrison

survives periods of complete inundation and exhibits

moderate tolerance of saline and alkaline soils.

Aristida purpurea

Purple Threeawn or

Red Threeawn __________________

Synonyms

Aristida fendleriana

Aristida glauca

Aristida wrightii

Aristida longiseta

Description

The perennial Aristida complex has been treated as

five separate species as listed above. The morphological characters used to distinguish the species involved

a continuum of features including beak length (awn

column), beak twisting, awn length, panicle length,

panicle branching, lemma roughness, and leaf position (Cronquist and others 1977). These characters

apparently vary from one extreme to another in any

given population. Some even appear to differ on the

same plant from year to year depending upon changing environmental conditions. Considerable differences in plant stature and leaf and stem numbers have

also been noted. Due to this extensive variability,

Cronquist and others (1977) included all taxa in

Aristida purpurea and recognized three varieties:

purpurea, glauca, and robusta. Regional ecotypes occurring in specific environments have also been described (USDA Forest Service 1937).

Plants of the threeawn complex are tufted perennials that range from 0.5 to 2.1 ft (1.5 to 6.5 dm) tall.

Culms are smooth and hairless and sometimes

branched at the lower nodes. There are long, soft,

white hairs at the apex of the sheath. Leaf blades are

inrolled or involute, and rarely flat, glabrous to slightly

roughened on the upper side, 0.04 to 0.08 inch (1 to 2

mm) wide, and 2 to 3.9 inches (5 to 10 cm) long. They

are often curved or flexuous on small plants and

straight on large ones. Ligules are scarcely 0.02 inch

(0.5 mm) long and consist primarily of a fringe of hairs.

Panicles are 2.4 to 9.8 inches (6 to 25 cm) long and

narrow to somewhat spreading. Panicle branches are

usually appressed and closely clustered. Spikelets

have narrow, linear glumes of uneven lengths that are

sometimes awned. The lemma is about 0.4 to 0.6 inch

(9 to 15 mm) long (including the awn column), often

grading into a thin, twisted awn column or beak. The

three awns that radiate from the awn column are

subequal in length and have a purple hue. Plants

flower from March to September (Cronquist and others 1977).
350 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Ecological Relationships and Distribution

Purple threeawn is commonly found in dry, sandy

soils of desert valleys, plains, mesas, and foothills. It is

distributed from British Columbia throughout the

Western and most of the Central States and south into

Central Mexico at elevations ranging from 2,500 to

10,000 ft (763 to 3,050 m) (fig. 14). In the United States

it is especially important in the Southwest. Purple

threeawn is often abundant along stock driveways,

roads, and abandoned farmlands. It invades disturbances where animals have burrowed, or where plowing or grazing have reduced or eliminated native

vegetation. It is often found on south- and west-facing

slopes where soil temperatures may be abnormally

high (Evans 1967). At midelevations it grows with

sand dropseed, needle-and-thread, and bluebunch

wheatgrass. At lower and warmer sites it occurs with

sideoats grama, Indian ricegrass, and bottlebush

squirreltail.

Purple threeawn often grows in nearly pure stands.

It may also be found intermixed with other native

bunchgrasses as a codominant species or as a minor

component of bunchgrass/sagebrush communities.

Daubenmire (1970) suggested that purple threeawn

distribution patterns indicate that its presence and

distribution are regulated by edaphic conditions. Purple

threeawn is reported to invade disturbances previously occupied by bluebunch wheatgrass, and it is

often considered an early seral species (Evans 1967).

However, this species is widespread and often present

in undisturbed situations (Küchler 1964). It is not as

heavily grazed nor eliminated as quickly by grazing as

bluebunch wheatgrass.

Plant Culture

Plants begin growth in early spring, generally by

April (Evans and Tisdale 1972). Flowering occurs in

mid-June and seeds mature in early September. Some

viable seeds are normally produced, even during dry

years (USDA Forest Service 1937). The divergent

awns and sharp callus are important for natural

dispersal and seeding, as they anchor and position the

seed in the soil (Evans 1967).

Purple threeawn is not widely used in revegetation

projects. At present, most seed is collected from wildland stands using modified hand or vehicle-mounted

equipment. Seed collection and processing is hindered

by the long awns. Removal of the awns is necessary to

reduce volume for storage and to plant seeds with

conventional drills or other seeding equipment. Fall

seeding at a rate of 6 to 10 pounds per acre (6.7 to 11.2

kg per ha) is recommended if the species is planted

alone. Drill seeding is generally required to place the

seed at a depth of 0.5 to 1.0 inch (1.3 to 2.5 cm). Evans

and Tisdale (1972) reported purple threeawn seeds

required treatment at high temperature to initiate

germination. Little seed germination was recorded

until seeds were subjected to air temperatures exceeding 104 ∞F (40 ∞C).

Purple threeawn has been successfully seeded in

mixtures with other native species. It competes well as

a seedling. Root systems develop rapidly, allowing

young plants to establish and persist, even during

periods of moderate drought. Plants may require 3 to

5 years to fully develop. They recover well following

disturbance, spreading from natural seeding. Developing stands and mature stands allow reestablishment of associated native shrubs and forbs.

Purple threeawn recovers quickly following burning

due, in part, to the production of some seed each year.

Seeds tolerate high temperature resulting from fires

(Sampson 1944), and some may persist in the soil as a

seed bank, aiding in natural recovery over time. Numerous small seedlings of purple threeawn may be

found following fire and other disturbances. This plant

is especially adapted to rapidly establish and spread

due to the awns that facilitate dispersal and planting.

Small and developing seedlings are able to compete

with cheatgrass and other weeds. Seedling survival

and growth of young plants in weedy competition

indicates this species could be used to contain weeds

and facilitate natural community recovery. Mature

purple threeawn plants respond somewhat differently

to burning. Fire can kill clumps that have an accumulation of large amounts of litter, but in most situations

burning may remove dead material but few plants are

killed (Evans 1967).

Figure 14—Purple threeawn is distributed from British Columbia to Mexico (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 351

Chapter 18 Grasses

Uses and Management

Although not generally considered a highly palatable species, purple threeawn produces considerable

early spring forage that is used by livestock and big

game animals (Dittberner and Olson 1983). The USDA

Forest Service (1937) identified it as an important

forage plant in some areas, but noted that considerable differences existed among ecotypes. As awns

develop, plants become less attractive to grazing animals, and can cause injuries if the awn attaches to or

penetrates the skin, face, and eyes. Palatability is

rated as poor to fair as the awns appear. The nutritional value of the species becomes somewhat more

important in midsummer, fall, and winter months

(Evans 1967), as it maintains some green foliage when

other species are normally dry.

Purple threeawn is an important understory species

in many shrubland communities that occupy dry slopes

and sites critical as winter and spring-fall ranges for

big game. In addition, these sites are subject to invasion by annual weeds if the understory is disturbed.

The presence of annuals has resulted in a serious

decline of many woody species due to competition and

increased fire frequency. Consequently, it is important to maintain understory species that are compatible with associated shrubs.

Purple threeawn is adapted to disturbed sites and

soil conditions where few other species exist. Kindell

and others (1996) reported that ecotypes differ in their

adaptation to local conditions. In many areas attempts

to replace these ecotypes with more palatable introduced perennial grasses have diminished species diversity and resulted in a loss of habitat. Restoring and

maintaining purple threeawn is essential to the health

of many rangelands.

Purple threeawn has been successfully seeded on big

game ranges in central Utah, and on disturbed roadways and mined sites in the Intermountain West. It has

established and developed when seeded in mixtures or

alone. Seed prices remain high, as demand is sporadic.

Seed production fields have not been developed. Use of

this species will likely increase as reestablishment of

native species on foothill and lower elevation sites

becomes more important.

Varieties and Ecotypes

There are no releases.

Arrhenatherum elatius

Tall Oatgrass ___________________

Synonyms

Avena elatior

Avena bulbosa

Description

Linneaus (1753) described tall oatgrass (Arrhenatherum elatius), naming it Avena elatior. Its current

binomial, Arrhenatherum elatius, was assigned by

Presl (1819). Tall oatgrass is a short-lived perennial

bunchgrass, often rooting from the first nodes. The

plant is erect and not truly rhizomatous. Culms arise

from the bulblike base and are loosely tufted, ranging

from 2.3 to 5.9 ft (7 to 18 dm) in height. Leaf blades are

flat, smooth to somewhat roughened, and 0.1 to 0.2

inch (3 to 6 mm) wide. Ligules are 0.04 to 0.12 inch (1

to 3 mm) long, obtuse, and covered with very fine,

ciliate hairs. Panicles are 5.5 to 9.8 inches (14 to 25 cm)

long, narrow, and bear flowers to the base. Spikelets

are shiny and two flowered with the first floret staminate and the second perfect. The florets disarticulate

as a unit. Glumes are membranous and unequal,

measuring about 0.2 to 0.3 inch (5 to 7 mm) in length.

The awn of the first lemma is bent, about 0.4 to 0.6 inch

(10 to 14 mm) long, and it is attached midlength on the

lemma. The awn of the second lemma is small, straight,

and attached at the lemma apex. It is rarely absent.

Plants flower from late May through July (Arnow

1987).

Ecological Relationships and Distribution

Tall oatgrass is native to Eurasia, but it is now

circumboreal (Arnow 1987). Introduced to the United

States in 1807, it has been widely cultivated and is

now distributed across the United States (Weintraub

1953) and Southern Canada. It is found in pastures

and hayfields in the Pacific Northwest, the Intermountain and North-Central States, and the desert

Southwest. It is well adapted to subhumid and irrigated conditions (Hafenrichter and others 1949). Although primarily used as a pasture plant, it has

escaped cultivation and spread to waste areas, particularly those with rich soils. It has been seeded on

rangelands and watersheds in the Intermountain

region, and has spread to occupy sites in mountain

brush, aspen, and mountain herbland communities. It

tolerates shaded areas, and has spread to forested and

wooded types. Plants are moderately salt tolerant, and

have been planted in saltgrass meadows in the West

(Weintraub 1953) (fig. 15).

Plant Culture

Tall oatgrass is a rapidly developing species with

vigorous seedlings that is planted to provide pasture

and forage. It is also harvested for seed, as good seed

crops are generally produced each year in cultivated

pastures and on most rangeland plantings. Seeds are

borne on long, narrow panicles. Individual seeds are

light, fluffy, and short awned. Seed crops can be
352 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

difficult to harvest because seeds shatter readily when

mature. Light winds and combines or other machinery

can dislodge the seed and reduce the harvest. Seeds

are difficult to process.

Tall oatgrass can be difficult to seed through drills

because the seeds often clump together to form balls

that impede the flow through the drill. This problem

can be corrected by processing the seed to separate the

fluffy material. When seeded with seeds of other species, tall oatgrass seeds are often carried through the

planting device in the mixture. Tall oatgrass seed can

be distributed adequately by aerial or broadcast seeding methods. Plummer and others (1968) reported

that seeds germinate without special treatment, and

uniform stands are easily established from either drill

or broadcast seedings. New plants grow rapidly and

begin spreading from seed within 2 to 3 years if

rodents or insects do not destroy the seed crops.

Uses and Management

Tall oatgrass has been used primarily as a pasture

and hay crop in the Intermountain area. It is generally

seeded on areas such as mountain meadows characterized by cool, moist conditions. It is commonly seeded

as a mixture with other species to provide diversity

and improve forage quality and yields (Weintraub

1953). The hay is harvested early in the season, and it

is palatable and nutritious (Irwin 1945). Tall oatgrass

produces a reasonable amount of herbage in both

pasture and rangeland plantings. Yields of few other

species exceed those of this grass.

Tall oatgrass is palatable and grazed by game and

livestock. Its erect growth habit and abundant leaves

provide considerable herbage in early spring. Excessive use in the early stages of growth can weaken the

stand (Hafenrichter and others 1968). Although its

palatability diminishes in midsummer and fall on

rangeland sites (Plummer and others 1968), it cures

well when harvested as hay (Stubbendieck and Jones

1996). Ordinarily, tall oatgrass begins to grow a few

weeks earlier and remains green later in the fall than

most other native grasses, thus extending the grazing

period for livestock and wildlife (USDA Forest Service

1937). Plants retain only moderate amounts of edible

forage in late fall and winter on game ranges.

On Western rangelands, tall oatgrass has become an

important species for seeding subalpine, aspen, and

mountain brush communities. It is planted on range

and watershed disturbances to quickly stabilize and

protect the soil. It is also commonly used to seed forest

lands following wildfires. Plants are moderately salt

tolerant, and the species has been used for seeding

saltgrass meadows. Individual tall oatgrass plants are

not long lived, surviving for 6 to 10 years. Natural

recruitment occurs if associated vegetation does not

fill in and occupy openings (Hull and Holmgren 1964).

Grazing management can affect long-term survival of

individual plants and seedings.

Although the ecological impacts of seeding tall

oatgrass on deteriorated aspen and mountain herbland

communities are not well understood, it is apparent

that seeding this species in disturbed situations does

not seriously interfere with natural recovery of native

grasses and broadleaf herbs. Tall oatgrass was planted

on mountain herbland disturbances of the Wasatch

Plateau in the 1930s and 1940s. Excellent grass stands

developed and soil erosion was stabilized. Within 10 to

30 years tall oatgrass essentially disappeared. Natural recovery of native species began before the decline

of the seeded grass. Obviously this grass can better

serve to provide initial ground cover and facilitate the

recovery of native species than more persistent perennial grasses, such as smooth brome, that are commonly used in these situations.

Varieties and Ecotypes

‘Tualatin’ tall oatgrass was developed from materials collected in Chile, Italy, and Belgium (Alderson

Figure 15—Tall oatgrass has been widely

planted on saltgrass sites across the West

(RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 353

Chapter 18 Grasses

and Sharp 1994) in an effort to provide a cultivar that

does not shatter rapidly (Hanson 1972). It has been

seeded at sites throughout the Intermountain West in

Gambel oak thickets, aspen and conifer forests and

associated openings, and in subalpine communities to

provide soil stabilization, forage, and community enhancement. It is not aggressive and can be seeded with

other perennial species. Tualatin grows well in full

sunlight, and it also exhibits good shade tolerance.

Plants are relatively short lived, but they will spread

from seed if the seedheads are not grazed. Tualatin

was selected for its rapid establishment and excellent

forage characteristics. It has not been seeded to any

extent in the Intermountain West. This cultivar is

leafier than most common lines, but seed is not currently maintained or available.

Bouteloua curtipendula

Sideoats Grama _________________

Synonyms

Chloris curtipendula

Cynodon curtipendula

Andropogon curtipendula

Melica curtipendula

Description

Sideoats grama is a widespread, native, warm-season, perennial midgrass. It is the largest grama species, reaching heights of 0.7 to 3.3 ft (0.2 to 1 m)

(Stubbendieck and others 1986). Plants are tufted

with scaly, slender to stout rhizomes. The species is

descriptively named based on the characteristic spikes

that are predominately on one side of the seedhead

(fig. 16). Culms are slender, erect, and smooth with a

purplish tint at the nodes. Leaves are blue green and

flat to involute. The ligule is reduced to a fringe of

hairs, approximately 0.04 inch (1 mm) long. Panicles

are 5.5 to 8.0 inches (14 to 20 cm) long with 25 to 80

pendulous short-stalked, spikelike, branches that disarticulate readily as a unit when mature. Each branch

produces two to eight spikelets, each with one perfect

floret. There is sometimes a vestigial spikelet. Glumes

are papery and often purple tinged. The fertile lemma

is three nerved with the lateral nerves ending as short

awns that are 0.12 to 0.24 inch (3 to 6 mm) long (Arnow

1987; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969). There are two varieties of

sideoats grama that are segregated by growth form: B.

c. var. curtipendula arises from creeping rhizomes

while B. c. var. caespitosa forms clumps.

Ecological Relationships and Distribution

Sideoats grama is widely distributed throughout

much of North America. It ranges from Saskatchewan

east to Ontario, and from Michigan south to southeastern California through Mexico to Central America to

Andean South America (Arnow 1987; Cronquist and

others 1977; Hitchcock 1950; Hitchcock and others

1969; USDA Forest Service 1937), growing on plains,

prairies, and foothills. It often grows in canyons and on

low-elevation mesas, scrublands, weedy meadows, open

woodlands, and steep rocky slopes at elevations from

3,000 to 8,000 ft (915 to 2,440 m). It is especially

abundant in the central and southern mixed prairies

of North America. Rhizomatous B. c. var. curtipendula

occurs over the vast majority of the species range,

while nonrhizomatous B. c. var. caespitosa occurs in

the Southwestern United States and in Central and

South America.

Figure 16—Sideoats grama, a native warm season perennial, is named for the spikes arranged largely along one side

of the inflorescence (Hitchcock 1950).
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Chapter 18 Grasses

Sideoats grama is most abundant on fine-textured

soils ranging from deep to shallow, but it also occurs on

sandy to dry, rocky soils (Nicholson and Bonham

1977). It is better adapted to calcareous or moderately

alkaline conditions than to neutral or acidic soils.

It is fairly drought resistant; root growth continues

during dry periods (Weaver and Albertson 1956). Seminal roots rapidly extend deeper into the soil when

subsurface soils are wet, reducing plant dependency

on fluctuating moisture availability near the surface

(Simanton and Jordan 1986). Abundant rhizome reproduction may then occur following precipitation

events (Weaver and Albertson 1956).

Sideoats grama is climax-dominant or codominant

over much of its range (Great Plains Flora Association

1986). It also occurs in early seral communities on

disturbed sites, and it generally increases following

drought (Weaver and Albertson 1956). In the Intermountain West, sideoats grama can be a climax indicator in big sagebrush, pinyon-juniper, and ponderosa

pine communities, and an associated species in shrub

communities including black greasewood and true

mountain mahogany. It is often found growing with

other grasses such as bluebunch wheatgrass, western

wheatgrass, prairie junegrass, blue grama, and Idaho

fescue.

Plant Culture

Sideoats grama reproduces from rhizomes, tillers,

and seed. Rhizome production and tillering are most

common in sandy soils. Natural reproduction from

seed occurs when sufficient moisture is available for

seed production, germination, and establishment

(Simanton and Jordan 1986). Seed production and

quality varies with precipitation and grazing pressure

(USDA Forest Service 1937). Germination is generally

rapid when sufficient moisture is available. Seedling

vigor is good compared to that of other warm-season

grasses, even on harsh sites.

Seed can be cleaned and processed without damage.

Seeds are small, but easily planted with most seeding

equipment. Shallow planting at depths of 1 inch (2.5

cm) or less is required. Seeds flow freely through drills,

but they can be scattered by wind if broadcast aerially.

In more southern areas of the West, seedings conducted prior to summer rains are more successful.

Late fall and early spring seedings have been more

successful in the northern portion of its range. Drought

tolerance and good germination and seedling vigor

contribute to its establishment and success. Because

of its size and vigorous rate of growth, sideoats grama

is quite competitive, and can be planted in mixtures or

seeded alone. When seeded in mixtures with other

species, it most often develops in small patches. It can

be successfully seeded with a number of native shrubs,

as it does not restrict seedling establishment.

Because sideoats grama is naturally distributed

over a wide area, ecotypes with different areas of

adaptation are abundant. Local seed sources or varieties with known adaptation to the seeding area are

recommended for seeding.

Uses and Management

Sideoats grama provides valuable forage for all

classes of livestock and big game (Dittberner and

Olson 1983). Although plants are palatable and nutritious throughout most of the year, palatability is

greatest in summer and fall and declines in winter

(Stubbendieck and others 1986; White 1986). This

species differs from other warm-season species because it produces important green herbage in spring.

Mature stems are usually not eaten (Towne and

Owensby 1983). Birds seek out the seed, and small

mammals feed on the seeds and foliage (Wasser 1982).

Sideoats grama has been successfully used in rangeland seedings and on critically eroding areas, waterways, and mined lands. If competitive species such as

pinyon, juniper, and big sagebrush are reduced, density as well as forage and seed production of sideoats

grama tend to increase. Following drought, sideoats

grama generally increases dramatically. Although less

tolerant of grazing than blue grama, sideoats grama

tends to replace taller grasses on abused ranges. It will

gradually decrease if continually grazed during the

growing season. This species, however, has the ability

to recover naturally from destructive grazing, but

recovery can be slow in some situations. Strong natural recovery is an important characteristic of this

species because many disturbances within its range

cannot be artificially seeded. Established seedings are

fire tolerant. Sideoats grama is more site specific than

many other perennial grasses, and ecotypes should be

planted on soils and sites where they are adapted.

Sideoats grama has proven only moderately adapted

to harsh disturbances. It is receiving increased use as

an ornamental grass.

Varieties and Ecotypes

‘Killdeer’ was selected for its outstanding vigor,

leafiness, and fair seed production. It was originally

collected near Killdeer, ND, in an area that receives 15

inches (380 mm) of annual precipitation. It has been

seeded with good success in the Dakotas.

‘Niner’ originated near Socorro, NM, at 4,200-ft

(1,280-m) elevation on a site that receives an average

of 9 inches (230 mm) of annual precipitation. It demonstrates superior seedling vigor, strong establishment, and high seed and forage production in dry

areas. It is recommended for use in New Mexico,

Colorado, southern Utah, and northern Arizona.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 355

Chapter 18 Grasses

‘Pierre’ exhibits outstanding vigor, excellent leafy

forage production, and good disease resistance. It has

been successfully seeded in South Dakota and in the

surrounding States and Canadian Provinces. Its origin is near Pierre, SD, in a 16-inch (410-mm) precipitation zone.

‘Vaughn’ originated near Vaughn, NM, in a 13.5-

inch (340-mm) precipitation zone at 5,930 ft (1,810 m).

It is best adapted to areas in western Texas, New

Mexico, southern Colorado, southern Utah, and northern Arizona. It is a good seed and forage producer, and

it has received some use in the Intermountain West.

Bouteloua eriopoda

Black Grama ___________________

Synonyms

Chondrosium eriopodum

Bouteloua brevifolia

Description

Black grama is an important native, warm-season,

perennial grass that exhibits a variety of growth forms

over its range of occurrence. Tufted plants 10 to 28

inches (25 to 70 cm) in height develop from a hard,

knotty, woolly base. The slender culms are solid,

densely white-woolly, spreading, and sometimes

stoloniferous at the lower nodes. They remain green

up to the second node from the stem apex during some

winters. Clusters of leaves emerge from the nodes in

spring. Leaves are mostly basal, narrow, and pointed.

They are generally twisted and involute near the top,

hairy above, and smooth below. Roots are finely divided and generally confined to the upper 10 inches (25

cm) of soil (Arnow 1987; Cronquist and others 1977;

Great Plains Flora Association 1986; Judd 1962).

Inflorescences consist of one to six slender, comblike

spikes borne on the sides of the inflorescence (fig. 17).

The spikes are narrow, flaglike, and do not drop at

maturity.

Vegetative growth begins as soon as adequate moisture becomes available. The growth period usually

lasts from July to October, depending upon temperature and precipitation. The vigor of black grama plants

is largely determined by the amount of precipitation

received during the previous summer (Canfield 1939).

Flowering occurs from June or July through August.

Seeds usually mature in late September and disperse

in October and November (Nelson 1934).

Ecological Relationships and Distribution

Black grama is a major warm-season grass of arid

and semiarid grasslands in Texas, New Mexico, and

Arizona. It also occurs in southern California, Utah,

and Wyoming. It is a climax species in a number of

grassland communities and occurs in early seral communities on disturbed sites. It is most often found in

grasslands on dry, rocky hills and uplands and lower

elevation mesas on well-drained, sandy and gravelly

soils. Common associated species include dropseeds,

threeawns, broom snakeweed, mesquite, and yucca.

Black grama can survive on harsh sites characterized

by low rainfall, high temperatures and evaporation,

and strong winds (Nelson 1934). It is generally considered tolerant of drought. During dry periods, however,

large tufts have been observed to break up and form

smaller clumps, with a reduction in ground cover, but

an increase in plant number.

Plant Culture

Stands are maintained and increased through seed

production, stolon proliferation from old tufts, and

tillering (Heizer and Hassell 1985). Reproduction varies widely with precipitation and grazing pressure

(Canfield 1939). Two periods of flowering may occur in

moist years (Canfield 1939). During a drought year,

very little, if any, flowering occurs, and seed production is limited (USDA Forest Service 1937). In addition, uneven distribution of precipitation in summer

may prevent seed from maturing (Nelson 1934). Black

grama seed fill and germination are often poor. Heizer

Figure 17—Black grama inflorescences consist

of one to six slender, comblike spikes (Hitchcock

1950).
356 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

and Hassell (1985) reported seed fills of only 7 to 10

percent for seed from native stands, while Campbell

and Bomberger (1934) reported that germination may

be as high as 41 percent once every 10 years.

In favorable years ground cover is increased through

tillering, perhaps the most important means of regeneration (Campbell and Bomberger 1934). On ungrazed

ranges, black grama forms uneven stands of large

tufts that die in the center during drought, forming

many smaller tufts. When favorable conditions prevail, these small tufts enlarge through lateral spread,

and can again form a continuous cover (Nelson 1934).

Stolons required two growing seasons to establish new

plants. New stolons are produced during one growing

season, while tillers develop and root during the second growing season (USDA Forest Service 1937).

Uses and Management

Black grama is used by all classes of livestock and

wildlife (Stubbendieck and others 1986), and it was

once a mainstay forage for livestock production in

many areas of the Southwest. It is highly palatable

and nutritious in summer and winter. During winter

and early spring it can provide forage when other

sources are scarce or unavailable (Canfield 1934).

Although black grama can withstand recurrent grazing by livestock, it spreads very little on heavily grazed

ranges. Excessive use impairs its vigor, and overgrazed plants will die out during drought periods.

Tillering is dependent on plant vigor during the previous year. Repeated cropping of black grama to 2 inches

(5 cm) may prevent lateral spread. Damage may be

especially severe in overgrazed areas that have deteriorated as a result of wind erosion (Canfield 1939).

Regeneration by stolon production can occur during

favorable years on conservatively grazed ranges

(Nelson 1934). Stolon production is negligible on ranges

that are excessively grazed year after year. In addition, drought-induced mortality and competition from

less desirable and more competitive species can lead to

a loss of black grama plants and a reduction in ground

cover and forage.

Black grama is seldom used in range seedings because it is a slow starter and it is best adapted to harsh

environments. Heizer and Hassell (1985) recommended

that black grama be seeded only to stabilize soil on

critically eroding areas. Black grama has good potential for ornamental use.

Varieties and Ecotypes

As a result of low seed quality, high seed costs, and

the difficulty of establishment, black grama has not

been seeded to any great extent. Two varieties are

available; little research has been conducted to develop additional varieties.

‘Nogal’ is less erect than ‘Sonora’ and adapted to

drier areas. Seed production is generally low, and

supplies are limited and expensive (Alderson and

Sharp 1994).

Sonora demonstrates outstanding leafiness, vigor,

forage production, and vegetative spread, along with

good seed set and seed production. It is adapted across

much of the species range. Seed is expensive and

supplies are very limited.

Bouteloua gracilis

Blue Grama ____________________

Synonyms

Actinochloa gracilus

Atheropogon gracilus

Bouteloua oligostachya

Description

Blue grama is a low-growing, densely tufted, warmseason perennial bunchgrass. The short rhizomes that

develop from its base often create thick mats. Culms

are 6 to 12 inches (15 to 30 cm) tall. Leaves are grayish

green and turn to gray or straw colored with age. They

are persistent, numerous, mostly basal, and sometimes hairy around the collar. Leaf sheaths are smooth

to silky haired. Blades are flat to involute, characteristically curled or rolled inward with maturity, roughened above and smooth below, 0.04 to 0.08 inch (1 to 2

mm) broad, and 2 to 6 inches (5 to 15 cm) long. Ligules

are sometimes membranous at the base and often

fringed with hairs that are about 0.01 to 0.03 inch (0.2

to 0.7 mm) long. Inflorescences usually consist of two

short, comblike pedicellate spikes that range from 0.5

to 2.0 inches (1.2 to 5.3 cm) in length and become sickle

shaped at maturity (fig. 18). Each spike bears about 30

to 80 fertile spikelets, terminating with a single terminal sterile spikelet. Fertile spikelets produce one perfect floret and one or two sterile terminal florets.

Glumes are one nerved. The first glume is 0.08 to 0.14

inch (1 to 3.5 mm) long, linear, and translucent. The

second is subequal to the lemma, 0.16 to 0.24 inch (4 to

6 mm) long, purplish, and three awned. The terminal

awn arises between two teeth and is 0.02 to 0.06 inch

(0.5 to 1.5 mm) long. Lateral awns are 0.04 to 0.08 inch

(1 to 2 mm) long with short, stiff hairs along the nerves.

The callus, a swollen node at the base of the lemma, is

densely hairy (Arnow 1987; Hitchcock 1950). Spikes

become golden brown and tend to curve backward at

maturity or when the plant is suffering from drought.

Seeds do not disseminate rapidly at maturity.

Blue grama produces an efficient, widely spreading

root system (Briske and Wilson 1977). Numerous

roots develop from short rhizomes. Fine fibrous roots
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 357

Chapter 18 Grasses

are mostly concentrated near the soil surface, but

some extend deeper.

Ecological Relationships and Distribution

Blue grama is a widely distributed climax species

that occurs throughout the Great Plains from Alberta

and Manitoba south to Mexico, and from southern

California eastward from southern Nevada and Utah,

Arizona, New Mexico, and Texas (Great Plains Flora

Association 1986; Hitchcock 1950; Stubbendieck and

others 1986). Blue grama is a major component of the

short grass prairie. It is also found in scattered sites in

the Eastern United States. In the West, it is the most

widespread and important species of its genus. It

occurs in grass and sagebrush communities on dry

plains and foothills, and sometimes in pinyon-juniper

and ponderosa pine woodlands and mountain meadows

of the Intermountain area. At midelevations it grows on

sandy loam or rocky soils. It is often associated with

galleta, sand dropseed, western wheatgrass, bluebunch

wheatgrass, needle-and-thread, and broom snakeweed.

Blue grama exhibits considerable variability in its

growth habit and general appearance throughout its

north-south range. In the north it tends to form a sod

under favorable moisture conditions. In the south and

at lower elevations it tends to form patches and not a

complete cover. It is also a taller, more vigorous bunchgrass (Griffiths 1912). Blue grama normally remains

dormant in spring, greening quickly with summer

precipitation. It is peculiar in that it flowers from July

through September. Because of its root system, blue

grama is highly drought resistant (Ward and others

1990; Wilson and Briske 1979; Wilson and others

1976); it is one of the most important soil-binding

grasses where adapted in the arid West. It has the

ability to become dormant during drought periods, but

resumes growth when temperatures and moisture

conditions become favorable. It is a rapidly growing

species, reaching maturity in 60 to 70 days (USDA

Forest Service 1937). It produces highly palatable

forage in late summer and early fall when other forage

is scarce. Blue grama is resistant to grazing, but it is

most productive when lightly grazed in late summer.

It is highly fire tolerant, but it is not shade tolerant.

Blue grama is adapted to a variety of soil types, but

it most abundant on well-drained, sandy and sandy

loam soils of uplands. It can also be found on alkaline

and coarse-textured soils. In Arizona it is common on

infertile, acidic soils (Nicholson and Bonham 1977). It

is a common and important species on rocky outcrops,

including mesas, canyon slopes, deep sandy soil on

plains, and some midelevation grasslands.

Plant Culture

Blue grama reproduces from seed, rhizomes, and

tillers. Good seed crops are produced during years

with favorable moisture conditions. Seed quality is

generally low, varying with year and location. Wind,

water, and animals disperse the seed (Albertson and

Weaver 1944). Adequate surface soil moisture is required for a 2- to 4-day period to support germination

and initial seminal root growth. Lacking lateral seminal roots, seedlings die when the surface soil dries out

before the seminal root elongates to deeper soil

(Wicklow and others 1984). Critical factors for successful establishment of blue grama seedlings are

seminal root initiation, good soil moisture for at least

6 weeks after emergence, and initiation of adventitious roots (Bock and Bock 1986; Albertson and Weaver

1944).

The entire spikelet, including the awned fertile

floret and the awned and densely bearded rudimentary floret(s), is planted so the fertile florets are not

Figure 18—Blue grama spikes curve backward and

are golden brown when mature (Hitchcock 1950).
358 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

separated during cleaning. These units are light and

fluffy and cannot be seeded through a conventional

grain drill. Specially designed seed drills facilitate

uniform flow from the seed box to the soil. Shallow

planting on firm seedbeds improves seeding success.

As a result of its wide distribution, blue grama is

highly ecotypic. It is important to use seed sources

adapted to the edaphic and climatic conditions of the

area being seeded.

Uses and Management

All classes of livestock and many wildlife species

graze blue grama (Dittberner and Olson 1983). Highquality forage, drought tolerance, and a desirable

growth habit make it one of the most important range

grasses in areas where it is adapted. Although more

palatable than most co-occurring species, blue grama

has increased on many overgrazed ranges (Bock and

Bock 1986; Weaver and Albertson 1956). Its persistence is attributed to its vigorous root system, rhizomatous habit, and its ability to maintain adequate

root carbohydrate reserves following grazing (Buwai

and Trlica 1977). Following grazing, some lower leaves

generally remain near the soil surface and provide

photosynthetic surface for plant recovery. With continuous heavy grazing and trampling, particularly on

fine soils, blue grama may become sodbound and

decline in forage production.

Blue grama is a rapidly growing species and usually

matures in 60 to 70 days. It is tolerant of grazing, but

yields the greatest return when grazed lightly in

summer during its period of rapid growth. It can also

be utilized in fall and winter. Grazing has often eliminated the upright forms, leaving the less productive,

sodforming types. This is especially true in the Southwestern United States.

Blue grama can be nutritious throughout the year.

It greens up early and grows rapidly following summer

storms. Its fine leaves are low in fiber and high in

protein when green, making it an exceptional forage.

Protein and carbohydrate content are greatest during

early bloom, decreasing toward maturity. About 50

percent of its nutritive value, however, is retained at

maturity, making it an excellent fall or winter forage

(National Academy of Sciences 1971). In the Southwest, productivity of the shorter growth form of this

species is relatively low. Growth of the shorter variety

begins in May and June, providing very little green

forage during spring and early summer.

Because of its drought hardiness and ability to

protect soils, blue grama is used extensively for protection of critical eroding areas. It is also used for watershed stabilization and mined-land revegetation, as

well as for low maintenance turf plantings.

Varieties and Ecotypes

‘Hachita’ was collected from an elevation of 4,000 ft

(1,220 m) in the Hachita Mountains of southern New

Mexico in an area with an average annual precipitation of 10 inches (250 mm). Hachita exhibits greater

drought tolerance and somewhat better seed and forage production than ‘Lovington’. It is well adapted to

areas in New Mexico, southeastern Colorado, and the

panhandle area of Texas and Oklahoma that receive at

least 8 inches (200 mm) of annual precipitation. It has

been seeded with some success in the four corners area

of Utah, Colorado, Arizona, and New Mexico (Alderson

and Sharp 1994).

Lovington is a good forage and seed producer and is

generally well adapted in areas of eastern New Mexico,

southeastern Colorado, and northwestern Texas that

receive at least 12 inches (300 mm) of annual precipitation. It has not been seeded to any extent in the

Intermountain West.

Bromus anomalus

Nodding Brome _________________

Synonyms

Bromus frondosus

Bromus ciliatus var. montanus

Bromus ciliatus

Fringed Brome__________________

Synonyms

Bromopsis ciliata

Bromus richardsonii

Description

Nodding brome is a tall, nonrhizomatous perennial

existing as small tufts with culms 24 to 30 inches (6 to

10 dm) tall. Culms are typically puberulent, at least at

the nodes. Leaf blades are flat, 0.12 to 0.20 inch (3 to

5 mm) broad, sometimes slightly incurved or involute,

and smooth or sometimes with silky hairs. Ligules are

small, 0.02 to 0.04 inch (0.5 to 1 mm) long. and slightly

irregularly toothed. Panicles are 2.8 to 5.5 inches (7 to

14 cm) long, loose or open, and usually nodding.

Spikelets are 0.67 to 1 inch (17 to 25 mm) long and

slightly compressed. They produce 5 to10 flowers.

Glumes are mostly minutely pubescent, or sometimes

hairless. The first glume is 0.20 to 0.28 inch (5 to 7 mm)

long and three nerved, or rarely one nerved. The

second glume is 0.25 to 0.31 inch (6.5 to 8 mm) long and

three to five nerved. Lemmas are 0.35 to 0.47 inch
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 359

Chapter 18 Grasses

(9 to 12 mm) long, five to seven nerved, rounded on the

back, and hairy. The awns are relatively short, about

0.06 to 0.12 inch (1.5 to 3 mm) in length.

Nodding brome closely resembles fringed brome,

and is known in some cases to hybridize with this

species. Fringed brome is also a nonrhizomatous,

tufted, native perennial with drooping or ascending

branches (fig. 19). Arnow (1987) reported that although both species occupy similar habits in Utah,

they appear to be maintaining their integrity as distinct species. Only 3 percent of all specimens examined exhibited any intermediacy. Nodding brome typically has pubescent glumes and awns are 0.04 to 0.16

inch (1 to 4 mm) long. In comparison, the glumes and

culm nodes of fringed brome are glabrous, and its awns

range in length from 0.08 to 0.24 inch (2 to 6 mm) long

(Arnow 1987).

Ecological Relationships and Distribution

Both nodding brome and fringed brome range from

British Columbia and Saskatchewan south through the

Western United States and Mexico, and east to the

Dakotas and Texas (Arnow 1987; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969). In

the Intermountain region, nodding brome and fringed

brome are found on overlapping sites. Both occur on a

variety of habitat types including dry to moderately

moist sagebrush, mountain brush, aspen, spruce-fir,

ponderosa pine, lodgepole pine, mountain herblands,

grassy meadows, and riparian communities at 5,460 to

10,890 ft (1,680 to 3,350 m) elevation (Arnow 1987;

Bartos and Mueggler 1981; Bradley and others 1992a).

Nodding brome is more common in drier, more open

communities in the mountain brush, pinyon-juniper,

ponderosa pine, and aspen types than fringed brome.

Mueggler and Campbell (1986) reported that fringed

brome is more common in shaded aspen and conifer

communities, and nodding brome is more prevalent in

open communities at lower elevations. Both species

occur as early, mid, and late seral species. They often

occur as dominant understory species in subalpine fir,

Engelmann spruce, aspen, and blue spruce types

(Alexander and Ronco 1987; Bradley and others 1992a;

Moir and Ludwig 1979). Both nodding brome and

fringed brome are important species in aspen and

conifer forest types including open meadows and grasslands. In addition, nodding brome is equally important in mountain brush, pinyon-juniper, and sagebrush types, occurring in various shrub and herb

associations (Allman 1953; Bartos and Mueggler 1981).

Fringed brome is commonly encountered on moist to

seasonally dry habitats, including shaded and open

parks. It grows well on semiwet or moist soils in valley

bottoms and meadows. It is adapted to poorly drained

conditions (Mattson 1984), but also grows on welldrained and rocky substrates (Butterwick and others

1992; Crouch 1985).

Plant Culture

Nodding brome and fringed brome have not been

extensively used in restoration programs. Limited testing has been conducted to evaluate either species for

range or watershed plantings. Both species exhibit

similar characteristics in relation to seed germination

and stand establishment. Seed viability and germination of wildland collections are normally quite high.

Seeds of both species can be harvested, cleaned, and

seeded as easily and successfully as other species of

brome. Plummer and others (1968) reported that seeds

of nodding brome germinate easily without special treatment. Hoffman (1985) found that seeds of fringed brome

are nondormant, and nearly all seeds germinated with

conventional treatments. Initial establishment and

growth of both species is normally greater than for

smooth brome. Plants of both species tend to form small

clumps, and do not persist as long as sodforming species

of brome.

Both nodding brome and fringed brome usually form

full stands when seeded alone, even on irregular

terrain. They provide more open stands than

sodforming bromes, including smooth brome. Nodding

brome plantings established in central Utah persisted

as nearly solid stands for 10 to 30 years, but were slowly

invaded by associated grasses and broadleaf herbs.

Although stands decreased in density, the grass remained important and spread by natural recruitment

Figure 19—Nodding brome forms small,

nonrhizomatous tufts with loose, open, panicles

(Hitchcock 1950).
360 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

to adjacent openings. Natural spread of both nodding and

fringed brome is quite successful, equaling that of most

perennial grasses adapted to mountain communities.

Seeds of both nodding brome and fringed brome are

large and easily seeded through most drills. Seedlings

of both species also establish well from broadcast

distribution and harrowing to cover the seed. Seeds

can be aerial seeded under aspen, oak, or other shrubs

and trees prior to leaf fall. Planted in mixtures or

alone, both species can be seeded at rates used for

smooth brome, between 2 to 4 pounds per acre (2.2 to

4.5 kg per ha).

Nodding brome and fringed brome are highly compatible with other species and can be seeded in mixtures without suppressing establishment or growth of

associated herbs. Both species are less competitive

than smooth brome and could be planted in areas

where smooth brome has been recommended.

Uses and Management

Both species furnish excellent forage and ground

cover. They are rated as among the best forage grasses

on Western rangelands (Dayton 1931). Plants are

grazed readily in spring and summer by livestock,

deer, and elk (Humphery 1960). The foliage of both

species can become tall and rank at maturity and is

less palatable in late summer. However, both species

can be removed by close grazing (Humphrey 1960).

Livestock and wildlife graze seedheads before and

after the seeds have matured. New basal leaves are

produced in fall and selectively grazed. Domestic sheep

seek the plants in mid and late summer as seeds

mature and after fall greenup.

Nodding brome and fringed brome have often been

misidentified and planted in areas where they are not

adapted. Nodding brome is better suited to open aspen

communities and sites at lower elevations. It can be

planted in the upper sagebrush and mountain brush

types to provide forage for wildlife. This species should

be more widely used to plant oak brush sites following

wildfires to control erosion and reestablish a native

understory. It can also be planted in pinyon and

juniper restoration projects where it is native. Nodding brome is not as productive as smooth brome or

mountain brome. It is less palatable than smooth

brome, but equal to mountain brome. Fringed brome

normally grows at somewhat higher elevations than

nodding brome, and it is better adapted to shaded

communities. It is also more common in tall forb types

at high elevations.

Nodding brome and fringed brome can be planted on

open disturbances, including forested sites and mountain brush fields burned by fires. They can be mixed with

other species, including natives, and planted by

aerial seeding to control erosion and restore native

communities. Where the distributions of these two

bromes overlap, mixed plantings are recommended. Although neither nodding brome nor fringed brome form

dense sod, both establish quickly and provide satisfactory cover. Both prefer deep and rich soils, but they are

also encountered on shallow, well-drained soils.

Nodding brome and fringed brome have been grown

to a limited extent in hay meadows, providing yields as

high as 2 tons per acre (2,250 kg per ha) (Dayton 1931).

Seed production fields have not been established for

either species, thus large seedings are not feasible.

Observation and management of wildland stands suggest both species could be grown under cultivation

without special care. Both species grow well in association with other herbs, and are useful grasses for

restoring disturbed watersheds. Plants establish

quickly from direct seeding, and spread to stabilize

small openings. Neither species is invasive; both can

be seeded in mixtures with native broadleaf herbs to

restore complex communities.

Both nodding brome and fringed brome should be

utilized in restoration plantings, and could be used to

replace smooth brome where reestablishment of native communities is required. To date, ecotypes of

these species have not been identified, but seed should

be collected and planted within areas of occurrence.

Moving high-elevation collections to lower and drier

sites is not advised.

Varieties and Ecotypes

There are no releases.

Bromus carinatus

Mountain brome, California

brome _________________________

Synonyms

Ceratochloa carinata

Bromus marginatus

Bromus breviaristatus

Bromus polyanthus

Description

Mountain brome is a member of a highly polymorphic complex of related taxa (Arnow 1987). It has

sometimes been recognized as three or more separate

species (Hitchcock 1935), usually including B.

marginatus, B. polyanthus (both perennials with awns

less than 7 mm long), and B. carinatus (an annual or

biennial with awns more than 7 mm long). B.

marginatus is pubescent throughout, while the spikelets of B. polyanthus are glabrous (Cronquist and

others 1977). B. marginatus and B. carinatus are
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 361

Chapter 18 Grasses

morphologically quite variable but scarcely distinct

from each other. B. marginatus interbreeds with both

B. carinatus and B. polyanthus (Wasser 1982). Plants

with characteristics of all three species can sometimes

be found within a single population. Hitchcock and

others (1969) concluded there was no basis for the

division of these taxa based on their annual or perennial growth habits. The following description of B.

carinatus follows Arnow (1987) and Cronquist and

others (1977) and includes these taxa.

Mountain brome is a tall, short-lived native perennial that often flowers during its first growing season

(fig. 20). Culms are solitary or tufted, usually erect, 1.0

to 2.3 ft (6 to 14 dm) tall, and smooth to minutely hairy.

Leaf blades are flat, 0.08 to 0.55 inch (2 to 14 mm) wide,

rarely involute, and smooth to roughened or silky

haired. Ligules are 0.04 to 0.12 inch (1 to 3 mm) long,

smooth or with silky hairs, and irregularly serrate

along the margins. Panicles are usually 4 to 12 inches

(1 to 3 dm) long, loosely contracted, and nodding.

Spikelets vary from one to four per branch. They are

0.6 to 2.4 inches (1.5 to 6 cm) long, strongly compressed, and produce 4 to 16 flowers. Glumes are

lanceolate and strongly keeled. The first glume is 0.27 to

0.43 inch (7 to 11 mm) long and three- to five-nerved.

The second glume is 0.35 to 0.51 inch (9 to 13 mm) long

and five to seven nerved. Lemmas are 0.43 to 0.67 inch

(11 to 17 mm) long, carinate keeled, and smooth to

roughened. The apex is slightly notched to entire.

Awns are 0.16 to 0.31 inch (4 to 8 mm) long. Anthers

are small and included within the lemma.

Ecological Relationships and Distribution

Mountain brome is widespread in North America,

occurring from Alaska and Canada south to California, Texas, and most of Mexico. It has been introduced

in some Midwestern States and naturalized in Europe

(Arnow 1987; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969). Mountain brome is

common in both early and late seral communities. It

occurs in the subalpine fir-Engelmann spruce zone

and in high-elevation sedge meadows (Tisdale and

Bramble-Brodahl 1983). In the Intermountain area, it

is commonly found in aspen and conifer communities

and in associated herblands and meadows (Ellison

1954; Mueggler and Campbell 1986). It also occurs in

mountain big sagebrush communities. It is more common on cooler sites (Mooney 1985) in the lower elevations of these communities (Hironaka and others 1983;

Tisdale and Bramble-Brodahl 1983).

Plant Culture

Mountain brome reproduces primarily by seed

(USDA Forest Service 1937), although some tillering

occurs (Stubbendieck and others 1992). Plants flower

from late May to early August. Harlan (1945) reported

that cross-fertilization is most common when growing

conditions are optimum, while adverse conditions

stimulate self-fertilization. In California, both open

pollinating and cleistogamous florets are commonly

found on the same plant (Hickman 1993).

Native stands frequently produce heavy seed crops,

and a limited amount of wildland-harvested seed is

sold each year. Agricultural production is required to

meet the demands for seed of this species, and highly

productive mountain brome seed fields have been

established on low-elevation farmlands. Seeds are

harvested by combining, and yields average about 250

pounds per acre (280 kg per ha) (Smith and Smith

1997). There are about 103,700 seeds per pound

(228,000 per kg) (Smith and Smith 1997).

Seed germination of mountain brome is relatively

high, normally exceeding 85 percent. Plummer (1943)

reported a higher percentage of seeds germinated in a

shorter time when exposed to alternating rather than

constant temperatures during germination. Paulsen

(1970) found that optimum germination occurred over

a wide range of temperatures.

Satisfactory wildland stands can be established from

either fall or spring plantings in weed-free areas.

When planted alone, a seeding rate of 4 to 10 pounds

per acre (4.4 to 11.2 kg per ha) is recommended. Seeds

should be planted at depths of 0.25 to 0.50 inch (0.6 to

1.3 cm) (Jensen and others 2001). Seeds are large and

can be planted with most drills or broadcast seeders.

Figure 20—Common on many high-elevation sites,

mountain brome occurs in subalpine, mountain

meadow, aspen, conifer, and mountain big sagebrush communities (RMRS photo).
362 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Broadcast seed should be covered by light harrowing.

This species establishes better than many other grasses

when broadcast seeded. Mountain brome is frequently

seeded in mixtures with other native species. It is also

seeded with introduced species to provide ground

cover and quickly stabilize severely disturbed sites.

Consequently, the seeding rate for mountain brome

may be adjusted so that it will provide adequate

ground cover, but also permit establishment of other

seeded species (Sours 1983). Seedings should be protected from grazing until plants are well established

and seedheads are produced.

Uses and Management

Seedlings of mountain brome are vigorous and establish quickly, but plants may not produce seed for 2

to 3 years (Cronquist and others 1977). Stands of

mountain brome generally consist of scattered plants,

but they sometimes develop a dense sod. Mountain

brome can increase in density when growing with

other perennials. Plants, however, are short lived and

must be allowed to set seed every 2 or 3 years to permit

natural reseeding. If natural seeding does not occur,

the species can be replaced by longer lived species.

Hafenrichter and others (1949) reported that longevity and persistence are important characteristics to

consider when selecting plant material of this species.

Mountain brome is seeded as an early successional

species and establishes quickly on disturbed and weedy

sites at mid to upper elevations. It is seeded alone or in

mixtures on depleted livestock and game ranges, burns,

roadways, and mined lands in forested areas. The

species is effective for providing rapid, cold-tolerant

cover and erosion control in moderate to high elevation watersheds, as it grows rapidly and produces

deep, well-branched root systems (Stevens and others

1992). Mountain brome is also an excellent grass for

pasture or hay (Sours 1983).

Mountain brome tolerates a wide variety of soil

textures, but it most often occurs on medium-textured

to loamy soils in the pH 5.5 to 8.0 range (Sours 1983).

It is most productive on deep, rich, moist soils, but it

can grow on dry, shallow, and infertile soils on disturbed sites. The species is winter hardy, moderately

tolerant of shade and salinity, and weakly to moderately tolerant of drought and heat (Hassell and others

1983; Steele and Geier-Hayes 1993). It does not endure long periods of flooding or high water tables.

Mountain brome is palatable to livestock and big

game (Stubbendieck and others 1992). It produces

moderate amounts of high-quality herbage and receives considerable use in spring and early summer

prior to seed dispersal (Hassell and others 1983). Elk

use it in summer (Leege 1984; Stubbendieck and

others 1992). Seedheads are used by birds and small

mammals. The herbage becomes coarse and fibrous at

maturity. Regrowth in summer is moderate, providing

late fall forage. Fall regrowth is fair. Mountain brome

is tolerant of grazing, but excessive use weakens the

stands (Stubbendieck and others 1992).

Mountain brome is tolerant of wildfires (Brown and

DeByle 1989), recovering from seed and surviving

plants. Stands may be reduced initially, but they

recover to prefire levels after a few years. Success of

postfire seedings of this species has been variable.

Established seedings are useful for stabilizing soil and

providing cover and forage, but they may prevent

recovery of native species or coseeded natives (Steele

and Geier-Hayes 1993).

Varieties and Ecotypes

‘Bromar’ mountain brome was developed from native material collected near Pullman, WA (Alderson

and Sharp 1994). It is adapted to streambanks, semiwet

meadows, subalpine conifer forests, aspen, and upper

mountain brush communities in the Intermountain

West. Bromar has been widely used to stabilize logging, mining, roadway, and other disturbances. It does

not spread aggressively, and it is compatible with

other seeded or indigenous species. Livestock and big

game use its forage, especially prior to the production

of seedstalks. This variety exhibits good seedling vigor

and spring greenup. It is, however, susceptible to head

smut and fairly short lived (6 to 10 years). Bromar is

a good seed producer; stands reseed naturally if not

grazed excessively. Bromar should be fall seeded by

drilling or broadcasting

‘Garnet’ mountain brome germplasm, released in

2000 by the Upper Colorado Environmental Plant

Center, Meeker, CO (Englert and others 2002), originated from material collected near Garnet in Powell

County, MT (Alderson and Sharp 1994), at an elevation of 5,800 ft (1,770 m). Garnet has been evaluated

throughout the Intermountain West. It exhibits good

stand longevity and excellent head smut resistance,

two characteristics lacking in many populations of

this species. Garnet is recommended for seeding on

deep, fertile, mesic, medium-to fine-textured soils in

areas receiving at least 15 inches (380 mm) of annual

precipitation. It thrives in climates with cool, dry

summers and good winter precipitation. It does not

tolerate high water tables or flooding, but it does

exhibit good shade tolerance and fair fire tolerance,

and it is very winter hardy. Garnet is adapted to

mountain brush and mountain big sagebrush communities. It is also seeded in openings in aspen and

conifer forests and in subalpine areas. It is not aggressive and can be seeded in mixtures.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 363

Chapter 18 Grasses

Bromus inermis

Smooth Brome, Hungarian

Brome _________________________

Synonyms

Festuca inermis

Bromopsis inermis

Bromus pergans var. purparescens

Bromus pergans var. longispicata

Bromus pumpellianus

Bromus ciliatus var. coloradensis

Description

Smooth brome is a long-lived, strongly rhizomatous,

sodforming perennial with the few culms about 5.9 to

17.7 inches (1.5 to 4.5 dm) tall. Leaf sheaths are

smooth to silky haired. Blades are flat and broad,

about 0.16 to 0.51 inch (4 to 13 mm) wide, rarely

involute, and hairless or rarely silky haired on the

upper side. Leaf margins and nerves are rough and

hairy. Ligules are small, about 0.06 to 0.20 inch (1.5 to

5.0 mm) long, and irregularly serrate or reduced to a

marginal fringe of minute hairs. Panicles are 2.8 to 9.1

inches (7 to 23 cm) long, moderately open, oblong, and

erect. Panicle branches are ascending; the upper ones

are appressed. Spikelets are 0.6 to 1.3 inches (1.6 to 3.2

cm) long, 4 to 10 flowered, very narrow and round

when young, but becoming somewhat compressed with

age. Glumes are hairless and smooth. The first glume

is 0.18 to 0.31 inch (4.5 to 8.0 mm) long, narrowly lance

shaped, and one to three nerved. The second glume is

0.24 to 0.39 inch (6 to 10 mm) long, lance shaped, and

three to five nerved. Lemmas are 0.35 to 0.47 inches (9

to 12 mm) long, rounded on the back, smooth or rarely

minutely hairy, and greenish to purplish tinged. The

lemma apex is bidentate to slightly rounded. Awns,

when present, are up to 0.08 inch (2 mm) long. Smooth

brome flowers from June to August, and sometimes

into early September.

Smooth brome hybridizes readily with native B.

pumpellianus, a taxon that Elliott (1949) suggested

might be reduced to a subspecies of B. inermis due to

extensive introgression. Arnow (1987) stated that she

had never seen material that could be identified as

belonging to B. pumpellianus, which is distinguished

from B. inermis by its pubescent culm nodes and

lemmas.

Ecological Relationships and Distribution

Smooth brome is a European introduction that has

been widely seeded for hay and pasture in irrigated

and nonirrigated fields. In the Western United States

it has been planted on overgrazed rangelands, roadways, ski areas, mined lands, burned areas, unstable

watersheds, and other disturbances at mid to upper

elevations (fig. 21). It is now distributed from Canada

to Southern California and east to New Mexico and

much of the North-Central and Northeastern United

States (Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969).

Smooth brome is adapted to streambanks, valley

bottoms, and upland sites (Hardy BBT Limited

1989) in exposed to partially shaded areas that

receive 14 to 20 inches (356 to 508 mm) of annual

precipitation (Jensen and others 2001; Plummer

and others 1968) and sites that are subirrigated or

intermittently wet. It grows on a variety of welldrained, medium-textured clay loam to sandy loam

Figure 21—A strongly rhizomatous introduced

species, smooth brome receives wide use for watershed stabilization and postfire seedings on forested sites, but it does preclude recovery of native

species (RMRS photo).
364 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

soils of at least moderate depth; it is less productive

on fine or coarse soils and on shallow soils. Smooth

brome is cold tolerant; tolerant of poorly drained

soils, saline or acidic conditions, and drought; but it

does not tolerate high alkalinity (Coulman 1987;

Hafenrichter and others 1968; Wambolt 1976).

Smooth brome becomes dormant in winter, and

initiates growth in early spring (Smoliak and others

1981). Plants are productive, developing abundant

vegetative growth and large seedheads (Jensen and

others 2001). They become dormant during dry periods, but initiate regrowth with adequate precipitation

in fall.

Following wildfires, smooth brome regenerates by

sprouting from rhizomes and tilllering. Productivity is

increased by early- or late-season fires that remove

litter, particularly in sodbound stands (Higgens and

others 1989; Hughes 1985). Fires occurring in late

spring are generally more damaging (Masters and

Vogel 1989).

Plant Culture

Smooth brome is generally a cross-pollinating species (Knobloch 1944), but self-fertilization can occur

between flowers in different spikelets (McKone 1985).

Because of insect herbivory, seed set is lower for

smooth brome than for many other brome species

(McKone 1985; Nielson and Burks 1958). Seeds mature in early to midsummer. They are harvested by

combining and cleaned with an air screen machine.

There are 136,100 to 149,700 seeds per pound (300,000

to 330,000 per kg) (AOSA 1999). Smooth brome seed

has remained viable for periods of 22 months to more

than14 years in storage (Hafenrichter and others

1968; McAlister 1943; Rincker and Maguire 1979).

The AOSA (1999) rule for testing germination of smooth

brome stipulates an incubation temperature alternating between 68 and 86 ∞F (20 to 30 ∞C) (16 hrs/8 hrs).

A moist prechill at 41 to 50 ∞F (5 or 10 ∞C) for 5 days is

recommended for fresh or dormant seed.

Smooth brome should be planted in late fall or early

spring at depths of 0.25 to 0.75 inch (0.6 to 1.9 cm),

depending on soil texture, but seedlings do emerge

from greater depths (Plummer 1943). Seeds may be

drilled or broadcast and covered. The elongate seeds

sometimes plug drills (Jensen and others 2001). Seedlings can germinate at low temperatures under snow

(Bleak 1959) and develop rapidly as temperatures

increase in spring. Smooth brome requires fertile soil.

Fertilization or seeding with a nitrogen fixer may be

required to maintain stands when natural soil fertility

is inadequate (Jensen and others 2001). Seedlings are

not strong, but once established they grow rapidly

(Hardy BBT Limited 1989; Hassell and others 1983).

Grasshoppers sometimes damage or destroy developing stands. Mature plants spread vegetatively, while

spread from seed is moderate (Plummer 1977). Although stands may require 3 years to become fully

established, they are aggressive and may dominate

seedings or recovering native species. Because of its

ability to spread vegetatively, smooth brome plants

are long lived. Plantings have persisted for 60 years or

more (Plummer and others 1968). Established stands

of the species invade and dominate local disturbances

(Boggs 1984; Boggs and Weaver 1992; D’Antonio and

Vitousek 1992).

Uses and Management

Smooth brome is one of the most commonly seeded

exotic grasses for pastures and wildlands in North

America (Mitchell 1982, 1987b). Selected varieties can

be used on a wide range of irrigated and nonirrigated

sites with varying soil conditions.

A number of smooth brome varieties with differing

areas of adaptation have been developed to provide

erosion control on disturbed lands and nutritional

forage or hay when planted alone or in combination

with legumes. Smooth brome regrowth following cutting is slow. Sixteen to 18 inches (41 to 46 cm) of annual

precipitation are generally required to maximize forage production. Pastures and fields that become

sodbound require application of fertilizer and cultivation for renovation (Jensen and others 2001).

Smooth brome has been planted in a wide variety of

plant communities to provide palatable forage for

wildlife and livestock. Seeded stands, particularly

those on fertile soils (Currie and Smith 1970), are

palatable and tolerant of heavy use. The species has

been seeded in saltgrass, pinyon-juniper, quaking

aspen, and subalpine communities (McGinnies

1960a,b, 1975; Sampson and others 1951).

Wildlife use of smooth brome varies among animals

and season. Deer browse the species (Stubbendieck

and others 1992); elk take it primarily in winter

(Hobbs and others 1981). Ducks and upland birds use

it for nesting cover, while geese graze the new shoots.

Rodents eat the forage and seeds.

The rhizomatous habit and aggressive growth of

smooth brome make the species a valued erosion

control plant (Hardy BBT Limited 1989). It is used for

stabilization of degraded streambanks, mined lands,

logged areas, and other disturbances (Elliott and others 1987; Ward and others 1986). Loss of native vegetation, however, may occur because of the aggressiveness of this species (Smoliak and others 1981).

Smooth brome has been widely used to hold surface

soil and produce forage rapidly following wildfires

(Clary 1988; Crane and Habeck 1982). Because
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 365

Chapter 18 Grasses

recovering native grasses, forbs, and shrubs are often

as effective in holding the soil and controlling erosion

and because the aggressiveness of smooth brome may

preclude recovery of native species, its value for

postfire seedings has come under question (Clark

and McLean 1979; Crane and Habeck 1982).

Varieties and Ecotypes

Smooth brome varieties are generally divided into

three types: southern, intermediate, and northern.

Southern varieties include ‘Lincoln’, ‘Saratoga’,

‘Baylor’, ‘Beacon’, ‘Tempo’, ‘Signal’, ‘York’, ‘Radisson’,

‘Badger’, ‘Achenbach’, and ‘Southland’. Intermediate

forms include Tempo, ‘Magna’, and Signal. ‘Manchar’,

‘Carlton’, ‘Elsberry’, ‘Polar’, ‘Jubilee’, ‘Bravo’, and ‘Rebound’ are northern forms (Alderson and Sharp 1994).

Southern varieties are strongly rhizomatous

spreaders that are not compatible with natives.

Their long, wide, and somewhat coarse leaves are

primarily basal, but some occur along the culm. The

panicles are somewhat contracted with green to

light-red spikelets. Southern varieties generally

exhibit good seedling vigor. They require less moisture than do the northern types, and they can be

planted in areas receiving full sunlight or considerable shade. They green up early in spring and are

good forage producers.

Northern varieties are also aggressive spreaders

that are not compatible with natives. Their long leaves

are intermediate in width, soft, smooth, more or less

vertical, and occur well up the culms. Panicles are

open with red to purple spikelets. Northern varieties

exhibit good seedling vigor. They require more moisture than southern varieties, begin growth about 5 to

10 days later, and are generally less productive.

Lincoln and Manchar are two of the most commonly

seeded smooth brome varieties in the Intermountain

West. Lincoln, a southern variety, is derived from

material thought to be of Hungarian origin (Alderson

and Sharp 1994). It is seeded in aspen, mountain

brush, ponderosa pine, upper pinyon-juniper, and big

sagebrush communities that receive 13 or more inches

(330 mm) of annual precipitation. It is also well adapted

to semiwet meadows and streambanks. Lincoln establishes quickly from drilling or broadcasting and is a

very aggressive spreader that eventually dominates

mixed seedings.

Manchar, a northern variety from Manchuria, China

(Alderson and Sharp 1994), is adapted to openings in

aspen stands, conifer forests, mountain big sagebrush

communities, semiwet meadows, and riparian areas.

It is less well adapted to mountain brush communities. Manchar is aggressive and will dominate seeded

areas, especially those that are heavily grazed.

Bromus riparius

Meadow Brome _________________

Synonyms

Bromus erectus

Bromus biebersteinii

Bromus macounii

Description

Meadow brome, earlier described as Bromus erectus

(Hitchcock 1950), is now recognized as Bromus riparius

(Vogel and others 1996). Based on descriptions provided by Tsvelev (1984), Vogel and others (1996) concluded that most North American introductions and

cultivars of B. biebersteiniiare also B. riparius. Meadow

brome is an introduced, long-lived, cool-season, perennial bunchgrass. Culms are tufted, 2 to 3 ft (6 to 9 dm)

tall, erect, slender, and glabrous to silky haired. Leaf

blades are narrow and sparsely hairy. Ligules are 0.06

inch (1.5 mm) long. Panicles are narrow and 4.0 to 7.8

inches (10 to 20 cm) long with ascending to erect

branches. Spikelets are usually 5 to 10 flowered.

Glumes are tapered to a point, the first is about 0.24 to

0.31 inch (6 to 8 mm) long, and the second about 0.31

to 0.39 inch (8 to 10 mm) long. Lemmas are 0.39 to 0.47

inch (10 to 12 mm) long and smooth or evenly rough

hairy on the back. Awn length is 0.20 to 0.24 inch (5 to

6 mm) (Hitchcock 1950).

Meadow brome is somewhat similar in appearance

to smooth brome (fig. 22). The main difference is the

Figure 22—Meadow brome can be distinguished

from smooth brome by the pubescence on the

awns and leaves. Its panicles rise well above the

leaves (RMRS photo).
366 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

pubescence on the awns and leaves of meadow brome

(Knowles and others 1993). In addition, its rhizomes

are much shorter than those of smooth brome, and its

stems and seeds are more pubescent (Vogel and others

1996).

Ecological Relationships and Distribution

Meadow brome is a European introduction

(Weintraub 1953), and has been planted or has established in localities from Maine to New York, as well as

in the Pacific Northwest and the South (Hitchcock

1950). In recent years it has been seeded on range and

watershed sites in the West. Meadow brome has proven

adapted to mountain brush, ponderosa pine, aspen,

and subalpine communities in the Intermountain region. It is well-suited to acidic soils of the Idaho

Batholith, including disturbances where subsoils have

been exposed. This species is marginally adapted to

drier regions occupied by pinyon-juniper and big sagebrush communities. In the Intermountain region, the

plant has not persisted on sites receiving less than 12

to 14 inches (30 to 36 cm) of annual precipitation.

Plant Culture

Seeds of meadow brome are similar in size to those

of smooth brome and other brome species. Knowles

and others (1993) reported that compared to other

bromes, the seeds of meadow brome shatter more

easily and require more processing to remove the awns

and pubescence to assure that seeds can be seeded

uniformly. Meadow brome establishes quickly, and

dense, uniform stands generally develop. Plants grown

under cultivation produce reliable and abundant seed

crops in most years. Successful stands have established on range and wildland sites in the West by drill

seeding or broadcasting and harrowing to cover the

seed. When drill seeded alone, planting rates of 8 to 10

pounds per acre (8.9 to 11.2 kg per ha) are sufficient.

The rate should be decreased to 2 to 4 pounds per acre

(2.2 to 4.4 kg per ha) when seeded in mixtures with

other species. Rates should be increased slightly for

seedings on severely disturbed and eroding sites to

assure rapid development. Plants provide good ground

cover in the first and second seasons. Like mountain

brome, meadow brome is susceptible to head smut,

and seeds should be treated prior to planting to reduce

the chance of infestation.

Uses and Management

Meadow brome and smooth brome are the two species of brome cultivated for grazing and seed production in pastures in North America (Vogel and others

1996). Although there are native bromes in the West,

few have been developed for restoration plantings.

Instead, both smooth brome and meadow brome have

been used to provide effective ground cover and furnish palatable forage. Use of meadow brome in the

West began in 1966 with the release of ‘Regar’ for

planting as a forage and conservation species on rangelands and to provide watershed stabilization (Casler

and Carlson 1995; Knowles and others 1993).

Normally long lived, meadow brome is less winter

hardy and drought tolerant than smooth brome

(Knowles and others 1993). It has been widely and

successfully planted in irrigated pastures for grazing

and seed production. It is also well-suited to mountain

brush, aspen, conifer forests, and mountain park communities, and it is recommended for planting in intermediate and high-elevation woodlands (Hansen 1972).

In western Montana it has competed more successfully with cheatgrass than has crested wheatgrass. It

is less well adapted to more arid regions occupied by

big sagebrush and associated shrubs and bunchgrasses.

Meadow brome is not as productive as smooth brome,

but it is still regarded as an important forage species.

Spring greenup is rapid (Schwendiman and others

1946), and plants provide excellent early spring and

summer herbage for game and livestock. Regrowth is

more rapid for meadow brome than for smooth brome

(Knowles and others 1993); consequently, pasture and

rangeland stands supporting the species recover well

from early-season grazing. When seeded on wet and

semiwet sites in mountain communities this grass

also provides excellent summer and fall herbage, and

plants persist well. Herbage quality diminishes as

plants mature, but the foliage remains quite palatable

even though the leaves are quite pubescent (Hansen

1972). This species provides a valuable hay crop, even

on soils with low fertility (Hansen 1972). Rangeland

plantings have not diminished in productivity or become sodbound due to a decrease in available nitrogen,

as is commonly experienced with smooth brome.

Vegetative spread is not so aggressive for meadow

brome as for smooth brome. Plantings on high-elevation mountain herblands and in mountain brush communities have spread little from the original rows over

a 25-year period. In addition, established stands have

not seriously reduced the presence of associated native herbs. This grass is much less competitive with

native species than smooth brome, and could be planted

in areas where it is important to maintain the composition of native plants.

Varieties and Ecotypes

Regar is an early maturing bunchgrass that is

adapted to areas in upper pinyon-juniper, mountain

brush, and aspen communities that receive 14 inches

(356 mm) or more of annual precipitation. It was

developed from material collected at Zek, Turkey
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 367

Chapter 18 Grasses

(Alderson and Sharp 1994), and released by the Aberdeen Plant Materials Center in Aberdeen, ID (Foster

and others 1966). This variety is characterized by good

drought tolerance, excellent winter hardiness, and

moderate spread. Plants greenup early in spring.

They grow well in full sunlight and partial shade. They

are leafy and exhibit good regrowth, particularly in

midsummer. Cooper and others (1978) found that

yields were greater for Regar than for smooth brome

when seeded in grass-legume mixtures. Regar may be

planted in fall or early spring when seeds germinate

and establish readily. It does not spread as aggressively as smooth brome, and it is somewhat compatible

with other recovering natives. It is seeded in the

Intermountain West for forage and for stabilization of

disturbed sites.

Subsequent meadow brome cultivar releases, ‘Fleet’

and ‘Paddock’, were developed by Agriculture Canada.

Seed production of both cultivars generally exceeds

that of Regar by more than 65 percent (Vogel and

others 1996).

Dactylis glomerata

Orchardgrass___________________

Synonyms

Bromus glomeratus

Festuca glomerata

Description

Attempts have been made to place orchardgrass in

different generic complexes, but for the most part,

taxonomists in the United States and Europe agree

that it should remain in the monotypic genus Dactylis.

The species is a tufted perennial that grows 1 to 4 ft (3

to 12 dm) tall and produces short rhizomes. Leaf

sheaths are shorter than the internodes. Ligules are

0.16 to 0.31 inch (4 to 8 mm) long and oblong to ovate

with irregularly lacerated margins. Leaf blades are

0.16 to 0.39 inch (4 to 10 mm) wide, lax or loosely

arranged, and roughened on the surface. Panicles are

1.8 to 7.9 inches (4.5 to 20 cm) long, laterally compressed, and mostly interrupted. The primary panicle

branches, one to three at each node of the rachis or axis

of the panicle, measure about 1.2 to 4.7 inches (3 to 12

cm) long or longer, and are erect to widely spreading or

reflexed. The branches are naked below, but with one

to several dense clusters of subsessile to shortpedicillate spikelets borne near the branch tips and

arranged on one side of the axis. Spikelets are 0.20 to

0.35 inch (5 to 9 mm) long, two to five flowered, and

occasionally purple tinged. Glumes are subopposite,

lance shaped, often asymmetrical, and keeled to

rounded on the back. Their margins are straight, and

they are acute to awn tipped apically. The keel is

smooth to variously pubescent. The first glume is 0.20

to 0.25 inch (5 to 6.5 mm) long, keeled, and acute or

tapered to an awnlike tip that is about 0.08 inch (2

mm) long. The second glume is 0.12 to 0.24 inch (3 to

6 mm) long and membranous. Florets disarticulate

above the glumes, which remain attached to the spikelet. Flowering occurs from June through August (Arnow

1987, 1993; Cronquist and others 1977; Hitchcock

1950; and Hitchcock and others 1969).

Ecological Relationships and Distribution

A European introduction, orchardgrass is widely

distributed throughout North America. In the West it

is found in sagebrush, mountain brush, ponderosa

pine, aspen-fir, and occasionally in desert shrub communities. Orchardgrass was introduced as a hay and

pasture grass, and in the West, it is commonly grown

in irrigated areas or on sites receiving a minimum of

18 inches (457 mm) of annual rainfall. Although it

generally occurs as a seeded species, orchardgrass

volunteers on moist nonirrigated sites (Vallentine

1985). It requires well-drained, medium-textured, deep

to moderately deep soils that are moderately acid to

moderately alkaline. It will grow on shallow, gravelly,

or stony soils (Vallentine 1961). Orchardgrass initiates growth in early spring, and it is highly productive during the cool season. Due to its deep rooting

habit it is also capable of considerable summer growth

under favorable conditions (USDA Natural Resources

Conservation Service 2000). Orchardgrass produces

palatable, nutritious forage, and it tolerates heavy

grazing. It is reported to increase or remain stable

after burning (Cocking and others 1979).

Plant Culture

Orchardgrass seed matures evenly and is ready for

harvest in mid-August (USDA Natural Resources

Conservation Service 2000). There are 480,000 to

650,000 seeds per pound (1,058,200 to 1,432,990 per

kg). Orchardgrass can be spring seeded, but later

dates can be successful with adequate irrigation. Fall

seedings may be more appropriate on sites that are

likely to dry early in the season. A clean, firm, weedfree seedbed is recommended. Seed should be planted

about 0.5 inch (1.3 cm) deep. Seed increase fields

should be planted with the rows at 28- to 40-inch

(71- to 102-cm) spacings. Orchardgrass establishes

well in mixtures with other competitive introductions

where adequate precipitation, at least 12 inches

(305 mm), is received annually (Hardy BBT Limited

1989).
368 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Uses and Management

Orchardgrass is used in seed mixes with other grasses

and forbs for revegetating overgrazed or otherwise

disturbed lands (Hardison 1980) (fig. 23). Success of

these seedings depends on the selection of appropriate

cultivars. Orchardgrass has been planted in logged

over or burned areas, and it provides cover and forage

for livestock and wildlife (Clary 1975a). It has also been

used for mine reclamation, ski slope stabilization, and

suppression of annual weeds (Behan 1983; Hardy 1989).

Orchardgrass is highly shade tolerant and has been

seeded under aspen and open conifer stands.

Orchardgrass is recommended for moderate earlyseason grazing (Humphrey 1960). It provides excellent, highly digestible forage (Vallentine 1961). Rotational grazing practices are best for irrigated pastures.

Orchardgrass is grown with alfalfa for hay crops

because of its ability to recover from grazing or clipping. It responds well to nitrogen fertilizers and to

being integrated with legumes (Shaw and Cooper

1973). In areas with dry, cold winters, a hardy variety

of orchardgrass should be used to avoid losses from

winterkill (Vallentine 1961).

Vallentine (1961) reports that all classes of livestock

readily use orchardgrass, and deer, elk, and bighorn

sheep graze it. The basal rosette of the plant is green

through fall and into the winter and provides a good

source of forage for wildlife (Dalke 1941). Wildlife use

of seedings on burns and other disturbances increases

when orchardgrass is seeded (Evanko 1953; Leege and

Godbolt 1985; Lowe and others 1978). Seedings also

provide habitat for rodents (Reynolds and Johnson

1964) and nesting and brood rearing habitat for upland

birds as well as escape and winter cover. It is used by

rabbits and Canada geese (Crawford and others 1969).

Although orchardgrass is widely seeded following

wildfires or other disturbances, it interferes with

natural regeneration of conifer and native understory

species and with establishment of planted conifers

(Anderson and Brooks 1975; Elliott and White 1987;

Lyon 1984; McLean and Clark 1980; Seidel and others

1990). It should not be included in seeding mixes with

native species.

Varieties and Ecotypes

Varieties that have been developed and for which

seed is available include: ‘Able’, ‘Akaroa’, ‘Amba’, ‘Ambassador’, ‘Arctic’, ‘Benchmark’, ‘Cambria’, ‘Chinook’,

‘Comet’, ‘Dactus’, ‘Dart’, ‘Dawn’, ‘Hallmark’, ‘Haymate’,

‘Juno’, ‘Justus’, ‘Kay’, ‘Latar’, ‘Napier’, ‘Pomar’,

‘Potomac’, ‘Prairial’, ‘Rancho’, ‘Rapido’, ‘Sumas’, and

‘Summer Green’ (Alderson and Sharp 1994). The only

two rangeland varieties are ‘Berber’ and ‘Paiute’.

Berber orchardgrass was developed for California

rangelands from material obtained from South Australia (Alderson and Sharp 1994). It has not been

seeded in the Intermountain West.

Paiute orchardgrass was introduced from Ankara,

Turkey (Alderson and Sharp 1994). It is a droughttolerant accession that is adapted to basin big sagebrush, pinyon-juniper, mountain brush, mountain big

sagebrush, and ponderosa pine communities receiving 14 or more inches (355 mm) of annual precipitation. Paiute is characterized by an abundance of thick,

succulent basal leaves that tend to remain green

throughout the winter. It begins growth in spring as

the snow melts. Plants are palatable and sought out by

big game and livestock. Seed should be fall planted on

a firm seedbed and covered. Germination is slow, but

once established, Paiute forms strong bunches that

are very persistent. Plants are shade tolerant and

compatible with some seeded and indigenous species.

Latar, an introduction from Russia, and especially

Potomac, developed from collections made in old

pastures on the east-central coast of the United States

Figure 23—Orchardgrass seedings provide forage,

stabilize disturbances, and suppress the spread of

invasive species, but it interferes with the recovery

and establishment of natives (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 369

Chapter 18 Grasses

(Alderson and Sharp 1994), are used extensively in the

Intermountain West for seeding streambanks and

disturbed sites in mountain brush, aspen, ponderosa

pine, and mountain big sagebrush communities. These

two varieties produce an abundance of succulent leaves

just after snowmelt in spring and remain green until

late fall. Both varieties are long lived and exhibit

considerable shade and grazing tolerance. They should

generally be seeded in fall, and they can be planted

with other rapidly developing species. Germination is

generally slow.

Deschampsia caespitosa

Tufted Hairgrass ________________

Synonyms

Aira cespitosa

Deschampsia pungens

Deschampsia holciformis

Deschampsia beringenis

Description

Tufted hairgrass is a caespitose, mat forming perennial with culms 0.7 to 4.3 ft (2 to 13 dm) tall, and

smooth, hairless leaf sheaths. Leaf blades are firm,

flat, or folded, 0.04 to 0.20 inch (1 to 5 mm) wide, and

often roughened ventrally. Ligules are 0.12 to 0.37

inch (3 to 9.5 mm) long, lacerate along the margins,

and gradually tapering to a sharp point. Panicles are

1.6 to 11.8 inches (4 to 30 cm) long, loose, often

nodding, and open to narrow, with slender, hairlike

branches that are fine hairy. Spikelets are two or three

flowered and usually shiny and purplish. Glumes are

lance shaped, acute, and smooth or roughened. The

first glume is single nerved and 0.08 to 0.20 inch (2 to

5 mm) long. The second glume is 0.10 to 0.20 inch

(2.5 to 5.2 mm) long and one to three nerved. Lemmas

are 0.10 to 0.18 inch (2.5 to 4.5 mm) long, often

purplish basally, and five nerved with the four lateral

nerves ending in apical teeth. The awns are 0.08 to

2.5 inches (2 to 6.5 cm) long and attach below midlength on the lemma. The callus is densely hairy.

Paleas more or less equal the lemmas. Plants flower

from late July to September (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969; USDA Forest Service 1937).

Tufted hairgrass has been described as 30 or more

different races, variants, or species. Recent western

floras have included these in the highly variable

species D. caespitosa (see Cronquist and others 1977)

(fig. 24).

Ecological Relationships and Distribution

A circumboreal species, tufted hairgrass is distributed in the Western Hemisphere from Alaska to Newfoundland, south through the Western and Eastern

States and into Northern Mexico. In the more southerly portions of its range it grows at higher elevations.

Its distribution in the Great Plains is limited (Hitchcock

1950; Larson 1993; Stubbendieck and others 1992).

Tufted hairgrass occurs in coastal prairies, bogs,

marshes, wet to mesic meadows, forest openings, dry

slopes, aspen stands, high-elevation valley bottoms,

and open areas above timberline, particularly on sites

where snow accumulates (Gould and Shaw 1983;

Johnson and Billings 1962; Ward 1969). It is also

found along ditches, streams, and lakeshores.

Figure 24—Tufted hairgrass exhibits considerable

ecotypic variability; individual populations are adapted

to a wide variety of soil types (Hitchcock 1950).
370 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Tufted hairgrass is a long-lived, cool-season species

that exhibits considerable ecotypic variation (Ward

1969). It occurs at elevations from sea level to over

14,000 ft (4,270 m) (Brown and others 1988) that

receive 14 to 24 inches (356 to 609 mm) of annual

precipitation (USDA Natural Resources Conservation

Services 2002). It is more common on sites that receive

at least 20 inches (508 mm) of annual precipitation

(Brown and others 1988). Tufted hairgrass is adapted

to fine- to medium-textured neutral to somewhat acidic

soils that are of moderate depth and fertility. Some

populations occur on coarser sandy or gravelly soils

(Gates 1962; Johnson 1962; Lewis 1970). Plants are

tolerant of anaerobic conditions (USDA Natural Resources Conservation Services 2002). They are not

drought tolerant, but they exhibit good winter hardiness (USDA Natural Resources Conservation Service

2002). Individual populations tolerant of salinity, alkalinity, or specific heavy metals have been identified

(Frenkel and Morlan 1991; Hardy BBT Limited 1989;

Rabe and others 1999).

Tufted hairgrass occurs in a large number of late

successional or climax communities (Walsh 1995),

growing mixed with other species or as nearly pure

stands on open sites, particularly at higher elevations.

It rarely occurs as an understory species in shaded

areas (Chambers 1987). Tufted hairgrass is also a

common species on disturbed areas within its range,

appearing in early- to midsuccessional communities

(Brown and others 1988).

Plants begin growth in early spring when temperatures are near freezing. In moist areas the leaves

remain green throughout the growing season (Bell

1974; Lewis 1970; Vallentine 1961). New leaf growth

is initiated prior to the onset of dormancy, and immature leaves are protected over winter by dry leaves of

the previous season.

Tufted hairgrass is fairly fire tolerant, as the leaves

are generally green during the growing season and

protect the root crown (Great Plains Flora Association

1986). Stands recover through resprouting from the

root crown or through establishment of new seedlings

from the soil seed bank (Chambers 1993; Gehring and

Linhart 1992). Severe fires can kill the plants

(DeBeneditti and Parsons 1979, 1984).

Plant Culture

Tufted hairgrass is self-sterile. Fruits ripen in summer. Wildland stands may be hand harvested in summer, but early shattering can reduce seed yields.

Combining or swathing followed by combining are

options for harvesting cultivated seed fields and extensive wildland stands on level terrain. Harvested

seed is cleaned using air screen machines and a

debearder or huller-scarifier. There are about 2,500,000

seeds per pound (5,500,000 seeds per kg). Fill and

viability of wildland collections are highly variable

(Chambers 1989). Germination requirements may vary

among seed lots. Archibald and Feigner (1995) reported that seeds do not require moist prechilling for

germination. Other investigators, however, found that

exposure to light or transfer from a warm (68/50 ∞F) (20/

10 ∞C) (16 hrs/8 hrs) environment to a cooler environment (59/41 ∞F) (15/5 ∞C) (16/8 hrs) after 6 weeks of

incubation improved germination of their seed lots

(Chambers and others 1987; Guerrant and Raven

1995; Sayers and Ward 1966).

Seed fields are planted at a rate of 1 pound per acre

(1.1 kg per ha). Irrigated fields may be planted in

spring or fall. Seeds are very small and should be

drilled no more than 0.25 inch (0.6 cm) deep. Weeds

must be controlled, as the early growth rate of tufted

hairgrass seedlings is only moderate. Mature stands

are quite competitive. Seed yields average 40 pounds

per acre (44.8 kg per ha) (Smith and Smith 1997).

Wildland seedings are conducted in late fall (Brown

and others 1988; Chambers and others 1987) or in

spring at higher elevations. Seeds can be drilled or

broadcast and covered at 0.8 to 1.6 pounds PLS per

acre (0.9 to 1.8 kg per ha) or greater on highly disturbed sites (Pawnee Buttes Seed, Inc. 2002).

Plugs, sod, and container stock have also been used

to establish tufted hairgrass on disturbances and

provide soil stabilization more rapidly. Applications of

fertilizers and soil amendments aid in hastening establishment on infertile soils (Brown and Chambers

1989, 1990; Brown and others 1988).

Tufted hairgrass is a valuable revegetation species,

particularly for disturbances at high elevations

(Vaartnou 1988). Because of the considerable ecotypic

variability, local seed sources or sources known to be

adapted to the planting site should be used. This is

particularly true on sites with unusual soil characteristics. Populations have been identified that are tolerant of heavy metals such as copper, lead, and manganese; other populations will grow on peat, while still

others have been found colonizing coal mine spoils

(Hardy BBT Limited 1989; Russell 1985; Vitt and

others 1990). Some populations are adapted to relatively dry sites (Arnow 1987).

Uses and Management

Tufted hairgrass provides an abundant source of

forage from spring through summer. It provides goodquality summer range and is one of the most valuable

forage species at high elevations. Tufted hairgrass

meadows are sometimes cut for hay. Late summer and

fall regrowth are moderate.

Although fairly tolerant of close grazing, the species

does decrease with excessive grazing and associated

trampling. It is considered an indicator species for

grazing pressure in many high-elevation communities.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 371

Chapter 18 Grasses

Stands should be allowed to set seed periodically to

perpetuate the community. Management activities that

alter the water table can also impact tufted hairgrass.

Increased shading with development of woody species

in later successional communities reduces tufted

hairgrass populations.

Tufted hairgrass is used to seed mined lands, dredge

spoils, coal mine overburden, burned forest lands,

degraded livestock and game ranges, ski areas, and

other disturbances at high elevations (McVay and

others 1980; Pawnee Buttes Seed, Inc. 2002). Seed

sources adapted to local soil conditions and with appropriate cold tolerance should be used for these

projects.

Varieties and Ecotypes

‘Norcoast’ was selected from collections made in

Cook Inlet, Alaska (Alderson and Sharp 1994), and

released for revegetation of disturbances and to provide forage. It has been used on adapted sites from

Alaska to California, but it has not seen use in the

Intermountain West.

‘Nortran’ was developed from materials collected in

Alaska and Iceland (Alderson and Sharp 1994; Mitchell

1988). It is used primarily for the revegetation of

disturbed areas and for pasture seedings. It is hardy in

alpine and boreal regions south of the Arctic. It has not

been seeded in the Intermountain West.

‘Peru Creek’ was developed from mass collections

made in subalpine meadows along Peru Creek, near

Dillon, CO (Englert and others 2002), at an elevation

of about 10,200 ft (3,120 m). It has been tested and

seeded in the Intermountain West for reclamation of

mined lands and other disturbed sites with acidic soils

and for restoration of wet meadows at high elevations.

Distichlis spicata

Inland Saltgrass ________________

Synonyms

Uniola spicata

Uniola stricta

Distichlis maritima var. laxa

Distichlis dentata

Briza spicata

Description

Inland saltgrass is a native, dioecious, warm-season, strongly rhizomatous perennial halophyte (Ungar

1974) that is widely distributed across the Western

United States and Canada (fig. 25) (Hitchcock 1950).

It is a relatively short grass that grows to 16 inches

(40 cm) in height. Inland saltgrass produces vigorous

scaly rhizomes at shallow depths, generally in the

upper 8 inches (20 cm) of the soil profile. The culms

sometimes bear long, soft, straight hairs that are

about 0.02 to 0.12 inch (0.5 to 3 mm) long at the collar

and upper margins of the sheath. Leaf blades on the

middle or upper portions of the culm are two ranked,

loosely involute and 0.08 to 0.16 inch (2 to 4 mm) wide,

with the upper surface sometimes roughened or long

hairy. Ligules are 0.01 to 0.02 inch (0.2 to 0.6 mm) long

with a fringe of cilia along the margins. Panicles are

1.2 to 4.0 inches (3 to 10 cm) long with a few somewhat

congested and strongly compressed spikelets. The

male panicle often extends above the leaves, while the

female panicle is normally enclosed within them. Staminate spikelets are 6 to 14 flowered and yellowish, while

the pistillate spikelets are 3 to 10 flowered. Glumes

are lance shaped, hairless, and five to seven nerved.

Lemmas are firmer than the glumes and seven to nine

nerved. Glumes and lemmas of pistillate flowers are

larger than those of the staminate flowers. The paleas

of the pistillate florets are slightly shorter than the

lemma, and their winged keels have irregular, shallowly serrate, and minutely ciliolate margins (Arnow

1987; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969).

Figure 25—A strongly rhizomatous warm-season perennial, inland saltgrass can grow on sites

with shallow water tables and on

highly saline soils (photo courtesy of Marshall Haferkamp,

USDA ARS, Miles City, MT).
372 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Saltgrass is best adapted to medium- and finetextured soils (Shaw and Cooper 1973), but it can also

be found in sandy and gravelly soils. It often occurs in

conjunction with a shallow water table and on sites

that are periodically flooded. On highly saline soils,

plants may be dwarfed, and the culms are generally

reclining with the tips ascending, or they may be

decumbent at the base and somewhat stoloniferous.

Saltgrass has a number of physiological adaptations

that allow it to occupy saline soils. Vesicular-abuscular

mycorrhizal fungi have been observed on inland

saltgrass roots, and are thought to enhance salt tolerance (Ho 1987). Plants contain salt glands that are

active in the extrusion of salt (Hansen and others

1976). The lacunae tissue of the roots is apparently

continuous with the rhizome and leaf sheath. This

allows for gas exchange under partial inundation and

in heavy soils (Hansen and others 1976).

Ecological Relationships and Distribution

Inland saltgrass is distributed from the interior of

British Columbia to Saskatchewan and southward to

California, Texas, and Mexico. It also ranges eastward

to Minnesota. Inland saltgrass is dominant or codominant in some communities and may be found in varying densities in a wide range of communities. Pure to

nearly pure stands occur on sites where soils are saline

and fine textured, and where the soil is wet at least

part of the year (Shantz and Piemeisel 1940; Ungar

1974). In the Great Basin saltgrass is a dominant

species in lower elevation desert shrub communities.

It spreads rhizomatously and forms a sod under black

greasewood and shadscale. Black greasewood and

saltgrass occurring together indicate the presence of

soil with a high moisture content. Where soil salinity

becomes greater than 1 percent, black greasewood

gives way to pure stands of saltgrass (Brotherson

1987).

Saltgrass does especially well on fine-textured soils

in areas with high water tables that receive seasonal

runoff. Salt flats, desert playas, valley bottoms, intermittent ponds, saline meadows, and borders of springs,

streams, and lakes commonly support this species.

Inland saltgrass forms a dense sod only on salt soils

with fine texture and good soil moisture throughout

most the year. It may also occur as a pioneering

species. The spreading rhizomes allow saltgrass to

colonize salt flats, playas, lakeshores, shale cliffs, and

other barren saline sites (Cronquist and others 1977;

Unger 1966) not tolerated by other grasses. Its pointed

rhizomes are well adapted to heavy clays and shales

(Hansen and others 1976).

Saltgrass has moderate to high drought tolerance

and can remain dormant for long periods of time

(Shantz and Piemeisel 1924). It exhibits exceptional

tolerance of flooding (Platts and others 1987). Although flooded plants may show a loss of vigor, they

generally recover within 30 days after the water has

receded. Saltgrass tolerates burning, as the rhizomes

are generally insulated from the heat of most fires.

Plant Culture

Inland saltgrass reproduction is mainly through

vegetative spread from a well-developed system of

deep underground rhizomes. It is well adapted to

spreading into a wide variety of soils. The species is a

poor seed producer. Seed production is generally restricted to better sites that support dense healthy

stands (Cluff and Roundy 1988). Fill and viability of

saltgrass seed is often quite high. Seed can remain

dormant for at least 4 years (Shaw and Cooper 1973).

This allows soil seed banking to occur and permits

plants to establish in large numbers when favorable

conditions exist for germination. Saltgrass seed requires relatively high temperatures, low salinities,

and moist soil to germinate (Roundy 1987). Saltgrass

has been successfully seeded with a Brillion seeder

and with broadcast seeding. Seed must be planted

about 0.5 inch (1.3 cm) deep.

Saltgrass has been established by planting rhizomes. Because rhizomes are sensitive to desiccation,

they should be planted only in wet, saline areas, unless

irrigation is being used during establishment. Planting rhizomes at depths of 1 to 2 inches (2.5 to 5 cm) has

proven successful (Delzell 1972).

Uses and Management

Saltgrass is not greatly preferred by livestock and

wildlife. Cattle generally avoid its harsh foliage. Because it usually remains green all summer long, it is

most often grazed after other grasses have been consumed or have dried out (Humphrey 1970; Shaw and

Cooper 1973). Saltgrass is low in nutritive value. For

cattle, a pure saltgrass diet should be avoided because

severe rumen compaction can occur (Stubbendeick

and others 1985).

Small mammals and birds use saltgrass for cover,

nesting, and feeding. It often forms a dense sod that

provides good escape and security cover. Saltgrass

stands can provide nesting cover for pheasants and

many species of ducks and shore birds. Seeds of

saltgrass are eaten by waterfowl and small mammals

(Ohlenbusch and others 1983). Saltgrass sod around

lakes often indicates alkaline conditions that are unfavorable for fish (Hansen and others 1988a,b).

Saltgrass is highly resistant to grazing and trampling, and it provides a resistant sod in areas where

cattle concentrate, such as watering sites, corrals, and

along trails. Attempts have been made to rehabilitate

saltgrass meadows by killing the saltgrass and planting
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 373

Chapter 18 Grasses

more palatable, salt-tolerant species (Bowman and

others 1985; Ludwig and McGinnies 1978). Plowing,

disking, and other mechanical treatments have proven

ineffective because inland saltgrass reproduces profusely from rhizomes (McGinnies 1974, 1975).

Plant Culture

Reestablishment of saltgrass is often required to

restore depleted wetlands. Seed, however, is not widely

available. In addition, removal of weedy competition

and preparation of a seedbed can be difficult on inland

saltgrass sites. Seeding on sites with some surface

litter is advised because the soils often crust, reducing

seedling emergence. Inland saltgrass develops slowly,

requiring 2 to 3 years to become fully established.

Seeding pure stands is normally recommended.

Varieties and Ecotypes

There are no releases of inland saltgrass.

Elymus canadensis

Canada Wildrye _________________

Synonyms

Hordeum patulum

Clinelymus canadensis

Sitanion brodiei

Elymus glaucifolius

Elymus robustus

Elymus brachystachys

Description

Canada wildrye is a highly variable, perennial, coolseason bunchgrass. Plants are coarse and open with

erect to decumbent, tufted, hollow culms 2.6 to 6.5 ft (8

to 20 dm) tall. Rhizomes are short and usually absent.

Leaves are mostly cauline. The blades are flat, 0.12 to

0.59 inch (3 to 15 mm) wide, and roughened above.

Auricles are well developed, clasping, and about 0.04

to 0.08 inch (1 to 2 mm) long. Ligules are 0.01 to 0.08

inch (0.2 to 2 mm) long, truncate, and fringed with

minutely ciliate hairs. Spikes are thick, bristly, and

usually nodding or drooping (Asay and Jensen 1996b).

Spikelets are 0.47 to 0.59 inch (12 to 15 mm) long

excluding the awns, two to four flowered, and two to

four per node. Flowers are widely separated, leaving

the axis of the spikelet visible. Glumes are subequal,

narrowly lanceolate, about 0.39 to 0.98 inch (10 to

25 mm) long, three to five nerved, broadest above the

base, and tapering to a long awn. The nerves on the

glumes are roughened to somewhat ciliate. Lemmas

are about 0.33 to 0.55 inch (8.5 to 14 mm) long,

strongly roughened, five to seven nerved, and gradually

tapering into a long, rough, flexible, sharp awn that

measures 0.63 to 1.18 inches (16 to 30 mm) in length.

Flowering occurs from late June through August

(Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969).

E. canadensis has been found to hybridize with

Agropyron trachycaulum and E. interruptus (Arnow

1987; Great Plains Flora Association 1986). It has

formed synthetic hybrids with Sitanion hystrix and

some Agropyron species (Dewey 1967).

Ecological Relationships and Distribution

Canada wildrye is distributed from Alaska south to

California, and eastward across much of Canada and

the United States except for the extreme Southeastern

United States (Great Plains Flora Association 1986).

It generally occurs in areas receiving 20 to 45 inches

(508 to 1143 mm) of annual precipitation. The species

occurs widely on tall-grass and mixed-grass prairies of

the Great Plains and Midwestern States, particularly

in sandy areas (Barker and others 1985). It also grows

in riparian areas and wetlands, occurring with eastern cottonwood, green ash, redosier dogwood, and

other riparian species in the Missouri River system of

the Great Plains (Boggs 1984). Canada wildrye is

tolerant of shade and occurs in oak savannas of Minnesota (Tolstead 1942), ponderosa pine communities

in the Southwest (Lavin 1953), and mixed evergreen

and hardwood forests of the Coast Range in California

(Hickman 1993). It is less common in the Intermountain West, usually growing in low, moist valleys or

along streams, irrigation ditches, and roadsides.

Canada wildrye is adapted to soils of low to moderate fertility. It can be found on soils ranging from

gravelly to clayey and from neutral to slightly alkaline

(Great Plains Flora Association 1986). Pure stands

can be found on wet to moist, alkaline, or sometimes

saline meadows (Arnow 1987).

Canada wildrye develops rapidly and is rather short

lived. It is a good seed producer and depends upon seed

for spread and persistence (Weaver and Zink 1946).

The species is considered early to midseral and often

occurs on riparian and upland disturbances. Canada

wildrye begins growth in early spring, flowers in

midsummer, and matures seed in late summer or

early fall. Plants may enter dormancy in summer and

resume growth in fall if moisture is received.

Fire may top-kill Canada wildrye. Its buds are

located beneath the soil surface where they are somewhat protected from burning. The coarse leaves and

stems are not highly flammable. Early spring fires are

most harmful, while the best postburn response follows summer fires. Regeneration following fire occurs

through recruitment from the seed bank (Howe 1994;

Mowat 1990; Robocker and Miller 1955; Wright 1971;

Wright and Bailey 1982).
374 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Plant Culture

Canada wildrye may outcross or self-fertilize (Gabel

1984). It produces numerous reproductive stems and

is generally a good seed producer. Seed maturation

occurs from July in the southern portion of the species

range to August in the northern portion (Smith and

Smith 1997; Wheeler and Hill 1957). Seed production

is generally good, and shatter during harvest is generally not problematic. However, threshing and cleaning

are complicated by the presence of the long awns

(Hafenrichter and others 1949). Swathing or windrowing during the hard-dough stage and combining

are the recommended methods of harvest (Smith and

Smith 1997). Seed is cleaned using a using a debearder,

a four-screen fanning mill, and a two-disc indent

cleaner (Smith and Smith 1997). Seeds average 115,000

seeds per pound (254,000 per kg). Seed yields from

nonirrigated fields range from 150 to 300 pounds per

acre (168 to 336 kg per ha), while yields from irrigated

fields range from 400 to 1,100 pounds per acre (448 to

1,232 kg per ha) (Smith and Smith 1997).

Appropriate seed sources for proposed planting sites

should be selected due to the considerable ecotypic

variation within the species. Seed should be planted in

late fall or early spring at a depth of about 1 inch

(2.5 cm) by broadcasting and covering the seed or by

drilling. Seeds germinate rapidly. Dry storage or a

2-month moist prechill improve germination (Blake

1935; Greene and Curtis 1950). Seedlings are vigorous

and stands develop rapidly. Peak production occurs in

2 or 3 years (Wasser 1982). Stand life for seed fields is

7 or 8 years (Smith and Smith 1997). In native stands

Canada wildrye may eventually be replaced by longer

lived species. It is not competitive with aggressive

exotic grasses. Canada wildrye is susceptible to infestation by ergot, which may cause illness or death if

consumed by livestock (Vallentine 1961).

Uses and Management

Canada wildrye is used by wildlife and livestock.

Forage value and palatability are greatest in early

spring, but decrease as plants mature (Hoover and

others 1948; Morris and others 1950; Stubbendieck

and others 1986). Stands of Canada wildrye are harvested for hay in the boot stage, just as the seedheads

emerge (Wheeler and Hill 1957). Plants tend to be

avoided after this stage. Stands decrease with heavy

grazing because the growing points are damaged

(Ehrenreich and Aikman 1963).

Canada wildrye seedings provide cover, litter, and

organic matter; control erosion; and provide habitat

for birds and other wildlife (Barker and others 1985;

Noyd and others 1995). It is tolerant of heavy metals,

and can be used to revegetate abandoned mine tailings

(Chambers and Sidle 1991; Hardell and Morrison

1983). It may also be used alone or in mixtures to seed

disturbances in sand dunes, riparian areas or wetlands, or native prairie communities. Canada wildrye

provides good cover for birds and many other small

animals. Its foliage tends to be rather coarse and not

highly palatable.

Varieties and Ecotypes

Mandan was released in 1946 by the Agricultural

Research Station Northern Great Plains Research

Center in Mandan, ND, the Natural Resources Conservation Service and the North Dakota Agricultural

Experiment Station. It originated from material collected near Mandan, ND, and was selected for its rapid

establishment, leafiness, productivity in young stands

of mixed species, and soil stabilizing capabilities

(Alderson and Sharp 1994; Baker and others 1985). It

is shorter and more persistent than many Canada

wildrye populations. Mandan currently receives little

use, and seed is generally not available.

Elymus cinereus

Great Basin Wildrye _____________

Synonyms

Leymus cinereus

Elymus condensatus var. pubens

Elymus piperi

Description

Great Basin wildrye is a coarse, robust, densely

tufted perennial bunchgrass that often forms clumps

up to 3.3 ft (1 m) across. It is the largest cool-season

bunchgrass native to the Western United States

(fig. 26) (Abbot and others 1991; Anderson and others

1995; Cash and others 1998). Plants sometimes produce short, stout rhizomes. Culms are usually more

than 2.3 ft (7 dm) tall, often growing to heights of 6.6

ft (2 m). They are hairless, or more often minutely

hairy, especially at the nodes. Leaf sheaths are hairless to soft hairy with appressed or spreading hairs.

Auricles are well developed to lacking. Ligules are

long and membranous or translucent, measuring about

0.08 to 0.28 inch (2 to 7 mm) in length. Leaf blades are

flat, or nearly so, 0.17 to 0.59 inch (4.5 to 15 mm) wide,

firm, and sometimes strongly nerved. Spikes are 2.8 to

9.0 inches (7 to 23 cm) long, 2.8 to 4.7 inches (7 to 12 cm)

wide, stiff, erect, and sometimes compound and branching. Spikelets are 0.35 to 0.79 inch (9 to 20 mm) long

and two to five flowered. There are usually three to six

per node, but sometimes only two. Glumes are subequal,

0.28 to 0.63 inch (7 to 16 mm) long, narrow, somewhat

awl shaped, tapered, and often as long as the spikelet.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 375

Chapter 18 Grasses

Lemmas are 0.28 to 0.47 inch (7 to 12 mm) long;

pubescent with very short, stiff hairs or rarely hairless; and usually nerveless below and five to seven

nerved above. The awn is short, usually less than 0.20

inch (5 mm) in length, or absent. Flowering occurs

from June to early August (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969). Great Basin wildrye plants in the

Western United States are generally tetraploid with

2n = 4x = 28, while those in Canada are octoploid with

2n = 8x = 56 (Asay and Jensen 1990b). Great Basin

wildrye forms natural fertile hybrids with beardless

wildrye and bottlebrush squirreltail in areas where

the species overlap (Arnow 1987; Asay and Jensen

1996b).

Ecological Relationships and Distribution

Great Basin wildrye occurs throughout the Western

United States and Canada and reportedly as far east

as Minnesota (Arnow 1987; Asay and Jensen 1996b).

In the Intermountain West it occurs in a wide range of

community types from salt desert shrublands to ponderosa pine communities (Arnow 1987). In this area it

commonly occurs with big sagebrush, rabbitbrush,

junipers, and wheatgrasses. In the Great Plains it is

associated with grama grass, buffalo grass, and mixedgrass prairie communities (Shiflet 1994). Great Basin

wildrye may occur scattered among other species or it

may form pure stands or stringers along riparian

areas, dry washes, roadsides, and on floodplains or in

other areas that receive runoff water or subirrigation,

or that have high water tables (Jankovsky-Jones and

others 1999; Wasser 1982). It also grows on upland

prairies, hillslopes, and mountainous sites. Great Basin

wildrye invades disturbed sites, and is an important

colonizer of steep, eroded slopes and gullies where it is

adapted.

Great Basin wildrye exhibits considerable ecotypic

variation with populations growing at elevations ranging from 1,970 to 9,840 ft (600 to 3,000 m) on sites

receiving 8 to 20 inches (200 to 500 mm) of annual

precipitation. Some populations grow on sites receiving as little as 5 inches (130 mm) of annual precipitation (Cash and others 1998; Davenport Seed Corporation 1997; Hickman 1993). Great Basin wildrye

commonly occurs on deep, silty to clayey soils, but it

also occurs on soils that are sandy or gravelly or that

have claypan layers or high potassium concentrations.

Various ecotypes are moderately tolerant of acid, alkaline, or saline conditions (Asay 1987; Wasser 1982).

Great Basin wildrye plants begin growth in early

spring (Hafenrichter and others 1968). Flowering

occurs in early to midspring, and seeds ripen in late

summer or early fall (Arnow 1987; Meyer and others

1995). Plants tolerate water stress; growth continues

into summer and sometimes after seed ripening (Anderson and others 1987). Some ecotypes are moderately

tolerant of intermittent high water tables, moderate

soil drainage, and winter or spring flooding (Barkworth

and Atkins 1984; Lesperance and others 1978; Morris

and others 1950). Reproduction is from seed or by

tillering or sprouting from rhizomes.

Plants are resistant to fires, particularly those occurring late in the growing season following the onset of

dormancy (Range and others 1982; Zschaechner 1985).

The bunching habit and the presence of coarse, dried

leaves covering the growing points in late summer

provide protection if burns are not severe (Range and

others 1982; Zschaechner 1985). Damage is greater

earlier in the season, particularly if soils are dry,

limiting the potential for resprouting from the root

crown or rhizomes (Bradley and others 1992a,b; Wasser

1982). If moisture conditions are favorable, production

may increase and remain high for several years following fire (Klebenow and others 1976). In addition to

resprouting following fire, stand increase may also

occur from the seed bank, as smoke has been found to

increase the growth of germinants (Blank and Young

1998).

Plant Culture

Wildland stands of Great Basin wildrye may be

hand harvested. Extensive wildland stands and seed

fields may be swathed and combined. Shattering is

only moderate, but lodging may create problems. Seeds

are cleaned by combining or by chopping and fanning

(Plummer and others 1968). There are about 95,000 to

166,000 seeds per pound (210,000 to 366,000 per kg)

Figure 26—Great Basin wildrye provides good winter cover for birds and small mammals (photo courtesy of John Kinney, USDA FS, Rocky Mountain

Research Station, Boise, ID).
376 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

(Vallentine 1971). Seed fill and viability are variable

and often low. Seeds may be marketed with 85 percent

purity and 80 percent germination (Wasser 1982).

Great Basin wildrye is usually difficult to establish

due to low or erratic germination, poor seedling survival, and slow establishment (Evans and Young 1983;

Roundy and others 1983). It should be seeded in late

fall in the drier portion of its range. Seeds should be

drilled or broadcast and covered to a 1-inch (2.5-cm)

depth (Wasser 1982). Seeds are nondormant and do

not require afterripening, but they will germinate

more readily in spring following exposure to cool,

moist seedbeds in winter (Meyer 1994). This allows

the seed to emerge more uniformly in spring and

maximizes survival in the face of drying seedbeds.

High-elevation and spring-flooded sites can be seeded

in spring as soon as they become accessible.

Great Basin wildrye can be drill or broadcast seeded

alone or with other nonagressive species in areas

where it is most likely to establish. It should not be

seeded in mixtures with more aggressive species or in

areas where weeds have not been controlled. High soil

salinity or soil crusting may reduce emergence. Stand

establishment requires two or three growing seasons.

New seedings of Great Basin wildrye should not be

grazed or harvested until after the first two growing

seasons.

Uses and Management

Great Basin wildrye is an important forage species

for wildlife and domestic livestock. It is palatable and

nutritious in spring and fall and receives considerable

use during these seasons (Jarecki 1985). It is also

valuable in winter because of its height and the accessibility of its forage (Cook and others 1954; Jarecki

1985). Palatability and nutrient content decrease in

summer and winter as the leaves dry and become

coarse (Cook and others 1954; Daubenmire 1970;

Jarecki 1985; Krall and others 1971; Walker and

Brotherson 1982; Wasser 1982).

Big game animals including mule deer, elk, and

bighorn sheep graze Great Basin wildrye (Austin and

others 1994; Keating and others 1985; Majerus 1992).

It provides excellent cover for small mammals, waterfowl, and upland game birds, and bedding cover for big

game (Batzli 1974; Sours 1983; Wasser 1982).

All classes of livestock use Great Basin wildrye. It

provides valuable forage and thermal protection for

livestock in winter. Native stands provide spring and

fall grazing and may be cut for hay. Poor grazing

practices, however, have degraded many native stands.

The timing, frequency, and extent of grazing all have

an impact on plant vigor and regrowth (Holzworth and

Lacey 1991; Jarecki 1985). Forage is of high quality in

early spring. Excessive grazing during this period

depletes carbohydrates and is detrimental to the plants,

especially when they are grazed during the boot stage

(Tweit and Houston 1980). Plants should not be grazed

until they reach heights of 12 to 15 inches (30 to 38 cm),

and 20 percent of the production should be left to

maximize regrowth (Hafenrichter and others 1968).

Livestock should be removed before the soil dries out

to permit regrowth and replacement of root reserves.

This approach should also be followed for fall grazing.

Great Basin wildrye is susceptible to infection by

ergot, which may cause sickness or death of livestock

that consume it (Cronquist and others 1977). Plants

are also sometimes severely infested with leaf rusts.

Great Basin wildrye is seeded to provide forage and

cover for livestock and wildlife. It is also planted in

pastures, sometimes with tall wheatgrass, on saline

and alkali soils that are subirrigated or that may flood

intermittently to provide fall, winter, or spring livestock grazing and for calving. These seedings are also

harvested for their hay and silage production.

Great Basin wildrye is valuable for erosion control

plantings in areas where it is adapted because of its

deep and spreading fibrous root system. It is useful for

stabilization of drainage ditches, mine spoils, highway

rights-of-way, burned sites, eroding slopes, and other

disturbances (Barker and others 1985; Cronquist and

others 1977; Walker and Brotherson 1982). Great

Basin wildrye is also seeded as a component of windbreaks and conservation plantings. A major use of the

species is for seeding disturbances on alkaline and

saline sites that once supported the species. Monsen

(1983) recommends that basin wildrye be included in

seedings on disturbances in or adjacent to riparian

areas including wet meadows, stream terraces, and

mountain brush, big sagebrush, desert shrub, and

saltgrass communities. Slow establishment, however,

limits the value of Great Basin wildrye during the first

two growing seasons following seeding.

Varieties and Ecotypes

‘Magnar’, an octoploid cultivar, originated in

Saskatchewan, Canada (Alderson and Sharp 1994),

and has been seeded extensively throughout the Intermountain West in upper pinyon-juniper, mountain

brush, and mountain big sagebrush communities. It

also establishes well in canyon bottoms, along road

rights-of-way, and in depressions where extra moisture accumulates. It is adapted to soils ranging from

loams to clays, and from slightly basic to alkaline.

Magnar performs best on sites receiving 15 inches

(380 mm) or more of annual precipitation, but stands

establish in areas receiving as little as 13 inches

(330 mm). It is not suited to lower, drier pinyonjuniper or sagebrush grass communities.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 377

Chapter 18 Grasses

Magnar foliage is blue green. Plants grow to 7 ft

(2.1 m) in height and produce wide, coarse leaves and

large stems and seed heads. Seedlings are vigorous

and establishment is generally rapid. Magnar is long

lived, nonaggressive, and suitable for mixed seedings.

‘Trailhead’ was collected near Roundup, MT

(Alderson and Sharp 1994). Plants are tetraploid and

dark green with broad, somewhat lax leaves. Trailhead

is considerably more drought tolerant and productive

than Magnar. It has established successfully in areas

receiving as little as 11 inches (280 mm) of annual

precipitation. Trailhead is adapted to soils ranging

from saline-alkaline to acidic. It can be seeded from

valley bottoms up through the mountain brush zone.

Trailhead is seeded to provide soil stabilization, early

spring and winter forage, and wildlife habitat.

Elymus glaucus

Blue Wildrye ___________________

Synonyms

Elymus sibericus var. glaucus

Elymus villosus var. glabriusculus

Elymus nitidus

Description

Blue wildrye is a green to blue waxy cool-season

native perennial bunchgrass that forms small loose to

dense tufts with only a few culms. The root system is

deep and fibrous. Plants sometimes produces short

stolons. Culms grow from 1.9 to 4.6 ft (6 to 14 dm) in

height. Leaves are scattered along the culms. Leaf

sheaths may be glabrous or with downward pointing

hairs; the collar is often purple. Leaf blades are flat to

sometimes slightly involute, 0.16 to 0.47 inch (4 to 12

mm) wide, and mostly roughened to sparsely silky

haired, or sometimes hairless below. Auricles are well

developed, about 0.08 inch (2 mm) long, and clasping.

Ligules are short, 0.01 to 0.04 inch (0.3 to 1 mm) long,

squared at the apex, and irregularly lacerate and

ciliate along the margins. Spikes are 2.4 to 6.3 inches

(6 to 16 cm) long, excluding the awns, erect, interrupted below and overlapping above, or dense throughout and long exerted. Spikelets are mostly two per

node, 0.39 to 0.63 inch (10 to 16 mm) long, and two to

four flowered. Glumes are nearly parallel, concealing

the base of the enclosed florets and tapering to a short

awn. They are subequal, 0.25 to 0.75 inch (6.5 to 19 mm)

long, three to five nerved, narrowly lance shaped, and

broadest above the base. The glumes are hardened

below and membranous above with glabrous, smooth

to roughened surfaces. Lemmas are 0.33 to 0.55 inch

(8.5 to 14 mm) long, five nerved, hairless to roughened

along the nerves, and tapering to long, slender, mostly

straight awns that are about 0.39 to 1.18 inches (10 to

30 mm) long. Plants flower from June to August

(Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969).

Blue wildrye is often confused with Agropyron

trachycaulumvar. unilaterale. Blue wildrye, however,

has the Elymus characteristics of two spikelets per

node, narrow glumes, and a distorted rachilla. It is,

however, extremely variable in its morphological and

genotypic characters. Some biotypes cross readily with

Sitanion hystrix, while others do not. These hybrids

are referred to as X elysitanion hansenii. Some manuals recognize blue wildrye varieties such as E. g. var.

breviaristatus, E. g. var. glaucus, and E. g. var. jepsonii

(Cronquist and others 1977). The species is

allotretraploid with 2n = 28 (Dewey 1982).

Ecological Relationships and Distribution

Blue wildrye is distributed from Alaska to California and into northern Arizona and New Mexico. It is

found less commonly from Ontario to New York, and

south to Arkansas (Arnow 1987, 1993; Cronquist and

others 1977). Blue wildrye is most abundant in timbered areas from sea level to 10,000 ft (3,050 m) (Asay

and Jensen 1996b). It occurs in shaded or open forests,

grasslands, shrublands, and on moist soils of riparian

areas and wetlands in mountain brush, aspen, ponderosa pine, spruce-fir, and lodgepole pine communities

(Arnow 1987; Hitchcock 1950) (fig. 27).

Blue wildrye populations are adapted to well-drained

soils with clay loam to sandy loam textures (Hassell

and others 1982) that are moderately acid to neutral

(pH 5.2 to 7). Populations are common on moist sites,

but some are moderately drought tolerant. The species

is adapted to soils of low fertility, but it does not

tolerate shallow soils, and it is moderately sensitive to

saline soils (Hassell and others 1983).

Plants are short lived, but stands reseed themselves

readily (Asay and Jensen 1996b). The species is favored by disturbances and is often most abundant in

early seral communities on open, disturbed sites where

it may form dense stands (Cole 1982; Hitchcock and

others 1969; USDA Forest Service 1937). Plants also

grow in shaded areas (Plummer and others 1955).

Blue wildrye forms small bunches with coarse foliage that burn rapidly and can survive wildfires. Burned

plants regenerate by resprouting from the root crown

(Simmerman and others 1991). New plants establish

from the seed bank, as burns appear to provide favorable microsite conditions for germination (Stickney

1989). Blue wildrye seedlings and cover develop rapidly, and stands of the species are common on burned

over sites where competition is low. Stands may decline after the first few years (Brown and DeByle 1989;

Hafenrichter and others 1949, 1968).
378 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Plant Culture

Blue wildrye tends to be a good seed producer.

Shattering can reduce yields, but seed can be safely

harvested shortly before reaching maturity and allowed to ripen while drying (Link 1993). Seed must be

dried before hammermilling to facilitate removal of

the awns (Hafenrichter and others 1949, 1968; Link

1993). Awns of seeds in large lots may be detached

with a debearder or brush machine. Debris is then

removed using an air screen machine.

There are about 131,000 seeds per pound (288,800

per kg) (Hafenrichter and others 1968). Seeds maintain viability for 2 to 4 years. They do not require

a pretreatment for germination (McLean 1967).

Seeds should be incubated at 68/86 ∞F (20/30 ∞C)

(16 hrs/8 hrs) for germination testing (Haferkamp

and McSwain 1951).

Some seed is commercially available. Libby and

Rodrigues (1992), however, suggested that use of

commercial seed outside its area of origin could contaminate local populations. Hassell and others (1983)

recommended use of local seed collected or produced

from populations growing within 300 miles (483 km)

and 1,640 ft (500 m) elevation from the planting site

because of the high variability within the species.

Blue wildrye can be drill or broadcast seeded on a

firm seedbed in fall or early spring. Seed should be

covered to a depth of about 0.5 inch (1.3 cm) (Plummer

1943). Establishment and growth are rapid and

seedings are persistent.

Uses and Management

Although blue wildrye provides fair to good forage

for big game and livestock early in the growing season,

the foliage is coarse and provides only poor protein

content and good energy value (Dittberner and Olsen

1983). The species produces a fairly extensive root

system, but plants are not tolerant of heavy grazing

(Vallentine 1961). Blue wildrye recovers fairly rapidly

when grazing is removed (Sampson 1944). A widespread species, it often provides considerable forage

for wildlife. Early spring growth is most readily used;

the awned seedheads are avoided (USDA Forest Service 1937).

Blue wildrye establishes rapidly and is a good seed

producer (Plummer and others 1968). Consequently,

it is used for postfire and postlogging seedings because

it generally establishes well and can be planted in

mixtures with other natives (Frishchknecht and

Plummer 1955). It is a pioneering species and can be

seeded to provide cover rapidly, spread from seed, and

control erosion on steep slopes, roadsides, and other

disturbances (Darris and others 1996; Hafenrichter

and others 1949, 1968). Seedings of blue wildrye do not

compete excessively with establishing tree seedlings,

but the grass does restrict invasion of weedy species

(Asay and Jensen 1996b). Blue wildrye has also been

used to provide an understory cover in windbreak and

conservation plantings.

Varieties and Ecotypes

Blue wildrye contains considerable intraspecific variability manifested by differing site requirements.

Plants are highly self-fertile. Some ecotypes are crosssterile, while others are compatible (Asay and Jensen

1996b; Snyder 1950). Seed transfer zones for the

species have been identified for National Forests of

north-central Oregon.

‘Arlington’ originated in Snohomish County, WA

(Alderson and Sharp 1994). It has been seeded primarily for soil stabilization and to provide forage. It grows

in shaded areas under the canopies of aspen groves

Figure 27—Blue wildrye is a native, shade-tolerant

bunchgrass that can be seeded as an understory in

tree and shrub plantings (photo courtesy of Kevin

Jensen, USDA ARS, Logan, UT).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 379

Chapter 18 Grasses

and conifer forests, but it also does well on the forest

edge and in openings, especially on moist sites and

along stream edges. Arlington exhibits good seedling

vigor, establishes readily, and provides cover for erosion control on burned and cutover forest areas. Plants

are short lived, but the variety does spread from seed.

Seed may be drilled shallowly or broadcast following a

fire or on snow in winter. Seed is now rarely available.

Elymus junceus

Russian Wildrye ________________

Synonyms

Elymus cretaceus

Psathyrostachys juncea

Description

Russian wildrye is a densely tufted, perennial, coolseason bunchgrass with the leaves mostly basal. Culms

are 1.3 to 3.6 ft (4 to 11 dm) tall. Leaf sheaths are

hairless; the auricles and collars are well developed.

Blades are flat, waxy, roughened, involute, and 0.06 to

0.16 inch (1.5 to 4 mm) wide. Ligules are very short,

0.01 to 0.04 inch (0.2 to 1 mm) long, squared at the

apex, and ciliate along the margins. Spikes are erect,

1.6 to 5.1 inches (4 to 13 cm) long, and 0.16 to 0.47 inch

(4 to 12 mm) wide with the rachis disarticulating

between the spikelets. Spikelets are three or sometimes

two per node. They are 0.31 to 0.39 inch (8 to 10 mm)

long and two to three flowered. Glumes are subequal,

short, 0.12 to 0.27 inch (3 to 7 mm) long, awl shaped,

roughened or short haired, but often hairless and

shiny at the base. Lemmas are small, 0.26 to 0.39 inch

(6.7 to 10 mm) long, roughened to stiff haired, five to

seven nerved, and tapering to an awn that is 0.02 to

0.12 inch (0.5 to 3 mm) long (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969).

Russian wildrye is easily identified by the readily

disarticulating axis of the rachis at maturity. This

results in the accumulation of old leaf sheaths at the

base of the plant that eventually separate into fibers.

Plants flower from May to July (Arnow 1987, 1993;

Cronquist and others 1977; Goodrich 1986; Hitchcock

1950; Hitchcock and others 1969).

Ecological Relationships and Distribution

Russian wildrye, a native of the steppes and deserts

of Russia and China, was introduced to the United

States in 1927. It has been seeded on the Northern

Great Plains and the Intermountain and Rocky Mountain areas as a pasture grass and erosion control

species since the 1950s (fig. 28). Russian wildrye

provides nutritious and palatable forage nearly year

long on arid and semiarid rangelands (Plummer and

others 1968). It is now distributed from Canada south

to Arizona, and eastward to Nebraska (Arnow 1987).

Scattered naturalized plants have been found in Utah,

Wyoming, Saskatchewan, and Manitoba (Cronquist

and others 1977). In the Intermountain area the species is adapted to salt desert shrub, sagebrush, grasslands, pinyon-juniper, oak scrub, mountain brush,

aspen, and ponderosa pine communities. It grows in

openings and in the shade of shrubs, but it is not

adapted as an understory species in the shade of

Gambel oak and mountain brush (Plummer and others 1968). Russian wildrye is adapted to fertile loam

and clay soils, but it will grow on a variety of soil types

including moderately saline or alkaline soils. It can be

seeded on soils that are too alkaline for crested wheatgrass. Russian wildrye is more heat tolerant than

crested wheatgrass in areas receiving summer precipitation, and matures later, providing a longer period of grazing. It is best adapted to areas with winter

and summer precipitation or a predominance of summer precipitation. The species is drought and cold

tolerant. It does not tolerate spring flooding (Jensen

and others 2001).

Plant Culture

Russian wildrye seed matures in July to August.

Seed production is generally low. Crops can be lost to

wind or rainstorms occurring during the ripening

Figure 28—Russian wildrye is drought tolerant and

produces palatable forage throughout the summer

(RMRS photo).
380 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

period, and seeds tend to shatter shortly after reaching maturity. Fields are windrowed or direct combined. They can be grazed following harvest to remove

litter (Jensen and others 2001). There are about 170,100

Russian wildrye seeds per pound (375,000 per kg)

(Heady 1975). Fresh or dormant seed becomes more

germinable following a 5-day prechill at 41 or 50 ∞F (5

or 10 ∞C) (AOSA 1999). Seed is incubated at an alternating temperature regime of 68/86 ∞F (20/30 ∞C) (16

hrs/8 hrs) for 14 days for germination testing.

Wildland seedings can be established in areas receiving as little as 8 inches (20 cm) of annual precipitation. Seed should be drill seeded or broadcast and

covered in fall at a depth of about 0.75 inch (1.9 cm)

(Ludwick 1976). Seeds are easily handled and planted.

Seedling vigor is low to moderate, and slow establishment can preclude stand success when weedy competition is present and during dry periods because few

seminal roots are produced (Jensen and others 2001).

Stands are seeded at wide row spacings, as established

plants are strongly competitive and self-thinning until widely spaced. The species is difficult to establish in

mixes with species having more rapidly developing

seedlings.

Uses and Management

Russian wildrye is seeded primarily to provide forage in dry areas because of its basal leaf production.

Grazing of newly seeded stands should be delayed

until plants are mature. Established stands are long

lived and tolerant of grazing on fertile sites; stands

decline more rapidly on sites of low fertility. The

species begins growth early in the season, and the

numerous basal leaves develop rapidly. Unlike many

cool-season grasses of dry areas, it is palatable and

nutritious from spring into fall and exhibits good

regrowth following use (Jensen and others 2001).

Seedings of the species provide good quality and quantities of fall and winter forage when other usable

forage is often not available. Although seedlings of this

species develop slowly, established stands are more

competitive than associated native species.

Russian wildrye is seeded in pastures and maintained with fertilizer treatments in some areas (Jensen

and others 2001). Legumes may be added to improve

productivity. The species is not generally used for hay

production. Wildlife use the species, and it can provide

valuable forage in early spring and also in late summer when other grasses have dried out (Dittberner

and Olson 1983; Plummer and others 1968). It provides some cover for small mammals and birds. Because plants tend to be widely spaced, it is not often

used to control erosion; however, it has been used on

roadside areas in southern Idaho to reduce blowing

sand and dust.

Varieties and Ecotypes

‘Bozoisky-select’, an introduction from Russia

(Alderson and Sharp 1994), was selected for its large

seed size, good seedling vigor and establishment, and

enhanced forage production. It is seeded throughout

the Intermountain West in the same areas as ‘Vinall’,

but exhibits better establishment and productivity.

‘Cabree’ was developed from Russian material for

use on the Northern Great Plains and Canadian prairies (Alderson and Sharp 1994). It was selected for its

seedling vigor, forage and seed production, and moderate drought tolerance. It has not been seeded extensively in the Intermountain West.

‘Mankota’, introduced from Alma-Ata in the former

U.S.S.R. (Alderson and Sharp 1994), was developed

for use on the Northern Great Plains and has received

little use in the Intermountain West. It is similar to

Vinall and Bozoisky-select.

‘Mayak’ was selected for use on the Northern Great

Plains and Canadian prairies (Alderson and Sharp

1994), and has only infrequently been seeded in the

Intermountain West. Mayak provides excellent forage

throughout the grazing season and has good curing

qualities that make it especially useful for late summer, fall, and early winter grazing. Its characteristics

include good forage and seed yield and resistance to

leaf spot disease.

‘Swift’, derived from ‘Saki’ and ‘Mandan 1546’

(Alderson and Sharp 1994), is adapted to areas of the

Intermountain West, even though it was developed for

the Northern Great Plains and Canadian prairies. It

was selected for its seedling vigor, early spring growth,

forage and seed production, and leafiness. It has not

proven superior to Bozoisky-select.

‘Tetracan’ is a colchicine-induced tetraploid (Alderson

and Sharp 1994) with large seedheads and seeds, wide

leaves, and good seedling vigor. It has performed well

on adapted sites, especially with regard to seedling

vigor and establishment. Tetracan has seen little use

in the Intermountain West.

Vinall was introduced from Russia (Alderson and

Sharp 1994) and has been seeded widely throughout

the Intermountain West, especially in pinyon-juniper, basin big sagebrush, Wyoming big sagebrush,

black greasewood, and salt desert shrub communities. It is adapted to clay and clay-loam soils, and it is

moderately tolerant of saline and sodic soils. A major

drawback of this variety is its slow and erratic germination and establishment. It does not do well when

seeded with species that establish and develop more

rapidly. Mature stands can prevent the establishment of other species. This variety is long lived and

drought tolerant.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 381

Chapter 18 Grasses

Elymus salinus

Salina Wildrye __________________

Synonyms

Elymus ambiguus var. salina

Elymus ambiguus var. salmonis

Leymus salinus

Elymus salina

Description

Salina wildrye is an erect, densely tufted perennial

bunchgrass that occasionally produces short rhizomes.

On some sites, particularly those with heavy clay soils,

the plants form large tufts that give them the appearance of being nonrhizomatous. Plants are coarse

stemmed with glabrous culms 1.1 to 3.3 ft (3.5 to 10

dm) tall. Leaves are glabrous to pubescent. Leaf blades

are flat or more commonly involute, 0.08 to 0.16 inch

(2 to 4 mm) wide, and often ciliate near the throat.

Auricles are well developed and often clasping. Ligules

are usually short, about 0.01 to 0.06 inch (0.2 to 1.5

mm) long, squared apically, and ciliate along the

margins. Spikes are erect, slender, 1.9 to 5.9 inches

(5 to 15 cm) long, and 0.10 to 0.31 inch (2.5 to 8 mm)

wide. Spikelets are usually solitary at each node and

slightly overlapping. They are 0.35 to 0.79 inch (9 to 20

mm) long and two to three or rarely six flowered.

Glumes are subequal, awl shaped, three nerved, smooth

to roughened, sometimes broader at the base, and 0.05

to 0.39 inch (1.2 to 10 mm) long. There is often an extra

glume at nodes producing only one spikelet. Lemmas

are five-nerved (at least above), smooth to slightly

roughened on the surfaces, roughened on the awn and

midnerve, and 0.28 to 0.41 inch (7 to 10.5 mm) long.

Lemmas taper to a short awn that is 0.01 to 0.08 inch

(0.2 to 2 mm) long. The species includes tetraploid

(2n = 28) and octoploid (2n = 56) populations that do

not differ morphologically. Plants flower in May to

July (Arnow 1987, 1993; Cronquist and others 1977;

Dittberner and Olsen 1983; Hitchcock 1950; Hitchcock

and others 1969).

Ecological Relationships and Distribution

Salina wildrye is distributed from California and

southern Nevada to southwestern Wyoming. It is

found from salt desert shrublands through desert

shrub, sagebrush-grass, mountain mahogany, aspen,

and conifer communities. It may be a common to

dominant species in Gambel oak and pinyon-juniper

communities (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969).

Salina wildrye is drought resistant and adapted to dry

sites with loamy soils, but it also occurs on rocky to

sandy sites and on clay soils. It is moderately tolerant

of alkalinity (Cronquist and others 1977). The species

grows on hillsides, alluvial fans, plateaus, bluffs, canyons, and mountainous areas (Baker and Kennedy

1985; Vallentine 1989) (fig. 29). Plants are generally

scattered, and normally do not form pure stands. The

species may be resistant to wildfire due to its rhizomatous nature and the widely spaced bunching habit

(Baker and Kennedy 1985).

Salina wildrye spreads from seed and tillers. Rhizomes are more commonly produced under mesic conditions. On clay soils, rhizomes may be very short and

inconspicuous.

Plant Culture

Seeds of salina wildrye mature in mid to late July.

Fruits are cleaned with an air screen machine. There

are about 130,000 seeds per pound (286,600 per kg)

(Link 1993). Seeds are estimated to remain viable for

2 to 3 years. Germination is enhanced by a 30-day

moist prechill (Link 1993).

Seedings of salina wildrye are often not successful

due to low seed germination and poor seedling vigor.

Established stands, however, are persistent and

drought tolerant under a wide variety of conditions

(Plummer and others 1968). The species has potential

for increased use due to its wide range of adaptation,

salt tolerance, drought resistance, and ability to stabilize disturbed slopes with clay soils (Goodrich and

Neese 1986; Plummer 1977; Vallentine 1989).

Uses and Management

Salina wildrye is moderately productive. Quality

and palatability of the coarse forage is only fair when

green; it becomes unpalatable when cured (Vallentine

Figure 29—Salina wildrye grows in mountainous areas and on hillsides, alluvial fans, and canyons from

California to southwestern Wyoming (RMRS photo).
382 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

1989). Excessive grazing has resulted in declines of

the species in northwestern Colorado (Baker and

Kennedy 1985).

Varieties and Ecotypes

The Upper Colorado Environmental Plant Center in

Meeker, CO, is advancing ‘Accession 9043501’ from

Colfax County, NM, toward release (UCEPC 2002).

This accession has potential for revegetation of dry,

moderately alkaline rangelands and other disturbed

sites in the West. Its forage quality is moderate to fair

for livestock and wildlife.

Elymus simplex

Alkali Wildrye or Low-Creeping

Wildrye ________________________

Synonyms

Elymus triticoides var. simplex

Elymus simplex var. luxurians

Leymus simplex

Description

Alkali wildrye is a waxy perennial that develops

from extensively creeping rhizomes. Culms are 1.3 to

2.3 ft (4 to 7 dm) tall and the rhizomes can be up to

16.4 ft (5 m) long. Leaves are mostly basal. Leaf

sheaths are hairless. The blades are usually involute

to more or less flat, 0.04 to 0.12 inch (1 to 3 mm) wide,

rigid, and roughened or sparsely hairy on the upper

surface. Auricles are poorly developed and usually less

than 0.04 inch (1 mm) long. Ligules are very short,

about 0.02 inch (0.5 mm) long, squared at the apex,

and ciliate along the margins. Spikes are 2.4 to 6.3

inches (6 to 16 cm) long, erect, and usually rather

slender (fig. 30). Spikelets are usually solitary, or

sometimes two per node. When two per node, one is

sessile and the other is short pedicellate. Spikelets

are 0.43 to 0.80 inch (11 to 20 mm) long, three to ten

flowered, sometimes well spaced and not overlapping,

but usually slightly overlapping. Glumes are subequal,

0.22 to 0.63 inch (5.5 to 16 mm) long, awl shaped to only

slightly broader at the base, hairless to roughened,

nearly nerveless, or one to three nerved. There is

sometimes an extra glume at a one-spikelet node.

Lemmas are 0.30 to 0.37 inch (7.5 to 9.5 mm) long,

usually hairless to minutely pubescent and shiny

green or rarely with a purplish tinge. They are nerveless

below and faintly five to seven nerved above. Glumes

taper to a 0.10- to 0.31-inch (2.5- to 8-mm) awn. Plants

flower from June to August.

Alkali wildrye is sometimes confused with small

plants of creeping wildrye with solitary spikelets. It is

also confused with salina wildrye plants that produce

rhizomes. Location is the best way to distinguish

between these two. Alkali wildrye is a grass of sandy

bottomlands, whereas salina wildrye usually grows on

mountainsides and in heavy clay soils (Arnow 1987,

1993; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969).

Ecological Relationships and Distribution

Alkali or low-creeping wildrye is found in moist,

sandy soils along riverbanks, and sometimes in drifting sands along the Green River in Daggett County,

UT. It also occurs in Wyoming and Colorado (Arnow

1987, 1993; Cronquist and others 1977).

Plant Culture, Uses, and Management

Alkali wildrye has received little use in revegetation

efforts. The species could be used on native sites to

provide cover, forage, soil stabilization, and diversity.

Varieties and Ecotypes

There are no releases.

Figure 30—Alkali wildrye spikes are elongate and

slender with the spikelets generally solitary or sometimes two per node (Hitchcock 1950).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 383

Chapter 18 Grasses

Elymus triticoides

Creeping Wildrye or Beardless

Wildrye ________________________

Synonyms

Elymus condensatus var. triticoides

Leymus triticoides

Elymus simplex var. luxurians

Elymus acicularis

Description

Creeping or beardless wildrye is a waxy blue or

sometimes green perennial that produces extensive

and aggressive creeping rhizomes. Culms are 0.5 to

3.9 ft (1.5 to 12 dm) tall. Leaf sheaths are glabrous,

roughened, or sometimes minutely hairy. Leaf blades

are flat or more commonly involute, and measure

0.10 to 0.28 inch (2.5 to 7 mm) in width. They are

coarse, stiff, roughened, and sometimes glabrous below. Auricles are well developed and often clasping.

Ligules are short, 0.01 to 0.28 inch (0.2 to 0.7 mm) long,

squared, and irregularly lacerate and ciliate along the

margins. Spikes are slender and about 1.4 to 3.5 inches

(3.5 to 9 cm) long, erect, loose, open to rather dense,

and sometimes compound. Spikelets are paired or

sometimes solitary at the nodes, and 0.39 to 0.70 inch

(10 to 18 mm) long. They are three to eight flowered

and greenish, brownish, or purplish. Glumes are

subequal, 0.16 to 0.43 inch (4 to 11 mm) long, narrow

and awl shaped, firm, one to three nerved, and mostly

roughened. Lemmas are small, 0.24 to 0.39 inch (6 to

10 mm) long, often shiny and smooth, hairless, minutely hairy, or sometimes minutely hairy only at the

apices. They are faintly to prominently five to seven

nerved, rounded on the back or keeled toward the tip,

and awnless or with a short awn about 0.02 to 0.10 inch

(0.5 to 2.5 mm) long. Plants flower from May through

August.

Considerable ecotypic variation occurs within the

species. Creeping or beardless wildrye can be confused

with western wheatgrass when there is only one spikelet per node. Very narrow glumes and a twisted rachilla

that cause the florets to be out of their normal position

are characteristic of creeping wildrye (Arnow 1987,

1993; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969), and although of differing

growth habits, creeping wildrye and Great Basin

wildrye share similar genomes and hybridize in nature (Dewey 1984). Creeping wildrye also hybridizes

with salina wildrye (Arnow 1987).

Ecological Relationships and Distribution

Creeping wildrye is distributed from Washington to

Montana, and south from Baja California to Texas. It

is especially common in the Humboldt River drainage

of Nevada (Jensen and others 2001). It is a long-lived,

salt-tolerant, sodforming perennial that grows in dry

to moist places, valley bottoms, saline seeps and meadows, subirrigated sites including alkaline areas with

high water tables, and sites exposed to spring flooding.

It occurs on saline meadows and salt desert shrub

lands receiving 8 to 12 inches (20 to 30.5 cm) of annual

rainfall. The species is cross-pollinating and frequently

does not set seed. Stands are generally not extensive;

patches may represent single clones (fig. 31). Creeping

wildrye often forms natural hybrids with Great Basin

wildrye and salina wildrye (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969; Knapp and Wiesner 1978).

Plant Culture

Creeping wildrye seeds mature between midsummer and early fall (Sampson 1924). Fields may be

combined or swathed and windrowed. Seed retention

at maturity is good, but lodging can interfere with

harvest. Yields from seed fields average 100 to 200

pounds per acre (112 to 224 kg per ha) (Smith and

Smith 1997). Stand life is about 5 years. Seeds are

cleaned with a fanning mill. There are about 175,000

seeds per pound (385,000 per kg) (Smith and Smith

1997). Dormancy results from an impermeable outer

seed coat. Germination is hastened by a 5-day prechill

at 1.5 ∞C (Wagner and Chapman 1970) with incubation at 15/20 ∞C (Wagner and Chapman 1970) or 20/

30 ∞C (16 hrs/8 hrs) (Haferkamp and McSwain 1951).

Creeping wildrye is fall planted to provide overwinter

exposure to cool, moist seedbed conditions. Seeds are

drilled or broadcast and covered. Slow germination,

Figure 31—Patches of creeping wildrye often represent individual clones (photo courtesy of Tom

Jones, USDA ARS, Logan, UT).
384 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

low seedling vigor, and resulting poor establishment

have limited use of this species. Although young stands

are not competitive, established stands are highly

persistent.

Uses and Management

Creeping wildrye is seeded to provide soil stabilization on highly erodible sites and can be used for

reseeding disturbances along waterways and on mined

lands, roadways, and disturbed rangelands. It is one of

the few species that can be seeded to provide forage on

wet, alkaline sites in low-precipitation areas, and it is

useful in saline areas and areas that are flooded in

spring. It can be used in native species restoration

efforts in salt desert shrub and pinyon-juniper communities. Native stands provide cover and forage for

wildlife and are sometimes cut for hay (Jensen and

others 2001).

Varieties and Ecotypes

‘Rio’ was released by the Lockford Plant Material

Center in 1991. It was selected for its vigorous spreading habit. ‘Shoshone’, released in 1980 by the Bridger

Plant Materials Center, is a productive and rhizomatous cultivar.

Festuca arundinacea

Tall Fescue, Reed Fescue, or Alta

Fescue ________________________

Synonyms

Bromus arundinaceus

Festuca elatior var. arundinacea

Description

Tall fescue is a stout, often strongly tufted perennial

with an extensive coarse, fibrous root system. Rhizomes are generally short and vary considerably in

their sodforming ability. Culms are 1.6 to 6.6 ft (0.5 to

2 m) tall and ascending or erect with smooth or

roughened leaf sheaths. Leaf blades are stiff, flat or

somewhat involute, 0.12 to 0.47 inch (3 to 12 mm)

wide, and roughened above. Auricles are prominent

and usually ciliate. Ligules are about 0.01 inch (0.2

mm) long. Panicles are narrow, 5.9 to 13.8 inches (15

to 35 cm) long, and often somewhat nodding. Spikelets

measure 0.4 to 0.6 inch (10 to 15 mm) in length and are

five to nine flowered. Glumes are lance shaped. The

first is 0.16 to 0.24 inch (4 to 6 mm) long and one to

three nerved; the second is 0.16 to 0.28 inch (4 to 7 mm)

long and three to five nerved with translucent margins. Lemmas are 0.3 to 0.4 inch (7 to 10 mm) long with

five to seven faint nerves. Awns, when present, are up

to 0.08 inch (2 mm) long. Plants flower from late May

through July (Arnow 1987, 1993; Cronquist and others 1977; Hitchcock 1950; Hitchcock and others 1969).

The species is allohexaploid with 2n = 6n = 42 (Sleper

and West 1996).

Ecological Relationships and Distribution

Tall fescue, native to Europe, is seeded as hay,

pasture, or turf grass in the Western Hemisphere. The

species contains considerable variability, and many

cultivars are available. Leaves of the forage varieties

are generally soft and lax compared to those of the turf

varieties (Jensen and others 2001). Tall fescue is

cultivated from Alaska and southern Canada throughout much of the United States (fig. 32). It is also

present in South America. The species has escaped

and become widely established in areas where it has

been seeded. Tall fescue can now be found in seeded

fields, moderately moist areas along irrigation ditches

and roadsides, and in fallow fields and seeded mountain meadows. Its best growth is on deep, moderately

fertile, fine silty to clay soils, but it will grow well on a

wide range of soil textures when moisture is adequate

(Wasser 1982). It can also be seeded on moderately

saline and alkaline soils (Jensen and others 2001).

Growth is good in areas that are irrigated, subirrigated,

or that receive at least 18 inches (460 mm) of annual

precipitation (Jensen and others 2001; Wasser 1982).

Tall fescue is a relatively drought- and heat-tolerant

Figure 32—Reed fescue grows on moist soils and

tolerates saline and alkaline conditions. Grown for

pasture and hay production, it has escaped and

become widely distributed in North America (RMRS

photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 385

Chapter 18 Grasses

pasture grass, but it does not survive long dry periods.

This species tolerates cold, winter flooding, high water

tables, and poorly drained soils, and it is moderately

shade tolerant. Plants are resistant to burning when

dormant, but the species does not commonly occur in

areas susceptible to wildfire.

An endophytic fungus, Acremonium coenophialum,

frequently infects tall fescue (Sleper and West 1996).

When infected forage is consumed in large quantities,

susceptible livestock may develop lameness and sometimes gangrene (Kingsbury 1964). Endophyte-free

cultivars should be used for forage production, but

these often require more fertile soils or fertilizer applications. Tall fescue is also an invader of lawns (Arnow

1987, 1993; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969).

Plant Culture

Seeds are harvested by direct combining or by swathing and combining. Some seed loss normally results

from shattering. Seeds are cleaned using an air-screen

machine. There are 181,000 to 230,000 seeds per

pound (399,000 to 507,100 per kg). Minimum purity

for seed purchases is 98 percent, and germination is 85

percent (Wasser 1982). Seeds are drilled or broadcasted seeded and covered to 0.25 to 0.5 inch (0.6 to 1.3

cm) deep depending on soil conditions. Germination

and initial seedling establishment and growth are

moderate to good (Hassell and others 1983; Plummer

and others 1968). Seedlings usually become established after one growing season (Jensen and others

2001). Plants are persistent, good seed producers, and

spread well from seed. Seed production is centered in

the Northwest, and yields of 2,510 pounds per acre

(2,240 kg per ha) are obtained in western Oregon

(Sleper and West 1996).

Irrigated tall fescue fields are planted in late summer. Weed infestations must be controlled until plants

are established. When legumes are added to improve

forage palatability and quality, seeding may be done

in early spring (Jensen and others 2001). The legumes

are often planted in alternate rows with the grass.

Ladino clover, alfalfa, white clover, and birdsfoot trefoil are used for this purpose (Sleper and West 1996;

Wasser 1982). Annual nitrogen applications are required when tall fescue is seeded without legumes.

Wildland sites in mountainous areas are seeded in

late fall or early summer. The resulting stands are

competitive and persistent when established, but they

may be somewhat slow to develop. New seedings

should not be grazed until plants are well established

Uses and Management

Tall fescue seedings provide pasture or hay in irrigated fields in valleys of the Intermountain region.

They are particularly useful on saline and heavy

alkaline soils where few other grasses can be seeded

(Arnow 1987). The species is persistent, palatable,

moderately to highly productive, and compatible with

a variety of management programs (Hanson 1979).

Tall fescue can be grown with selected legumes if

managed carefully (Wasser 1982). When grown alone,

it requires added nitrogen and irrigation to maximize

productivity (Thornburg 1982). Tall fescue is tolerant

of periodic or continuous grazing and can be grazed or

mown to a 2- to 4-inch (5 to 10 cm) stubble height

(Plummer and others 1968; Wasser 1982). Grazing

should be applied evenly to maintain a uniform stand.

Frequent grazing improves palatability, induces new

tiller production, and maintains a dense sod (Wasser

1982). Stands are more resistant to trampling than

most pasture grasses.

Tall fescue provides a long grazing season. It receives greatest use in spring and early summer when

leaves are young. Its productivity and palatability are

greatly reduced in late summer when the leaves become coarse (Hassell and others 1983). Summer and

fall growth retain digestibility and crude protein content better than many cool-season grasses (Matches

1979). Leaves remain green and provide good-quality

fall and winter forage for livestock. Tall fescue provides excellent quality hay (Jensen and others 2001).

Tall fescue increases over time when planted with

other pasture grasses due to its lower palatability

(Jensen and others 2001). Cutworms, slugs, and grasshoppers can reduce yields.

Tall fescue has been seeded to provide forage in

mountain meadows and in openings in aspen and

conifer forests (Vallentine 1971). It is used by elk and

deer, but its palatability to big game is generally low.

Seeds are consumed by songbirds, and the seeds and

foliage are used by small mammals. Sodforming cultivars are used to stabilize soils on road cuts and fills,

and disturbances in riparian areas and recreational

areas, and to provide cover crops. Fertilizer applications may be required when using the species on

infertile soils (Vallentine 1967).

Varieties and Ecotypes

‘Alta’ and ‘Fawn’ were developed from European

material (Alderson and Sharp 1994) and are adapted

to valley bottoms where they are seeded as pasture

grasses and for the renovation of saltgrass sites. Both

varieties are also used for soil stabilization and forage

production in semiwet meadows of upper elevation

areas and in openings in aspen and conifer forests.

They require at least 16 inches (410 mm) of annual

precipitation. Both green up early in spring. They are

compatible with other species and can be included in

mixtures.
386 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Several tall fescue varieties have been developed for

hay and pasture forage production. These include

‘Barcel’, ‘Barvetia’, ‘Cajun’, ‘Courtenay’, ‘Dovey’,

‘Festorina’, ‘Fuego’, ‘Georgia No. 5’, ‘Goar’, ‘Kentucky

31’, ‘Martin’, ‘Maximize’, ‘Mozark’, ‘Penngrazer’,

‘Phyter’, ‘Southern Cross’, ‘Stargrazer’, and ‘Stef’

(Alderson and Sharp 1994). None of these varieties

have proven adapted to mountain rangeland conditions. There are a number of other varieties that have

been bred for use as turf grasses.

Festuca idahoensis

Idaho Fescue or

Bluebunch Fescue ______________

Synonyms

Festuca ovina var. ingrata

Festuca ingrata

Festuca occidentalis var. ingrata

Festuca ovina var. oregona

Festuca ingrata var. nudata

Festuca ovina var. nudata

Description

Idaho fescue is one of the most common bunchgrasses in the West. It is a densely tufted

nonrhizomatous, long-lived perennial bunchgrass with

hairless to roughened culms that grow 1 to 3.3 ft (3 to

10 dm) tall (fig. 33). Leaves are numerous, fine, and

mostly basal, forming a tuft 6 to 12 inches (15 to 30 cm)

tall that is usually more than half the length of the

culms. Leaf blades are firm, involute, less than 0.04

inch (1 mm) wide, and without auricles. Ligules are

mostly 0.01 to 0.02 inch (0.3 to 0.6 mm) long, finely

ciliate, and longer on the sides than at the center.

Panicles are 1.9 to 5.9 inches (5 to 15 cm) long, loosely

compressed, and extending mostly from one side of the

rachis. Spikelets are 0.31 to 0.51 inch (8 to 13 mm) long

and four to seven flowered with visible joints on the

axis of the spikelet. Glumes are narrowly lanceolate

and taper to a pointed or rounded apex. The first glume

is 0.10 to 0.18 inch (2.5 to 4.5 mm) long, the second is

0.16 to 0.22 inch (4 to 5.5 mm) long. Lemmas are 0.20

to 0.28 inch (5 to 7 mm) long, rounded, and glabrous to

roughened distally. Awns are 0.08 to 0.24 inch (2 to 6

mm) long. Plants flower from May to August (Arnow

1987, 1993; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969). Idaho fescue is very

closely related to Festuca ovina. Some authors regard

it as the native, low-elevation phase of that species and

apply the name F. ovina var. ingrata.

Ecological Relationships and Distribution

Idaho fescue ranges from Southwestern Canada

south to northern California and east from central

Montana to northern New Mexico (Tisdale 1959). It

occurs at elevations ranging from 800 to 12,000 ft (244

to 3,659 m) on sites receiving 15 to 30 inches (380 to

760 mm) of annual precipitation (Daubenmire 1966).

On north slopes and cool areas receiving added moisture it may grow where annual precipitation is as low

as 10 inches (254 mm). Idaho fescue occurs over a wide

environmental gradient and is associated with a range

of soil conditions and plant communities. It grows on

dry, well-drained, fertile, moderately deep soils ranging from clayey or silty to sandy and gravelly loams

that are slightly acidic to slightly alkaline (USDA

Forest Service 1937; Wasser 1982; Vallentine 1971). It

also occurs on shallow, rocky soils and on weakly

saline soils where its productivity is lower. Plants are

cold tolerant and moderately drought tolerant, but

they are not tolerant of high water tables or flooding.

Idaho fescue is a common or codominant species in

grasslands, sagebrush-grass, mountain mahogany,

oakbrush, mountain brush, pinyon-juniper, antelope

bitterbrush, mountain meadow, conifer, riparian, and

alpine communities (Zouhar 2000). It occurs on meadows, plains, foothills, ridgetops, mountain brush communities, and open conifer forests. The species occurs

in open areas, but it is also moderately shade tolerant.

Associated species are big sagebrush, Sandberg bluegrass, bluebunch wheatgrass, and many native forbs.

In lower elevation and drier communities, the distribution of Idaho fescue is restricted to areas with better

availability of water (Johnson 1994). Many Idaho

fescue sites have been invaded by cheatgrass (Wasser

1982).

Seedlings may emerge in fall and develop during

favorable periods in winter and spring to the point that

the plants can become dormant and survive summer

Figure 33—Idaho fescue is one of the most important cool-season bunchgrasses in the West (RMRS

photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 387

Chapter 18 Grasses

drought (Daubenmire 1975b). Idaho fescue begins

growth in March or April, and seeds mature in mid to

late summer depending on location and altitude. Plants

become dormant during summer, but resume growth

in fall if adequate moisture is received (Wasser 1982).

Reproduction is by seed and tillers (Stubbendieck

1992). Plants produce shallow, fibrous root systems.

Goodwin and others (1996) found that about 40 percent of the root biomass occurs in the upper 4 inches

(10 cm) of soil. Roots are associated with vesiculararbuscular mycorrhizae (Goodwin 1992; Ho 1987).

Idaho fescue can be damaged by wildfires at any

season (Smith and Busby 1981; Wright and others

1979). Cool fires burning through stands with little

litter buildup may cause minimal damage. However,

the fine leaves and accumulation of litter can result in

prolonged high-temperature burning that may kill or

damage the root crown (Agee 1996), with the extent of

damage dependent upon fire conditions, fire history of

the site, extent of grazing, and other factors (Bunting

1984; Wright and others 1979). Rate of postfire recovery varies considerably among burns and depends on

the degree of plant damage and environmental factors

(Beardall and Sylvester 1976; Harniss and Murray

1973), with response to season of burn variable. Surviving plants recover through tillering if soil moisture

permits. Both protein content and digestibility of Idaho

fescue increase temporarily following burning (Beardall

and Sylvester 1976; Singer and Harter 1996). Fires

near the margins of the species range are damaging at

any season (Blaisdell 1953; Wright and others 1979);

fires in dry environments tend to reduce productivity

for longer periods because the shallow root systems

are heavily damaged. Seedling establishment is reduced following burns, likely due to reduction of the

seed bank when temperatures exceed 257 ∞F (125 ∞C)

for more than 5 minutes (Warg 1938). It is not clear

whether early- or late-season burning best protects

Idaho fescue plants, but Wright (1974) recommended

burning when species of concern are dormant and not

stressed by drought. Bunting and others (1998) found

that deferring grazing for at least 1 year following

burning improved recovery of Idaho fescue.

Plant Culture

Variable and often low seed production and viability

in wildland stands has limited the use of Idaho fescue

in wildland seedings (Ensign and others 1984; Wasser

1982). Overcoming this problem has been a major

emphasis in plant selection work with the species;

two cultivars are now available. Lodging is not normally a problem in seed fields. Harvesting is accomplished by swathing before seed shattering occurs in

early to midsummer. Seed can be combined when in

the hard-dough stage. Yields are about 300 pounds per

acre (336 kg per ha) from irrigated fields and 100

pounds per acre (112 kg per ha) from nonirrigated

fields. A debearder will remove awns; seeds are cleaned

using a fanning mill (Smith and Smith 1997). There

are about 425,000 to 460,000 seeds per pound (937,000

to 1,014,100 per kg) (Ensign and others 1984). Recommended quality standards are 90 percent purity and

80 percent germination (Wasser 1982). Seeds undergo

afterripening over a period of at least 6 months

(Goodwin and others 1996), thus germination is delayed until seeds have gone through at least one

winter. Idaho fescue can be germinated by incubating

the seeds at 50/68 ∞F or 68/86 ∞F (10/20 ∞C or 20/30 ∞C)

(16/8 hrs) (Haferkamp and McSwain 1951). Exposure

to light improves germination (Haferkamp and

McSwain 1951).

Idaho fescue is fall seeded on drier sites by broadcasting or drilling on a firm seedbed. Seed should be

planted about 0.5 inch (1.3 cm) deep. In irrigated seed

fields or on low-elevation wildland sites that receive

reliable moisture it can be spring seeded. Dry sites are

fall seeded. At high elevations it can be spring or early

summer seeded as soon as the site is open.

In seed fields, drilling into stubble or planting in

alternate rows with annual grains may establish seed

fields. Fallowing also reduces weed populations. Row

spacings of 30 to 36 inches (76 to 92 cm) are recommended for seed fields (Wasser 1982). Expected stand

life is about 5 years (Smith and Smith 1997).

Vigor of seedlings is poor to fair (Shaw and Cooper

1973; Vallentine 1971), and they are not competitive

with seedlings of more aggressive species (Hafenrichter

and others 1968). Stands require 2 to 3 years to

establish (Hafenrichter and others 1968; Plummer

and others 1968), and weeds must be controlled in seed

fields. Grazing should be restricted until stands are

established (Ensign and others 1984); grazing pressure must be carefully controlled to maintain stands.

Resting during the season or grazing on a rotational

basis may be required (Hormay and Talbot 1961).

Seedlings are susceptible to attacks by damping off

and other seedling diseases, grasshoppers, and rodents (Wasser 1982). Competitive ability increases

considerably as Idaho fescue stands mature. When

seeded on erosive sites, addition of mulch may be

required to stabilize the surface soil until plants become established (Hafenrichter and others 1968).

Uses and Management

Idaho fescue is very palatable and heavily used by

livestock and wildlife. It is a particularly valuable

species at mid to higher elevations in sagebrush

grasslands, open conifer forests, and alpine areas.
388 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Greatest use is received in spring when the forage is

succulent and palatable, but use, particularly by sheep,

declines later in the season as the forage becomes

coarse and dry. Animals will continue to use it if other

forage is not available. Plants cure well and provide

good fall and winter forage where accessible (Shaw

and Cooper 1973). The species withstands some use,

but stands are weakened or lost with excessive grazing

and trampling, particularly in early spring. Stands

maintained in healthy condition will reseed themselves. Deer use Idaho fescue in spring (Johnson 1994),

and elk use it throughout the year (Dragt and Havstad

1987; Kufeld 1973; Shaw and Cooper 1973).

Idaho fescue can be planted alone or in mixtures

with other native grasses to provide diversity. Although not deep rooted, it produces an extensive,

fibrous root system that provides soil stabilization and

ground cover on disturbances resulting from logging,

road construction, mining, and recreation (Ensign and

others 1984; Hafenrichter and others 1968). It can be

seeded in areas where tree seedlings have been planted

if the seeding rate is low to moderate (Ensign and

others 1984). Mature stands withstand or slow the

spread of invasive species (Borman and others 1990;

Hafenrichter and others 1968; Lilley and Benson 1979).

Varieties and Ecotypes

The varieties ‘Joseph’ and ‘Nezpurs’ were developed

from material collected in the Northwestern United

States and Southwestern Canada (Ensign and others

1984). They are adapted to elevations ranging from

1,000 to 9,000 ft (305 to 2,745 m) and communities

ranging from sagebrush grass to subalpine that receive 14 to 30 inches (355 to 760 mm) of annual

precipitation. Both varieties are cold and drought

hardy and moderately shade tolerant. They are best

adapted to silty to sandy loam soils that are slightly

alkaline to slightly acidic. They do especially well in

ponderosa pine and mountain big sagebrush communities on gravelly to well-drained loamy soils. Joseph

and Nespurs are seeded on summer ranges to provide

forage for livestock and wildlife and soil stabilization

on road cuts and fills, skid trails, cutover forest areas,

and recreational sites. These varieties perform well as

components of seeding mixes because they are not

overly competitive.

Joseph was selected for its greater leaf and culm

height (28 to 31 inches [72 to 80 cm]), basal leaf growth,

and good germination (Ensign and others 1984).

Nezpurs was selected for its shorter stature, 20 to 28

inches (50 to 70 cm), large seed size, productivity,

basal leaf growth, and olive-green color. Both varieties

have been seeded in the Intermountain West, especially

in Idaho, Wyoming, eastern Oregon, and western

Colorado. Nezpurs has received more use than Joseph.

Festuca ovina

Sheep Fescue or

Alpine Fescue __________________

Synonyms

Festuca brevifolia

Festuca minutiflora

Festuca saximontana

Description

Sheep fescue is a short, matforming perennial with

dense tufts of basal leaves. Culms are 0.3 to 1.6 ft (0.8

to 5 dm) tall. The tufts are 2.0 to 5.9 inches (5 to 15 cm)

tall and generally about half the height of the culms.

Sheaths are glabrous to minutely hairy. Leaf blades

are stiff, involute, and without auricles. Ligules are

scarcely 0.02 inch (0.4 mm) long, finely ciliate, and

longer on the sides than at the center. Panicles are 0.6

to 3.95 inches (1.5 to 10 cm) long, mostly strongly

compressed, but somewhat open in F. o. var. ovina.

Spikelets are 0.16 to 0.35 inch (4 to 9 mm) long and two

to five flowered. Glumes are narrowly to broadly lanceolate. The first glume is 0.08 to 0.14 inch (2 to 3.5

mm) long and one nerved. The second is 0.12 to 0.18

inch (3 to 4.5 mm) long and three nerved. Lemmas are

0.14 to 0.24 inch (3.5 to 6 mm) long, glabrous or

minutely hairy toward the tip. Awns are 0.04 to 0.20

inch (1 to 5 mm) long. Sheep fescue flowers in July

through August and disperses its seed from August to

October. Reproduction is solely by seed due to the

absence of rhizomes (Arnow 1987, 1993; Cronquist

and others 1977; Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution

Sheep fescue is a widespread, polymorphic

circumboreal species with many varieties, some of

which have been treated on the specific level. In North

America, sheep fescue is distributed southward from

the arctic region through mountainous areas to California, and east to northern New Mexico, Kansas, and

North Carolina. The species also occurs in Mexico and

South America. Plants of the two native varieties are

small and occur in rocky areas and on shallow, gravelly soils, and dry exposed sites on slopes and ridges.

They require annual precipitation of 8 to 14 inches (20

to 36 cm) or more (Arnow 1987), and may be abundant

in communities from sagebrush to pinyon-juniper,

mountain brush, aspen, conifer, and mountain grassland communities. F. o. var rydbergii is distributed

from foothill to subalpine areas and F. o. var. brevifolia

occurs in subalpine and alpine habitats (Cronquist

and others 1977).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 389

Chapter 18 Grasses

The three varieties commonly seeded are all European in origin: sheep fescue, hard sheep fescue)

(fig. 34), and sulcate sheep fescue (fig. 35). Sheep

fescue is distinguished by its open and spreading

panicle. Hard sheep fescue has leaf blades that are

smoother, wider, and firmer than those of sulcate

sheep fescue.

Sheep fescue is a long-lived, drought-hardy variety.

It grows on coarse, dry, infertile soils on sites receiving

10 to 18 inches (25 to 46 cm) of annual precipitation

(Jensen and others 2001). Sheep fescue is cold tolerant

and provides stability to surface soils. It is somewhat

tolerant of finer soils and shading. Hard sheep fescue

is seeded on sites receiving 15 inches (38 cm) or more

Figure 34—Hard sheep fescue plants are upright,

leafy, and slightly taller than those of sulcate sheep

fescue (RMRS photo).

Figure 35—Sulcate sheep fescue grows as distinct fine-leaved clumps (RMRS photo).

of annual precipitation (Wasser 1982) to stabilize soils

and revegetate disturbances. It is used for roadway

plantings, airports, and other sites where a competitive, low-growing ground cover is required. Although

seedlings are slow to establish, plants are long lived

and become competitive as they develop through the

growth of abundant fibrous roots. Forage production is

low, but of fair to good quality (Hassell and others

1983). Sheep fescue is adapted to shallow, moderately

to well-drained soils with pH 5.5 to 7.5. Plants are heat

tolerant.

Plant Culture

Sheep fescue seeds ripen in early fall (Sampson

1924). They can be combined directly or first windrowed and dried for 5 to 7 days (Smith and Smith 1997;

Wheeler and Hill 1957). Lodging is generally not a

problem, but seeds begin shattering after reaching the

hard-dough stage (Smith and Smith 1997). Seed is

cleaned using a debearder and fanning mill. An indent

cylinder may be used to increase purity. Seeds are not

generally harvested during the establishment year.

Stand life is 4 to 5 years. There are about 500,000 seeds

per pound (1,100,000 per kg) (Smith and Smith 1997).

Seed fields can be planted in fall or spring (Jensen

and others 2001; Plummer and others 1968). Seeds are

easily handled. They should be planted about 0.25 to

0.5 inch (0.6 to 1.5 cm) deep (Horton 1989; Plummer

1943) on a firm seedbed. Rows should be 12 or 24

inches (30 to 60 cm) apart in irrigated fields and 36

inches (90 cm) apart on drylands (Smith and Smith

1997). Wildland sites are drill seeded or broadcast.

Dry sites are best seeded in fall. More mesic areas can

be spring seeded. Seedlings are not highly vigorous,

and establishment occurs over a 2- to 3-year period as

plants develop extensive fibrous root systems. On

wildland sites, sulcate sheep fescue is more compatible with other species than hard fescue and can be

included in mixtures. Plants are not tolerant of highly

saline or alkaline soils (Jensen and others 2001); they

are useful on dry, sandy, or rocky sites (USDA Forest

Service 1937) where other components of the mixture

may not establish well. Mature plants are persistent

due to development of an extensive fibrous root system. Plantings reduce the spread of weeds and are

tolerant of grazing if properly managed. Excessive

grazing can damage the stands; recovery is slow (Jensen

and others 2001).

Uses and Management

Sheep fescue has been seeded successfully throughout the Intermountain West in mountain big sagebrush and mountain brush communities and in openings in aspen and subalpine communities that receive

14 inches (355 mm) or more of annual precipitation. Its
390 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

seedlings are slow to establish, but tenacious. The

extensive root system of mature plants makes them

competitive and fairly resistant to drought and trampling. Sheep fescue greens up early in spring and

receives some use by livestock, particularly sheep.

Grazing should be regulated to leave a 2- to 4-inch (5

to 10 cm) stubble (Jensen and others 2001).

Sheep fescue is used for postfire seedings, mined

land reseedings, recreation areas, ditchbanks, and

other disturbances in mountainous areas to reduce

soil erosion. Baron (1962) found conifer seedling survival was not decreased on areas planted with sheep

fescue relative to controls. Sheep fescue is used in lawn

seed mixtures and is particularly useful on infertile or

sandy soils.

Varieties and Ecotypes

‘Covar’ sheep fescue originated near Konya, Turkey

(Alderson and Sharp 1994). A fine-leaved fescue, it

was selected for erosion control and revegetation of

disturbed sites and depleted communities. It is adapted

to shallow, gravelly, and well-drained soils that receive at least 8 inches (203 mm) of annual precipitation (Wasser 1982). Covar has been seeded successfully in mountain big sagebrush, upper mountain

brush, ponderosa pine forests, and aspen openings. It

requires fall seeding, and it is slow to establish. Mature plants are persistent, competitive, winter hardy,

and drought tolerant. The variety is competitive and

will spread to dominate a community.

‘Durar’ hard fescue, a larger plant of European

origin (Alderson and Sharp 1994), is adapted to welldrained, basic to moderately acidic soils of pinyonjuniper, mountain brush, ponderosa pine, aspen, and

conifer forests receiving 14 to 28 inches (300 to 700

mm) of annual precipitation. The variety was developed for soil stabilization and forage production. Durar

is somewhat slow to establish, but mature stands

spread aggressively and outcompete most native and

some introduced grasses, forbs, and shrubs.

Hilaria jamesii

Galleta or Curly Grass ___________

Synonyms

Hilaria sericea

Description

Galleta is a strongly rhizomatous or stoloniferous

perennial warm-season grass with a dense, fibrous

root system. Rooting depth is about 12 inches (30 cm).

Culms are 0.5 to 2.1 ft (1.5 to 6.5 dm) tall and grow

horizontally along the ground before turning upward

(fig. 36). Nodes of the culms are often finely hairy.

Leaves are mostly basal with hairless to minutely

roughened sheaths, and a few shaggy hairs at the

throat. Leaf blades are short and curly, flat to folded,

0.06 to 0.12 inch (1.5 to 3 mm) wide, 0.8 to 2 inches (2

to 5 cm) long, and involute toward the tip. They are

rigid, hairless to roughened, often waxy, and sometimes covered with small stiff hairs. Ligules are 0.04 to

0.10 inch (1 to 2.5 mm) long with lacerate, long-ciliate

margins. Spikes are 0.08 to 0.28 inch (2 to 7 mm) in

length with long, shaggy hairs at the base of the

spikelet clusters. Glumes of the lateral spikelets are

subequal, roughened, and 0.20 to 0.33 inch (5 to 8.5

mm) long with an irregularly lacerated apex. The first

glume is asymmetrical with the midnerve displaced to

one side; awn length is 0.12 to 0.22 inch (3 to 5.5 mm).

The second glume is awnless, or very short awned.

Glumes of the central spikelet are subequal and 0.16

to 0.26 inch (4 to 6.6 mm) long with stiff hairs. The

nerves in the glumes extend into an irregular awn that

measures 0.06 to 0.28 inch (1.5 to 7 mm) in length.

Lemmas of the lateral and central spikelets are 0.18 to

0.35 inch (4.5 to 9 mm) long, lanceolate, and blunt

tipped. They are hairless basally, roughened apically,

and three nerved. The single lemma of the central

spikelet is narrowly lanceolate and bears a dorsal awn

from below the notched apex (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969). Galleta is sometimes referred to in

older literature as black grama. This common name is

misleading, giving the impression that it belongs to

the genus Bouteloua. Galleta flowers from May to

August. Chromosome number is 2n = 2x = 36 or 38

(Arnow 1987).

Figure 36—A strongly rhizomatous or stoloniferous

species, galleta stems grow along the ground before

bending upward (RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 391

Chapter 18 Grasses

Ecological Relationships and Distribution

Galleta is the most northerly distributed species of

its genus (Heizer and Hassell 1985). Its range extends

from southeastern California and southern Nevada

across Arizona and southern Utah to Wyoming, New

Mexico, Kansas, Texas, and the Oklahoma Panhandle

(Heizer and Hassell 1985). It grows from dry desert

scrublands of lower valleys to mesas, plains, and

pinyon-juniper woodlands of foothills. Galleta often

grows in association with blue grama in sagebrush

areas, often as the dominant species. It occurs from the

valleys to the lower ponderosa pine zone at elevations

from 4,000 to 8,000 ft (1,220 to 2,439 m) (USDA Forest

Service 1937; Wasser 1982). It is particularly common

and frequently dominant on the plains of New Mexico.

It grows with shadscale on salt desert shrub sites in

the Great Basin and on sandy and rocky sites in the

Colorado Plateau. It also occurs in creosote bush and

desert shrub communities in the Southwest.

Galleta grows on sites receiving 6 to 18 inches (150 to

460 mm) of annual precipitation with soils ranging

from coarse to fine and pH of 6.6 to 8.4 (Bridges 1942;

Horton 1989; USDA Natural Resources Conservation

Service 2003; Stefferud 1948). Plants have a low requirement for soil fertility (USDA Natural Resources

Conservation Services 2003); tolerance of salinity and

alkalinity varies among populations. Plants are

drought and cold tolerant, but not shade tolerant.

Galleta is moderately fire tolerant and usually recovers by the second year following burning (Arnold and

others 1964).

Plant Culture

Galleta initiates growth in spring if adequate moisture is received (Wasser 1982). If summer rains occur,

it may mature two seed crops. The first crop can be

harvested in June and the second in August to October. Many florets in wildland stands do not produce

seed, thus costs of wildland collections are high. The

release, ‘Viva’, was selected for its high percentage of

seed-bearing florets (Heizer and Hassell 1985). Native

and domestic seed fields can be harvested by combining (Wasser 1982). Seed is cleaned by hammermilling

and fanning. Recommended purity is 20 to 40 percent

and germination 80 to 98 percent for seed sales. There

are about 170,000 seeds per pound (374,780 per kg).

Seeds do not require prechilling. They can be incubated at 68/86 ∞F (20/30∞C) (16/8 hrs) to test germination (Haferkamp and McSwain 1951). Exposure to

light is required. Seeds are nondormant and will

germinate under a range of constant and alternating

temperatures (Knipe 1967).

Seed should be planted during the most reliable period

of rainfall, usually early summer (Link 1993). Seed is

planted about 0.5 to 1 inch (1.3 to 2.5 cm) deep using drills

designed to plant chaffy seed. Seed may be broadcast, but

it must be covered. Light irrigation helps to increase

germination and establishment in seed fields.

Wildland stands may be established using water

catchment techniques such as deep furrow drilling to

aid stand establishment on dry sites. Rate of seedling

development is slow to moderate and stand establishment may require 2 years or more. Seedlings are

drought and cold tolerant, but they are susceptible to

attacks by rodents and grasshoppers (Wasser 1982).

Galleta is adapted to fine, sometimes saline soils that

limit seedling establishment of many species. It can be

seeded in mixtures with other species adapted to such

sites. Grazing should be avoided during one or more

growing seasons to permit stand establishment (Merkel

and Herbel 1973).

Uses and Management

Galleta produces considerable nutritious forage and

withstands heavy grazing and drought. It is palatable

in pure stands, but receives less use when mixed with

more palatable grama grasses. When grazed heavily,

such mixtures gradually shift toward a dominance of

galleta. Galleta provides good, moderately palatable

forage for livestock and wild ungulates during the

summer growing season. Regrowth is good following

summer rainfall, but regrowth following fall rains or

frost is more limited. Dormant plants are dry, tough,

and unpalatable (Heizer and Hassell 1985; Vallentine

1971; West 1972). Plants receive little use in late fall

or winter. Although galleta is resistant to grazing,

excessive use results in reduced forage and rhizome

production (Heizer and Hassell 1985).

Galleta is seeded to provide forage, soil stabilization,

and watershed protection (Wasser 1982). Spread from

seed is slow, but the woody rootstocks that sometimes grow to 6 or 7 ft (1.8 to 2.1 m) in length permit

vegetative spread. They also protect plants from

trampling damage and hold soils in erodible areas.

Plants are bunchy and do not form a continuous sod.

Galleta is extremely drought resistant and provides a

persistent vegetative cover in adapted areas.

Varieties and Ecotypes

Viva was collected from a population near New Kirk,

NM, on a site at 5,740 ft (1,750 m) elevation that

receives an average of 9 inches (230 mm) of annual

precipitation (Alderson and Sharp 1994). It exhibits

good seedling vigor and drought and cold tolerance.

Viva provides forage for livestock and big game. It is

also used to revegetate mined lands and other disturbances. It is seeded in extreme southern Utah and

Colorado, northern Arizona, and northwestern New

Mexico. It has done well when seeded with other

warm-season grasses on the Great Plains from Mexico
392 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

to the Canadian border. It has not established or

persisted well in the Great Basin.

Koeleria macrantha

Prairie Junegrass _______________

Synonyms

Koeleria cristata

Poa cristata

Aira cristata

Koeleria gracilis

Koeleria nitida

Description

Prairie junegrass was originally and erroneously

named and described as K. cristata, based on a specimen of Poa cristata. The oldest valid name is believed

to be K. macrantha; however, Cronquist and others

(1977) prefer to use the oldest New World name, K.

nitida. Junegrass can be distinguished from Poa

fendleriana and Trisetum wolfii by its hairy panicle

axis and generally smaller spikelets. It is often found

in association with these grasses.

Prairie junegrass is a cespitose perennial bunchgrass with culms 0.6 to 2.2 ft (2 to 6.5 dm) tall. Most

leaves are basal, but a few small leaves are produced

on the culms. Sheaths are stiff haired or rarely hairless. Leaf blades are usually folded or involute, 0.04 to

0.10 inch (1 to 2.5 mm) wide, and stiff haired to

hairless. Ligules are 0.02 to 0.08 inch (0.5 to 2 mm)

long, irregularly lacerate to subentire, minutely ciliate, and sometimes sparsely short hairy. Panicles are

1.0 to 5.9 inches (2.5 to 15 cm) long, contracted, and

spikelike with short, appressed branches (fig. 37). The

spike axis is densely short hairy. Spikelets are 0.16 to

0.20 inch (4 to 5 mm) long and two to four flowered.

Spikelet axes are silky haired. Glumes are subequal

and roughened on the keel and sometimes throughout.

The first glume is 0.11 to 0.20 inch (2.8 to 5 mm) long,

lanceolate to narrowly lanceolate, and one nerved. The

second glume is 0.13 to 0.22 inch (3.2 to 5.5 mm) long,

lanceolate, and one to three nerved with the lateral

nerves barely visible. Lemmas are about as long as the

second glume, lanceolate, and five nerved with the

lateral nerves obscure. They are roughened and sometimes bear a short awn near the tip. Flowering occurs

in June and July (Arnow 1987, 1993; Cronquist and

others 1977; Hitchcock 1950; Hitchcock and others

1969).

Ecological Relationships and Distribution

Prairie Junegrass is the only species of Koeleria

native to Western North America. It ranges from

southern Ontario to British Columbia and southward

to Texas, California, and Washington. It is one of the

most common and widely distributed of the western

grasses, extending over a wide range of altitudes and

growing on a variety of dry to moist clay loam to sandy

or sometimes rocky soils with pH 6.5 to 8.0 (Hassell

and others 1983). Junegrass is an important mid- to

late-seral forage species of the sagebrush, mixed grass,

and open timber types (Coupland 1950). It requires 16

to 21 inches (41 to 53 cm) of annual precipitation, and

is usually found at elevations of 5,000 to 8,000 ft (1,524

to 2,438 m) (Parker 1975). The species does not yield a

great deal of forage, as it is low growing with abundant

but short and mostly basal leaves. Prairie junegrass

begins growth from seed or established plants very

early in the season (Simonin 2000). Seed is dispersed

from July to September. The generally low seed viability is counterbalanced by abundant production (Arnow

Figure 37—Prairie Junegrass panicles are spikelike

with short-appressed branches (photo courtesy of

John Kinney, USDA FS, Rocky Mountain Research

Station, Boise, ID).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 393

Chapter 18 Grasses

1987, 1993; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969). Plants are cold, heat,

and drought tolerant (USDA Natural Resources Conservation Service 2003).

Prairie junegrass production is improved by spring

or fall burns when adequate soil moisture is available,

and effects may be long lasting (Blaisdell 1953). Late

spring burns are most damaging (Towne and Owensby

1984). Plants are small and clumped with the growing

points near the soil surface; thus, they burn rapidly

with little heat transfer (Young 1983). Recovery is

from the crown or from seed.

Plant Culture

Native stands of prairie junegrass are collected by

hand stripping, while cultivated fields can be combined or swathed and combined (Smith and Smith

1997). Lodging and shattering are not generally problems with this species. The seeds are small, but readily

cleaned using an air-screen machine. Seed yields on

irrigated land average 150 pounds per acre (168 kg per

ha) and 75 pounds per acre (84 kg per ha) on

nonirrigated land (Smith and Smith 1997). There are

about 2,315,000 seeds per pound (5,090,000 per kg)

(Smith and Smith 1997). Blake (1935) found viability

of seed kept in dry storage decreased from 21 percent

after 3 years to 7 percent after 6 years (Blake 1935).

Prairie junegrass should be planted about 0.25 to

0.50 inch (0.6 to 1.3 cm) deep on a firm seedbed in fall.

Seed fields and upper elevation sites are planted in

early spring. Seed may be planted by drilling or broadcasting. Seed fields should be planted at 1 pound per

acre (1.1 kg per ha) in 24-inch (60-cm) rows, 2 pounds

per acre (2.2 kg per ha) in 12-inch (30-cm) rows, and

0.75 pound per acre (0.82 kg per ha) for 36-inch (90 cm)

rows (Smith and Smith 1997). Direct seeding and

seedling transplanting has produced poor results in

seed fields; however, transplanting 1-year-old plants

has been more successful (Nuzzo 1978). Commercial

growers do not grow junegrass often because they are

unable to acquire large amounts of native seed (Hassell

and others 1983). Available native seed is frequently

of poor quality, as it is generally harvested from

wildland sites. Use of locally adapted material may be

essential for reestablishment of healthy stands

(Simonin 2000). Some European material is now being

marketed in the United States.

Junegrass may be useful for reestablishing vegetation resistant to annual weed encroachment and invasion, largely due to the early spring emergence of its

seedlings. This species does well in colonizing disturbed soils, and seedling emergence and survival are

not affected by competition with Japanese brome (Romo

and Eddleman 1987; Simonin 2000). However, plugs

of prairie Junegrass were unable to compete with

medusahead, dogtail, bulbous bluegrass, ripgut brome,

or yellow starthistle (Borman and others 1991). In

irrigated seed fields the plants grow relatively slowly

compared to weeds (Smith and Smith 1997). Established plants are long lived (Hassell and others 1983).

Uses and Management

All classes of wildlife and livestock utilize prairie

junegrass with greatest use early in the season (Rose

and others 1998). It provides good-quality early spring

forage due to its rapid development, but palatability

decreases during seed production (Vallentine 1961).

Prairie junegrass is a key winter food source for

animals of the British Columbian prairies (Blower

1982). Although utilized by whitetail and mule deer,

prairie junegrass is not a primary source of forage

because of its scattered distribution.

Prairie junegrass can be used to revegetate highly

disturbed areas (Redmann and Schwarz 1986). Studies of succession in the Piceance Basin of northwestern

Colorado (Biondini and others 1985) and Wood Buffalo

National Park, Alberta (Redman and Schwarz 1986),

showed that prairie junegrass increased following soil

disturbances. The species has been included in seed

mixes designed to restore disturbed fescue communities in Glacier National Park (Laycock 1967).

Varieties and Ecotypes

There are no releases.

Oryzopsis hymenoides

Indian Ricegrass ________________

Synonyms

Achnatherum hymenoides

Stipa membranacea

Stipa hymenoides

Oryzopsis membranacea

Description

Indian ricegrass is a densely tufted perennial bunchgrass with a deep, fibrous root system. Culms are

hollow, thick walled, and 0.7 to 2.8 ft (2 to 8.5 dm) tall.

Leaf sheaths are persistent and glabrous to minutely

hairy. Old sheaths are fibrous and papery, sometimes

becoming partly buried on sandy sites. Ligules are

0.10 to 0.30 inch (2.5 to 7.5 mm) long, entire to lacerate

along the margins, and abruptly tapering at the apex.

Leaf blades are smooth, strongly involute, about

0.04 inch (1 mm) wide, and nearly as long as the

culms. Panicles are 2 to 7.5 inches (5 to 19 cm) long

with the branches and hairlike branchlets paired,

lacey, and spreading widely at maturity. Glumes are

ovate, fine hairy to nearly hairless, translucent along
394 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

the margins, and tapering to a point or short awn up

to 0.08 inch (2 mm) long. The three or sometimes five

nerves are prominent near the base, but become indistinct toward the apex. Length of the first glume is 0.18

to 0.31 inch (4.5 to 8 mm), and the second is 0.17 to 0.30

inch (4.2 to 7.5 mm). Lemmas are 0.10 to 0.20 inch (2.5

to 5 mm) long, subglobose to fusiform, hard, thickened,

shiny, and dark brown or nearly black when mature.

They are densely silky haired with the hairs nearly

exceeding the glumes. Awns are 0.12 to 0.22 inch (3 to

5.5 mm) long, straight, and readily deciduous. Paleas

resemble the lemmas (Arnow 1987, 1993; Cronquist

and others 1977; Hitchcock 1950; Hitchcock and others 1969). Plants flower from May through August.

Indian ricegrass is largely self-fertile, particularly in

hot weather. Its chromosome number is 2n = 2x = 48

(Johnson and Rogler 1943). Indian ricegrass is the

state grass of Utah and Nevada (USDA Natural Resources Conservation Services 2002). It was named for

its large, high protein seeds that were used by Native

Americans to make flour.

Arnow (1987) transferred Indian ricegrass from

Oryzopsis to Stipa based on similarities in embryological characteristics and floral structures and natural

hybridization between the two genera. More recently,

Barkworth placed the species in Achnatherum

(Barkworth and Everett 1987; Hickman 1993).

Ecological Relationships and Distribution

Indian ricegrass is one of the most common grasses

on semiarid and arid lands of the West. It is distributed across Western North America from British Columbia to southern California and Mexico and eastward to Manitoba, the Dakotas, and Texas (Arnow

1987; Stubbendieck and Jones 1996). It is one of the

most drought-tolerant native range grasses (USDA

Forest Service 1937), occurring on sand dunes and dry,

sandy soils of valley bottoms where it often forms

dense stands. Indian ricegrass is a dominant or associated species in creosote bush, salt desert, big sagebrush, black sagebrush, pinyon-juniper, ponderosa

pine, and mixed-grass prairie communities (Jensen

and others 2001; Plummer and Frischknecht 1952;

Young and Evans 1984). It also grows scattered among

other vegetation on high grassy plains, foothills, exposed ridges, and dry, sandy or rocky mountain slopes.

Indian ricegrass can be found at elevations ranging

from 2,000 to 10,000 ft (600 to 3,000 m) (Booth and

others 1980; Wasser 1982) on sites receiving 6 to 16

inches (15 to 40 cm) of annual precipitation (Natural

Resources Conservation Service 2002). Although it

generally occurs on sandy soils on drier sites, it is

found on soils ranging from sandy to heavy clays

where precipitation is greater (Jones 1990; Platou and

others 1986).

The growth cycle of Indian ricegrass is keyed to

survival in dry and uncertain environments. Growth

of mature plants is initiated at low temperatures in

early spring (Pearson 1979). Leaves and flowers develop rapidly, and good seed crops mature prior to

summer drought. Flowering may continue into summer in favorable precipitation years (Jones and Nielson

1989; Stoddart and Wilkinson 1938). Plants adapt to

unstable sandy soils by developing adventitious roots

from elongating basal internodes (Stoddart and

Wilkinson 1938). Fungal hyphae are associated with

the root system (Robertson 1976), and the rhizosheaths

fix nitrogen (Wullstein 1980).

Indian ricegrass is fire tolerant when dormant. Spring

and early summer burns may be more damaging, but

the effects of fire on the plant and its response to fire

are not well documented. Recovery occurs through

resprouting, tillering, or germination of seeds remaining in the soil seed bank or arriving from offsite

(Everett 1987; West 1994; Wright 1985; Wright and

others 1979).

Plant Culture

Rodent caching and burial of seeds by blowing sand

provide appropriate microsites for natural regeneration (McAdoo and others 1983; Smigelski 1968). Long

afterripening requirements may be met as seeds endure drought periods, permitting development of stands

during wet periods (Jones and Nielson 1989).

Seed production of native stands varies by year and

is dependent upon local weather conditions. In addition, flowering and seed ripening are indeterminate

and seed dispersal is rapid (Robertson 1977). Consequently, only a portion of the seed crop may be harvested on any collection date. Narrow glume angle has

been used as a factor for selecting genotypes that

might disperse seeds more slowly (Whalley and others

1990). Much of the seed produced under arid conditions fails to fill, but remains on the plant; thus,

production may be considerably less than anticipated

(Stoddart and Wilkinson 1938; Stubbendieck and Jones

1996). In cultivated fields with irrigation, good seed

yields are expected annually.

Seed may be harvested by hand collection from

wildland stands or by swathing and combining (Wasser

1982). Planted seed fields are combined. Harvesting

equipment such as brush harvesters for species with

indeterminate flowering and fruit ripening may permit multiple harvests, thus increasing the total annual yield (Stubbendieck and Jones 1996).

Seed is cleaned using a barley debearder or

hammermill. Air-screen machines remove the chaff.

Seed is marketed at about 95 percent purity and 11

percent germination (Wasser 1982). A high percentage of the seed is usually dormant. There are about

159,000 seeds per pound (350,000 per kg) (Smith and
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 395

Chapter 18 Grasses

Smith 1997). Seed can be stored in a warehouse for 5

years (Plummer and others 1968).

Emergence and establishment of seeded stands of

Indian ricegrass is slowed by complex seed dormancy

(Jones 1990). Mechanical restraint imposed by the

indurate lemma and palea restricts oxygen uptake.

Physiological dormancy is strongest in newly harvested seed and decreases over time. Acid treatments

and mechanical scarification have been applied to

relieve mechanical dormancy, while use of old seed

and moist prechilling treatments can reduce physiological dormancy. These treatments and combinations of these treatments, however, have not proven

satisfactory for ensuring stand establishment (Jones

1990), and they are not easily applied on a large scale.

Seed dormancy in this species is further complicated

by the presence of seed polymorphism (Jones 1990).

The presence of two or more likely heritable seed types

with differing germination requirements within a population serves as a natural survival mechanism. Seed

polymorphism, however, also increases the difficulty

of establishing uniform seeded stands.

Seeds should be fall planted in areas receiving primarily winter precipitation to provide exposure to

cool, moist winter conditions that serve to reduce

physiological dormancy (Smith and Smith 1997). Seed

should be drilled 1 to 3 inches (2.5 to 7.6 cm) deep, or

broadcast and covered to this depth. Seed may be

planted up to 4 inches (10.2 cm) deep in very sandy

soils (Wasser 1982). If pure stands are drill seeded, 5

to 10 pounds of pure live seed per acre (5.6 to 11.2 kg

PLS per ha) should be applied (Allison 1988; Pawnee

Buttes Seed, Inc. 2002). The rate should be increased

for broadcasting. Natural variation in dormancy among

seeds and seed types within a population often translates to erratic stand development.

Early growth is slow, and good weed control is

required in agricultural fields to permit establishment

of a good stand. Wildland seedings should not be

conducted in areas where dense stands of cheatgrass

or other weeds are likely to create considerable competition. Indian ricegrass should not be drill seeded with

highly competitive species. Stands should be protected from grazing until they are well established

because younger plants are easily uprooted. Stands

develop slowly, but they are persistent once established (Wasser 1982). Wildland seedings spread well

from seed if not grazed excessively (Plummer and

others 1968).

Uses and Management

Indian ricegrass is highly palatable to wildlife and

livestock (Beale and Smith 1970; Kufeld 1973; Lauer

and Peek 1976). Plants are productive and the herbage

cures well, providing a good energy source in winter

(Cook and others 1954; Stubbendieck and others 1985;

Young 1989) (fig. 38). Protein content of dry material

is low during this period (Welch 1981), but some

material near the base of the plants may remain green

and more nutritious. The species is extremely important on winter ranges, such as salt desert shrublands

where extensive stands occur (Young 1989). It is also

valuable in late winter and early spring because it

initiates new growth sooner than many associated

species (Green and others 1951; Hutchings and Stewart

1953; Quinones 1981). Indian ricegrass is moderately

resistant to grazing, but stands have been lost as a

result of excessive use (Rogler 1960; Stubbendieck and

others 1985). Deferment of livestock use in early

spring may be required in some years to permit recovery of plants that have been grazed excessively and to

permit seed production and dispersal. Summer dormant plants are more tolerant of use (Stubbendieck

and others 1985).

Seeds of Indian ricegrass are large and high in

protein. Big game, livestock, rodents, rabbits, birds,

and other small animals seek them out (McAdoo and

others 1989). Rodent gathering and caching of the

seeds aids in dispersal and establishment of the

species.

Because it is an early successional species, Indian

ricegrass is frequently used to revegetate disturbed

sites such as mine sites on sandy soils. It may be used

alone or in mixtures. Although initial growth is slow,

Figure 38—Indian ricegrass is an extremely important forage species that remains nutritious through

the winter (photo courtesy of John Kinney, USDA

FS, Rocky Mountain Research Station, Boise, ID).
396 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

established plants provide canopy cover and develop

extensive root systems that bind surface soils in areas

where few other species may be adapted. Indian

ricegrass is used for low-maintenance landscaping in

recreational areas, on roadsides, and in parks and

gardens. It is often used as an ornamental and in dried

bouquets because of its attractive growth habit and

seed heads.

Varieties and Ecotypes

Seed production, seed germination, and establishment traits vary considerably among populations of

Indian ricegrass. Specific ecotypes have been selected

to provide cultivars that can be successfully cultured

(Jones 1990; Jones and Nielsen 1989). The cultivars

released to date are all nonbred, native-site populations.

‘Nezpar’ originated near White Bird, ID (Alderson

and Sharp 1994), and has been seeded extensively in

the West. It is best adapted to the Snake River Plain

and Columbia River Basin.

‘Paloma’ originated from a site near Pueblo, CO

(Alderson and Sharp 1994), at an elevation of 5,000 ft

(1,530 m) that receives 10 to 12 inches (250 to 300 mm)

of annual precipitation. Paloma is adapted to sandy to

sandy loam soils where it exhibits good stand establishment, vigor, drought tolerance, and forage production.

It is seeded to provide soil stabilization on mined lands

and other disturbed rangelands. Paloma is adapted to

basin and Wyoming big sagebrush; black, Bigelow, and

sand sage; and pinyon-juniper communities. It has

performed well in Arizona, New Mexico, and the southern portions of Utah, Nevada, and Colorado.

‘Ribstone’ was collected near Taber, Alberta (Jones

and Larsen, in press; Jones and others 2002e, 2004c),

and selected for its good seed yield, germination, and

acute glume pair angle. The latter trait has been

associated with seed retention. Ribstone is adapted to

the dry mixed grass ecoregion characterized by brown

chernozem soils in southeastern Alberta and surrounding areas. The Utah Agriculture Experiment

Station recommended it for release as a Selected

Germplasm in 2002.

‘Rimrock’ originated at 3,600 ft (1,100 m) near Billings,

MT (Alderson and Sharp 1994; Jones and Larson, in

press; Jones and others 2002e, 2004a), on a site with

fine sandy loam soil that receives 10 to 14 inches (250

to 350 mm) of annual precipitation. It is adapted to

sandy soil, but it is more persistent in medium- and

heavy-textured soils than are Nezpar or Paloma. Rimrock is adapted to the Northern Great Plains, Rocky

Mountains, Great Basin, Snake River Plain, Columbia River Plateau, and prairies of southern Alberta

and Saskatchewan. Its intended uses are for revegetation of disturbed rangelands and as a forage for livestock, big game, birds, and other wildlife species.

Phalaris arundinacea

Reed Canarygrass_______________

Synonyms

Arundo colorata

Calamagrostis variegata

Description

A perennial sodformer, reed canarygrass was first

introduced into the United States in about 1880. By

the 1930s the first selections were released for use in

the Western States. The plant is now widespread and

regarded as a widely naturalized species. Reed

canarygrass grows to 9 ft (2.7 m) in height, and it is

strongly rhizomatous. Leaf blades are flat and 0.24 to

0.70 inch (6 to18 mm) wide. Ligules are 0.12 to 0.40

inch (3 to 10 mm) long, blunt or rounded at the apex,

sometimes hairy, and usually lacerate along the margins. Panicles are 2.8 to 7.9 inches (7 to 20 cm) long,

compact, and sometimes lobed at the base. Spikelets

are ovate and three flowered with the first two florets

sterile and the third perfect. Glumes are subequal,

0.16 to 3.0 inches (4 to 7.5 mm) long, and strongly

compressed, especially toward the tip. They are keeled

and three nerved, with the lateral nerves most prominent. The keel of the glume is roughened, tapering to

a point, and usually wingless. Lemmas are shiny,

appressed hairy, and faintly three to five nerved. The

fertile lemma is 0.12 to 0.16 inch (3 to 4 mm) long,

ovate, and brownish at maturity. The two sterile

lemmas subtending the floret are subequal, 0.04 to

0.08 inch (1 to 2 mm) long, awl shaped, and hairy

(Arnow 1987; Cronquist and others 1977).

Ecological Relationships and Distribution

Reed canarygrass is a common riparian species in

wet places along ditches, streambanks, marshes, and

meadows (fig. 39). It ranges from Alaska through

Canada and the Western States to California, Nevada,

northern Arizona, northern New Mexico, and Oklahoma. It occurs in the Great Lakes area of the Central

States and eastward to North Carolina. It is also found

in Mexico and South America. Reed canarygrass tolerates frequent and prolonged flooding and submergence, but it can also withstand periods of drought. It

is best adapted to fine-textured and poorly drained

soils, and it is moderately tolerant of saline or alkaline

soils (Marquis and others 1984; Wasser 1982). It has

little shade tolerance.

Reed canarygrass has become a weedy species in

many wetland areas. It is highly productive and persistent once established. It spreads vegetatively and

from seed. It is highly competitive with timothy, Kentucky bluegrass, and redtop, often invading areas
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 397

Chapter 18 Grasses

dominated by these species and developing persistent,

monotypic stands (Apfelbaum and Sams 1987).

Plant Culture

Reed canarygrass is highly productive and has been

successfully used to plant meadows, wetlands, and

other sites with poorly drained soils. It can be seeded

in fall or early spring when there is sufficient moisture

for establishment. Seeds of reed canarygrass germinate readily, but produce weak seedlings. Consequently, satisfactory stands often fail to develop. Problems arise due to the difficulty of preparing seedbeds

on wetland or wet meadow sites. Planting sites must

be allowed to drain or dry out prior to cultivation or

seeding. Soil crusting is common on these sites and

contributes to poor seedbed conditions and planting

failures. Reed canarygrass should be seeded alone, as

it is not compatible with most other grasses. Once

established, it reproduces mainly by rhizomes that

grow into dense mats within 1 year. Up to 74 percent

of new shoots in established stands arise from rhizomes; new tillers bud within 2 inches (5 cm) of the soil

surface (Apfelbaum and Sams 1987).

Uses and Management

Reed canarygrass has been used to seed wetlands,

meadows, riparian sites, and aspen and conifer stands

with degraded understories, and disturbed sites with

poorly drained soils. It is a useful soil stabilizer and

provides an abundance of forage for cattle and big

game. Waterfowl, upland game birds, riparian mammals, and fish all use reed canarygrass for cover and

food. Reed canarygrass hay fields have been planted to

provide goose grazing areas (Burgess 1969). Prairie

chickens use reed canarygrass for hiding and thermal

cover in winter (Tester and Marshall 1962). Reed

canarygrass has been rated as good forage for cattle,

sheep, and horses in Utah, Colorado, Wyoming, and

Montana (Dittberner and Olsen 1983). It is most

palatable when it is growing, becoming more coarse

and increasing in alkaloids in autumn (Boggs and

others 1990).

Reed canarygrass can be a desirable forage crop for

cattle. Grazing should begin when the grass is 12

inches (30 cm) tall and when soils are dry to minimize

trampling (Boggs and others 1990). Intense stocking

rates with a short rotation period are recommended.

Stands should not be grazed to less than 5 to 8 inches

(13 to 20 cm) in height. Reed canarygrass is often cut

for hay, providing an acceptable winter feed for cattle.

This grass has been successfully established as an

understory with aspen and conifers. Although it has

little shade tolerance, it can still dominate these communities. It has been seeded with other perennial

grasses in aspen parks and mountain herblands, meadows, and on wet sites, but it generally dominates the

stand. Reed canarygrass has been widely seeded to

control erosion and stabilize streambanks, as it produces a spreading ground cover that provides excellent soil protection. Reed canarygrass has been seeded

in the Intermountain West on areas with a high water

table, including sites that are inundated with water

for up to 4 months. It has been planted within waterways and riparian areas from the valley floor up to and

including the conifer forest. This grass will grow in

neutral to very basic salty soils. It has not proven

useful on most wildland plantings, and is now most

widely planted as a pasture species at lower elevations. It has often been seeded in riparian communities because seeds of native species were not available,

but it gains dominance to the exclusion of other species. It should not be planted where native recovery is

desired and wildlife habitat is important.

Varieties and Ecotypes

‘Ioreed’ reed canarygrass has been seeded in the

Intermountain West on areas with high water tables,

sites that are inundated for up to 4 months each year,

and along waterways and riparian areas from the

valley floor up through conifer forests. This grass will

grow in neutral to very basic, salty soils. Ioreed was

selected for its high forage productivity and its adaptation to wet sites. It is seeded to stabilize disturbed

Figure 39—Reed canarygrass, invasive in many

North American wetland areas, forms monotypic

stands that displace native species (RMRS photo).
398 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

sites, waterways, and severely eroding areas, and to

provide forage. Ioreed can be very aggressive and

exclude other species. Other, less used, varieties include ‘Castor’ and ‘Vantage’ (Alderson and Sharp

1994). Seed of other unknown sources has been planted

widely in the Intermountain West.

Phleum alpinum

Alpine Timothy,

Mountain Timothy _______________

Synonyms

Phleum pratense var. alpinum

Description

Alpine timothy is a densely tufted perennial with

culms 0.7 to 1.5 ft (2 to 4.5 dm) tall that are often

prostrate at the base. Leaf sheaths are glabrous. Leaf

blades are flat, about 0.10 to 0.28 inch (2.5 to 7 mm)

wide, roughened along the margins and sometimes on

the surfaces. Auricles, when present, are blunt to

rounded. Ligules are 0.02 to 0.18 inch (0.5 to 4.5 mm)

long, truncate, and subentire. Panicles are small and

ovoid to cylindric when fresh. When pressed flat they

are 0.4 to 2 inches (1 to 5 cm) long and 0.28 to 0.47 inch

(7 to 12 mm) wide (fig. 40). Spikelets are one flowered

and elliptic. Glumes are subequal, 0.10 to 0.14 inch

(2.5 to 3.5 mm) long, abruptly truncate, and tapering

to stout awns that are 0.05 to 0.13 inch (1.2 to 3.2 mm)

long. Margins of the glumes are roughened; the margin of the first glume is sometimes ciliate. Lemmas are

0.07 to 0.10 inch (1.7 to 2.5 mm) long, lanceolate to

ovate, truncate at the apex, glabrous, and shiny or

sometimes fine hairy. Alpine timothy is distinguished

from timothy by the sheath of the uppermost leaf being

typically inflated. Its panicles rarely exceed 2 inches (5

cm) in length, the awns are generally greater than 0.06

inch (1.5 mm) long, and the culms are not bulblike at

the base (Arnow 1987, 1993; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution

Alpine timothy is the only species of Phleum native

to North America. It is widely distributed throughout

the cooler portions of North America and ranges from

Alaska south to New Hampshire, northern Michigan,

South Dakota, New Mexico, and southern California.

It is also found in Mexico, Chile, Patagonia, Northern

Europe, and Asia. Alpine timothy grows mainly in

alpine and subalpine areas, and it also extends down

through the spruce-fir zone and into the aspen zone. It

can be found in moist or wet mountain meadows,

parks, on streambanks, around springs, and in the

mud of bogs and marshes. It can also appear on

relatively well-drained soils on grassy slopes, dry

meadows, and occasionally in moist sagebrush areas.

In Alaska and along the northwest coast it grows at sea

level. Alpine timothy may develop dense, monotypic

stands, but in many areas, it occurs mixed with other

species, forming only a minor portion of the vegetation.

Alpine timothy grows on deep clay loam to sandy

loam soils of pH 5.0 to 7.5 that are poorly to well

drained (Hassell and others 1983), but it sometimes

occurs on shallow soils in areas receiving 16 to 60

inches (410 to 1,520 mm) of annual precipitation.

Plants grow slowly and are moderately long lived

(USDA Natural Resources Conservation Service 2003).

They reproduce from shoots developing from the base

of the plants as well as from seed. The species is cold

Figure 40—Alpine timothy is the only Phleum species

native to North America. Panicles generally less than 2

inches (5 cm) in length aid in distinguishing it from

timothy (Hitchcock 1950).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 399

Chapter 18 Grasses

and fire tolerant, fairly shade tolerant, but not drought

tolerant (USDA Natural Resources Conservation Service 2003).

Plant Culture

Supplies of alpine timothy seed are limited, as seed

is generally harvested from wildland stands. Seed

matures in August or September (USDA Forest Service 1937) and is hand harvested (Rose and others

1998). Seeds remain on the plants for a short period

before dispersal (USDA Natural Resources Conservation Service 2003). Fruits are cleaned using a

hammermill and air-screen machine. Seeds are small

at 1,680,000 seeds per pound (3,703,730 per kg) (Hassell

and others 1983) and should be planted shallowly on

a firm seedbed in fall or spring by cultipacking or by

broadcasting. Seeds can be planted on or near the

surface if the soil is moist. Germination and emergence are good if moisture is available, but seedling

growth is slow to moderate (Hassell and others 1983;

Rose and others 1998).

Uses and Management

Alpine timothy produces a fair amount of nutritious

foliage that usually remains green throughout the

summer, making it especially valuable as a lateseason feed for livestock and big game (Hitchcock

1950; Stubbendieck and others 1992). Regrowth following use is slow (Natural Resource Conservation

Service 2003).

Alpine timothy spreads slowly from seed (Natural

Resources Conservation Service 2003). The species

naturally revegetates disturbances and openings in

mountainous areas (Brown and Johnston 1978a,b). It

has performed well when tested on alpine mine wastes

in Montana. Alpine timothy has potential for use in

improving wildlife habitat, forage for livestock, roadway plantings, and in seeding mixes for disturbed

areas within its range.

Varieties and Ecotypes

There are no releases.

Phleum pratense

Timothy _______________________

Synonyms

Phleum nodosum var. pratense

Description

Timothy is a tufted perennial with a usually bulbous

base, hairless leaf sheaths, and culms 0.7 to 4.9 ft (2 to

15 dm) tall. Leaf blades are flat and 0.12 to 0.31 inch

(3 to 8 mm) wide with roughened margins and sometimes with small auricles. Ligules are 0.06 to 0.20 inch

(1.5 to 5 mm) long, subentire, rounded at the apex and

sometimes lacerate along the margins. Panicles are

0.12 to 0.63 inch (3 to 16 mm) long, 0.20 to 0.40 inch (5

to 10 mm) wide when pressed flat, and compressedcylindrical when fresh. Spikelets are one flowered and

elliptical. Glumes are subequal, 0.08 to 0.13 inch (2 to

3.2 mm) long, abruptly truncate, and awned. Keels of

the glumes are strongly and evenly ciliate. Glumes are

three nerved, with the lateral nerves close to the keel.

The first glume is slightly narrower and sometimes

hairy along the margins. Awns of the glumes are 0.02

to 0.06 inch (0.6 to 1.6 mm) long. Lemmas are 0.05 to

0.08 inch (1.2 to 2 mm) long, ovate, faintly five nerved,

minutely hairy, truncate, lacerate, and sometimes

with a small awn at the apex (Arnow 1987, 1993;

Cronquist and others 1977; Hitchcock 1950; Hitchcock

and others 1969).

Ecological Relationships and Distribution

Timothy is widely distributed in Europe and Asia in

cool, humid habitats. Introduced to North America, it

is now widespread in the northern half of the United

States and southward in mountainous areas. It is

found on moist meadows, weedy or grassy parks,

streambanks, moist canyon bottoms, open grassy

slopes, woodlands, forest openings, and roadsides and

trails from sea level to 10,500 ft (3,200 m). Timothy is

widely adapted, but grows best on deep, fertile, welldrained, moist clay, silt, or loam soils (Wasser 1982)

that are weakly basic to weakly acidic. It also grows in

sandy, gravelly, or rocky areas if the soil is moist. It

withstands winter and early spring flooding, but it is

intolerant of flooding during the growing season

(Jensen and others 2001). Timothy is not drought

tolerant, but it is cold and frost tolerant (USDA Natural Resources Conservation Service 2003). Timothy is

moderately fire tolerant, particularly when dormant.

It occurs on many wet sites and high-elevation locations that rarely burn (Wasser 1982).

Plant Culture

Timothy blooms in late spring and produces large

numbers of seeds that mature in mid to late summer.

Seeds are small and dispersed by livestock, wildlife,

wind, and other forces (USDA Forest Service 1937).

Seed are harvested by combining, and are cleaned

with an air-screen machine. There are about 1,300,000

seeds per pound (2,865,980 per kg) (Wheeler and Hill

1957). Seeds remain viable for 4 to 5 years in dry

storage (Wheeler and Hill 1957). They become highly

germinable in about 1 month following harvest, and
400 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

germination remains high for 1 to 2 years. A potassium nitrate pretreatment and a 5-day prechill with

incubation at 59/77 ∞F (15/25 ∞C) or 68/86 ∞F (20/30 ∞C)

(16 hrs/8 hrs) and exposure to light during the high

temperature periods are recommended for maximizing germination (AOSA 1999). Germination occurs

rapidly. Seed lots may be purchased with 99 percent

purity and 90 percent germination (Wasser 1982).

Timothy is seeded in fall where early spring drying

of the soil is likely to occur. Higher elevation sites are

planted in mid to late summer, while irrigated fields

are planted in late summer (Wasser 1982). Seed should

be planted about 0.25 to 0.50 inch (0.6 to 1.1 cm) deep,

or slightly deeper in coarser or drier soils (Jensen and

others 2001; Wasser 1982). Seed can be surface planted

or barely covered on fine, moist soils. The small seeds

may require addition of a diluent to permit regulation

of the seeding rate. Seeding is accomplished by broadcasting, cultipacking, or use of a drill that will place

the seeds near the soil surface.

Timothy seedlings develop rapidly when adequate

moisture is available (Cornelius and Talbot 1955;

Hassell and others 1983; Jensen and others 2001;

USDA Natural Resources Conservation Service 2003).

They are susceptible to competition and soil drying

because of their shallow root systems. Plants are

generally fully established by the second growing

season, or sometimes by the end of the first growing

season (Wasser 1982). Transplants with well-developed root systems are sometimes more successfully

used in areas where frost heaving or drying are problems (Brown and Johnston 1978a,b; Brown and others

1976).

Uses and Management

Timothy is known best for its widespread use as hay,

primarily for horses, but it is also a forage plant where

seeded on Western ranges. It is frequently grown in

combination with other cool-season grasses and with

red clover, alfalfa, and other legumes (Berg and others

1996; Wasser 1982). Timothy hay can be harvested

two to four times per season, but regrowth is slow. The

seed is used by birds. It provides cover for small

mammals, waterfowl, and other birds. It is used as a

forage by big game animals.

Livestock use timothy in spring and fall, and it

persists well if properly grazed (Wasser 1982). Timothy should not be grazed prior to emergence of the

heads. Forage quality, quantity, and palatability are

greatest in spring. Plants recover slowly following

grazing, and forage quality declines rapidly as plants

mature. Plants can spread by tillering, but they are

not resistant to heavy grazing or trampling because

the sod and bulbous bases of the plants are easily

damaged (Arnow 1987, 1993; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969).

Stands are short lived if not permitted to reseed

naturally. Established seedings are long lived when

managed properly (USDA Natural Resources Conservation Service 2003).

Timothy can be seeded where soils are moist and the

growing season is adequate for seed production. It has

been used for reseeding irrigated mountain meadows

and to provide ground cover and forage on burned,

logged, or excessively grazed mountain rangelands

where annual precipitation exceeds 40 inches (102 cm)

per year (Jensen and others 2001; USDA Forest Service 1937). An early to midseral species, timothy will

encroach and dominate on disturbed sites. It is used in

seeding mixes to stabilize roadsides, mined lands,

recreational areas, and other disturbed sites (fig. 41).

Timothy establishes by the second year after seeding.

Adapted seed sources should be used (Wasser 1982).

Timothy can be invasive and limit the regeneration

and establishment of native species. It has reduced

native grass colonization in fescue grasslands of Glacier National Park and invaded natural vegetation

there (Tyser 1992). It competes strongly with seeded

and transplanted conifers where seeded following wildfires (Baron 1962).

Varieties and Ecotypes

‘Climax’ is the most commonly seeded timothy on

rangelands and wildlands in the Intermountain West.

It is adapted to aspen, ponderosa pine, conifer, and

subalpine communities. Climax is seeded to provide

forage and soil stabilization. It can be seeded in fall or

early spring. Germination occurs quickly, and seedling establishment and vigor are good. Climax is

Figure 41—Seeded on mountainous sites to provide forage or soil stabilization, timothy limits establishment of naturally recovering or seeded species

(RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 401

Chapter 18 Grasses

compatible with natives, providing rapid soil stabilization as slower developing species establish and

mature. Stands weaken with time; their longevity is

normally 5 to 9 years. Climax will reseed itself in weak

communities and on open, disturbed sites if seed is

allowed to mature.

The variety, ‘Clair’, has been seeded recently. It

appears to perform as well as Climax on Western

rangelands. Additional varieties developed for hay

and pasture production for which seed is available are:

‘Alexander’, ‘Alma’, ‘Argus’, ‘Basho’, ‘Bottnia II’,

‘Bounty’, ‘Carola’, ‘Champ’, ‘Drummond’, ‘Dynasty’,

‘Farol’, ‘Glenmor’, ‘Hokuo’, ‘Itasca’, ‘Korpa’, ‘Mariposa’,

‘Mohawk’, ‘Nike’, ‘Richmond’, ‘Salvo’, ‘Tiiti’, ‘Tiller’,

‘Toro’, and ‘Winmor’ (Alderson and Sharp 1994).

Poa ampla

Big Bluegrass __________________

Synonyms

Poa secunda

Poa confusa

Description

Big bluegrass is a robust, long-lived, densely tufted,

hairless to minutely hairy, green or blue-waxy perennial. Mature plants develop strong, fibrous root systems. Culms are 1.3 to 5.9 ft (4 to 18 dm) tall with a

basal cluster of leaves 8 to 16 inches (20 to 40 cm) long

that generally wither early. Leaf sheaths are smooth

to roughened, generally shorter than the internodes,

and open less than one-fourth their length. Auricles of

culm leaves are often hard and conspicuous. Leaf

blades are roughened along the margins, faintly bluewaxy, 0.06 to 0.14 inch (1.5 to 3.5 mm) wide, and

usually flat, but become involute as they dry. Ligules

are short, thick, 0.04 to 0.08 inch (1 to 2 mm) long, and

truncate or more often rounded. Panicles are narrow

and elongate, 0.31 to 0.87 inch (8 to 22 mm) long,

densely flowered, and usually pale green, but sometimes tinged with purple. Spikelets are 0.22 to 0.43

inch (5.5 to 11 mm) long, three to five flowered, and

relatively narrow to somewhat cylindrical. The glumes

taper to a point, the first glume is 0.12 to 0.18 inch (3

to 4.5 mm) long and the second 0.14 to 0.22 inch (3.5 to

5.5 mm) long and about as long as the first floret.

Lemmas are 0.17 to 0.24 inch (4.2 to 6 mm) long,

rounded at the apex, and usually rounded on the back

toward the base, becoming somewhat keeled towards

the tip. Lemmas are yellowish green with translucent

margins or sometimes brownish. They are usually

hairless, but sometimes slightly hairy below. Flowering occurs from May through July. Big bluegrass often

hybridizes with rhizome-producing Poa species such

as Canada bluegrass and Kentucky bluegrass. As a

result, some specimens may have short rhizomes.

(Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969). Arnow

(1987) placed Poa ampla in the P. secunda complex.

Ecological Relationships and Distribution

Big bluegrass is found on moderately moist to dry

sagebrush slopes, aspen and forest openings, and

midelevations and, less frequently, subalpine meadows (Wasser 1982). It ranges from Alaska to North

Dakota and south from northeastern California to

New Mexico (Arnow 1987, 1993; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969).

Big bluegrass begins growth in early spring. At low

elevations it matures in late spring or early summer,

while at higher elevations or on sites with greater

moisture availability, growth may continue later into

the summer or throughout the season (Cronquist and

others 1977; Shaw and Cooper 1973; Wasser 1982).

Big bluegrass grows in areas receiving 15 to 21 inches

(381 to 533 mm) of annual precipitation, but it also

occurs in areas receiving as little as 10 inches (254

mm) (Hassell and others 1983).

Big bluegrass is adapted to a wide range of soil

textures, but it is generally found on clay loam to

sandy loam soils that may be shallow and of low

fertility. It grows on soils with pH ranging from 6 to 8

(Hassell and others 1983), and it is moderately sensitive to salinity. Plants are only weakly tolerant of early

spring flooding, high water tables, and poor drainage

(Hassell and others 1983). Local ecotypes are tolerant

of cold or heat. The species is only moderately shade

tolerant and not particularly drought tolerant. Plants

are fairly fire tolerant when dormant (Hassell and

others 1983)

Plant Culture

Seed fields are harvested using a combine or stripper or by first windrowing (Archer and Bunch 1953).

Seeds are cleaned in a fanning mill. Acceptable purity

is 90 percent and germination 70 percent (Wasser

1982). There are about 917,000 seeds per pound

(2,022,000 per kg) (Heady 1975). Seed requires a

prechill treatment of 7 days at 41 ∞F (5 ∞C) for germination (Currie 1967; Haferkamp and McSwain 1951).

Seed can be stored for 5 years (Plummer and others

1943). Seed should be planted 0.25 to 0.50 inch (0.6 to

1.3 cm) deep on medium-textured soils. It can also be

broadcast and covered. Seedings should be conducted

in fall in drier areas. Spring or early summer seedings

may be successful on sites receiving greater annual

precipitation or in irrigated seed fields (Archer and

Bunch 1953; Currie 1967; Wheeler and Hill 1957).
402 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Seedling vigor is low to moderate and stands mature

slowly. Seedlings are easily pulled up, particularly on

wet or loose soils; thus, grazing should be avoided for

two growing seasons or until the roots are securely

anchored.

Uses and Management

Big bluegrass is seeded on disturbed mountain big

sagebrush sites and high mountain meadows (Jensen

and others 2001). It is used for postfire seedings in

forested areas. It is also seeded in hay fields and

pastures with alfalfa and other grasses in both irrigated and nonirrigated fields. Such seedings may

require weed control during the first season. Big bluegrass may be seeded in mixtures with other native

grasses, forbs, and shrubs to provide diverse native

vegetation and wildlife habitat. Established stands of

big bluegrass are persistent and competitive with

cheatgrass and other weeds.

Big bluegrass greens up early in spring. It can be

grazed in spring or fall, and it is highly palatable

during these periods. Palatability declines in early

summer when moisture is inadequate. Fall rather

than spring grazing and dense seedings that favor

vegetative development are approaches for minimizing pull-up problems resulting from grazing (Hyder

and Sneva 1963). Grazing in spring should be delayed

until plants reach a height of at least 8 inches (20 cm).

Plants are tolerant of grazing, but excessive use as

plants are heading out can reduce growth during the

subsequent year (Wasser 1982). Excessive grazing

and trampling can lead to loss of big bluegrass.

Big bluegrass provides forage for wildlife and is used

by ungulates in spring and fall. Elk use it throughout

the year. At higher elevations it provides summer

forage, remaining green throughout the season if adequate moisture is available and producing good fall

regrowth. Small animals use big bluegrass as escape

and thermal cover. Birds use it for nesting cover and

consume the seeds and foliage.

Varieties and Ecotypes

‘Sherman’, collected near Moro, Sherman County,

OR (Alderson and Sharp 1994), is a large, long-lived

bluegrass that greens up very early as the snow melts.

In the Intermountain West this variety is adapted to

mountain brush communities and openings in aspen

and conifer forests. Sherman is used for soil stabilization, forage production, revegetation of disturbed lands,

and postfire seedings (fig. 42). Big game and livestock

seek it out. It is nonaggressive and compatible with

other native species.

Poa compressa

Canada Bluegrass _______________

Synonyms

Paneion compressum

Description

Canada bluegrass is an introduced, perennial, coolseason, rhizomatous sodformer that has naturalized

throughout much of the United States and Canada.

Plants are wiry, erect, and bluish green throughout.

Culms are 8 to 31 inches (2 to 8 dm) tall, strongly

flattened, prostrate at the base, and often bent at the

nodes. Stems and leaf sheaths are characteristically

flattened. Leaf sheaths are open near the base, and so

strongly compressed as to appear winged. Leaf blades

are flat to folded, and 0.06 to 0.14 inch (1.5 to 3.5 mm)

broad with short, boat-shaped tips. Ligules are to 0.02

to 0.08 inch (0.5 to 2 mm) long, fairly stiff, more or less

entire, and rounded to nearly squared at the apex.

Panicles are 1.2 to 3.5 inches (3 to 9 cm) long, compact,

dense, and usually narrow, but occasionally somewhat open (fig. 43). Spikelets are 0.12 to 0.30 inch (3 to

7.5 mm) long, crowded, and usually three to seven

flowered. Glumes are subequal and 0.07 to 0.13 inch

(1.8 to 3.2 mm) long, the first one to three nerved, the

second broader and three nerved. Lemmas are 0.08 to

0.12 inch (2 to 3 mm) long, firm, sometimes leathery,

strongly keeled, more or less rounded, and often purplish at the tip. There are fine hairs on the keel and

along the marginal nerves, especially toward the base.

Figure 42—Sherman big bluegrass can be seeded

with other natives to provide wildlife forage and

increase species diversity (photo courtesy of Scott

Lambert, USDI BLM, Boise, ID).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 403

Chapter 18 Grasses

The web is lacking, or if present, it is very scant (Arnow

1987; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969).

Ecological Relationships and Distribution

Canada bluegrass was introduced into North America

from England. It is now widely distributed throughout

the cooler regions of North America from British

Columbia southward to California and east to Georgia. In the United States, it is most common from New

England to Ohio and West Virginia. It occurs sparsely

throughout Intermountain ranges in mountain meadows and along dry streambanks, and it is occasionally

abundant on sites where it has been seeded. In the

Southwest it occurs on higher mountain ranges. It has

spread to occupy extensive disturbances throughout

its present range (Van Dyne 1958), and also grows in

irrigated lawns and pastures.

Canada bluegrass grows on soils ranging from clay

to sand and gravel. It grows well on soils with low

fertility or poor drainage (Hardy 1989; Sampson and

others 1951). Plants exhibit moderate drought and

salinity tolerance, but they are not shade tolerant

(Van Dyne 1958). Canada bluegrass and Kentucky

bluegrass occur on similar sites. Canada bluegrass

tends to dominate only on soils that are too acid,

droughty, or nutrient deficient for Kentucky bluegrass

(Johnson and Nichols 1970).

Plant Culture

Canada bluegrass reproduces from seed and rhizomes. Seeding is best accomplished in fall. There are

more than 2.5 million seeds per pound (5.5 million per

kg). Seed may be drilled or broadcast, but it should not

be planted more than 0.12 inch (0.3 mm) deep, as light

is required for germination (Fulbright and others

1982). Seedlings may emerge, but they do not persist

with more competitive species on highly fertile soils.

Seedlings do, however, compete exceptionally well on

soil with low fertility (Sampson and others 1951), but

they develop slowly, requiring 2 to 4 years to reach

maturity. Once established, Canada bluegrass can

increase its area of occurrence by natural spread, even

amid considerable competition. In the Intermountain

West, Canada bluegrass has been seeded in big sagebrush, mountain brush, aspen, conifer forests, mountain meadow parks, and riparian sites. It requires at

least 14 inches (356 mm) of annual precipitation to

establish and persist. Because seedlings develop slowly,

Canada bluegrass can be seeded with tree and shrub

species on roadways and other disturbances.

Uses and Management

Canada bluegrass has been used extensively as a

turf grass, especially in mixtures with other turf

species. It is often used to provide cover and erosion

control on roadsides, road cuts and fills, borrow pits,

dam sites, and recreational areas. It is frequently

seeded in mixtures with legumes for revegetation of

mined areas (Hardy BBT Limited 1989). It is generally

slow to establish, but mature stands provide good

cover and remain green over a long period.

Canada bluegrass is used by cattle, horses, sheep,

deer, and elk (Crawford and others 1969; Hardy BBT

Limited 1989; Kufeld 1973; Kufeld and others 1973;

Sampson and others 1951). It greens up early in

spring, and growth continues in summer and fall.

Consequently, it is palatable to livestock during this

period (Hardy BBT Limited 1989). Because it matures

later than Kentucky bluegrass and is more drought

tolerant, Canada bluegrass tends to be more palatable

in summer. It provides fair to good cover for birds and

small mammals (Dittberner and Olson 1983). This

species is resistant to heavy use and trampling and

shows good fire tolerance.

Canada bluegrass is classed as an invader on overgrazed rangelands (Van Dyne and Payne 1964). It is

generally not recommended for seeding as a pasture

grass because of its low productivity. On poor soils,

mine wastes, and roadway disturbances it may be

Figure 43—Canada bluegrass panicles are

compact, dense, and usually narrow, but sometimes open (photo courtesy of Kevin Jensen,

USDA ARS, Logan, UT).
404 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

seeded to provide useful forage (Stubbendieck and

others 1985; Welsh and others 1987) and ground

cover. Although slow to establish, this species competes well with annual weeds. It is not recommended

in seedings where native species are desired. It can

be planted to furnish cover on harsh sites, but it

usually does not support establishment of herbaceous species. Native trees and shrubs are able to

establish into existing stands of the grass, particularly in areas receiving over 20 inches (508 mm) of

annual precipitation.

Varieties and Ecotypes

‘Foothills’ Selected Germplasm, a composite of eight

Canada bluegrass accessions (Englert and others 2002),

is adapted to the Northern Rocky Mountains of the

United States and Southern Canada. It was selected

for its forage production, seed production, and ability

to spread. Recommended uses include erosion control,

critical area plantings, grazing, and low maintenance

landscaping.

‘Reubens’ originated near Reubens, ID (Alderson

and Sharp 1994). It was selected for stabilization of

low-fertility soils with an irregular moisture supply.

In the Intermountain West it is seeded extensively in

mountain brush, mountain big sagebrush, three-tip

sagebrush, and ponderosa pine communities. Reubens

establishes well on sterile, coarse-textured soils ranging from slightly acidic to slightly basic. It is seeded on

mined lands, road cuts and fills, and other disturbances to stabilize the soil. Plants are low growing and

not highly productive. Reubens is compatible with

other species, and can be seeded as a component of

mixtures.

Poa fendleriana

Mutton Bluegrass _______________

Synonyms

Eragrostis fendleriana

Poa eatoni

Poa montana

Poa longiligula

Description

Mutton bluegrass is a perennial bunchgrass that

forms large tufts 0.7 to 2.5 ft (2 to 7.5 dm) tall. Stems

tend to stool at the edge of the tufts, and there are

occasionally a few short rhizomes. Leaves are basal,

erect, numerous, and strongly blue waxy. Basal leaf

sheaths are papery, persistent, somewhat roughened,

and open nearly their entire length. Leaf blades are

firm, stiff, usually roughened, 0.06 to 0.16 inch (1.5 to

4 mm) wide, sometimes flat, but more often folded or

involute with keeled tips. Culm blades are reduced or

lacking, revealing only the ligule on the bladeless

sheath. Ligules are highly variable, often truncate,

and range from 0.02 to 0.47 inch (0.5 to 12 mm) in

length to much elongate and tapering, sometimes

exceeding 0.4 inch (1 cm) in length. Most are about

0.12 to 0.31 inch (3 to 8 mm) long. Panicles are pale to

deep purple or sometimes tawny, 0.12 to 0.6 inch (3 to

15 mm) long, and 0.4 to 0.8 inch (1 to 2 cm) wide,

oblong, contracted, and usually situated well above

the basal leaves. Spikelets are 0.20 to 0.41 inch (5 to

10.5 mm) long and tightly three to six flowered. They

are strongly compressed, keeled throughout, and often

appear papery. The first glume is 0.11 to 0.18 inch (2.8

to 4.5 mm) long; the second is 0.14 to 0.22 inch (3.5 to

5.5 mm) long. Lemmas are 0.16 to 0.28 inch (4 to 7 mm)

long, blunt on the usually irregularly lacerated apex,

and with long, silky hairs on the prominently nerved

keel and marginal nerves. Specimens of mutton bluegrass from the Intermountain area are generally pistillate, posing doubt that fertilization actually takes

place. Some plants produce a few staminate or perfect

florets (Arnow 1987) (fig. 44). Plants flower from May

Figure 44—Mutton bluegrass flowers from May

to August. Flowers are generally pistillate, but

some plants produce a few staminate florets

(RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 405

Chapter 18 Grasses

to August (Arnow 1987, 1993; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969).

Muttongrass forms a highly polymorphic complex

extending across a broad elevational range and occupying diverse habitats (Arnow 1987). Soreng (1985)

described the species as including three subspecies

differing in production of rhizomes, ligule characteristics, and reproductive biology. Goodrich and Neese

(1986) consider the species highly variable, but suggested the subspecies be reduced to varieties. Although most plants are pistillate, they found a limited

number of staminate plants that produced long

rhizomes.

Ecological Relationships and Distribution

Mutton bluegrass is one of the most widely distributed native bluegrass species in the West (Wedin and

Huff 1996). It is distributed from southeastern British

Columbia to Manitoba and south to California, Northern Mexico, western Texas, Colorado, and western

South Dakota. In the southern part of its range it

occurs from the pinyon-juniper zone through oakbrush,

mountain brush, ponderosa pine, and aspen communities to elevations of 7,000 to 12,000 ft (2,134 to

3,658 m) on sites receiving 10 to 22 inches (254 to

559 mm) of annual precipitation (Link 1993; USDA

Natural Resources Conservation Service 2003). It appears mainly on mesas, ridges, and slopes, and in open

timbered areas, as well as in well-drained parks and

meadows. In the northern portion of its range it occurs

at lower elevations in the sagebrush zone where it

grows with other cool-season grasses and forbs. It

occasionally occurs in desert shrub communities

(Arnow 1987), but it may have been lost from drier

areas as a result of overgrazing.

Mutton bluegrass is best adapted to clay loam soils,

but will grow on well-drained, medium- to coarsetextured soils that are fairly neutral (USDA Forest

Service 1937). In oakbrush, mountain brush, and

aspen communities it grows as an understory species

on deep soils and tolerates some shade. In sagebrush

and pinyon-juniper communities it may occur on shallow and gravelly soils in addition to soils with moderately well-developed surface horizons (USDA Forest

Service 1937; Dittberner and Olsen 1983).

Mutton bluegrass is an important late successional

species in many shrub-dominated communities (Koniak

1985). It is generally tolerant of fires (Arnold 1950).

Although Koniak (1985) found some stands were damaged by wildfires, Gartner and others (1978) determined that growth of rhizomatous population was

actually stimulated by fire. Plants have responded

well following fire in sagebrush, pinyon-juniper, and

mountain brush woodlands in central and eastern

Utah.

Plant Culture

Muttongrass seed matures between late May and

mid-August (Sampson 1924; Smith and Smith 1997).

Seeds can shatter rapidly at maturity (Smith and

Smith 1997). Seed stocks remain erect at maturity and

extend well above the leaves. Fields can be direct

combined as lodging is generally not a problem. Yields

in irrigated seed fields vary considerably from year to

year, averaging about 35 pounds per acre (39 kg per

ha) and ranging from 25 to 75 pounds per acre (28 to

84 kg per ha) (Smith and Smith 1997).

Seeds can be cleaned to some extent during combining. Further processing using a debearder may be

required to complete separation of seed from the debris. Trash is removed with an air-screen machine.

There are about 890,000 seeds per pound (1,958,000

per kg). Germination tests can be conducted by incubating seeds at an alternating temperature regime of

68/86 ∞F (20/30 ∞C) (16 hrs/8 hrs) with exposure to light

during the high-temperature alternation (Haferkamp

and McSwain 1951). Seed can be kept in dry, cold

storage for up to 5 years (Smith and Smith 1997).

Mutton bluegrass seed fields can be planted in early

spring. Establishment may require 2 years, and weeds

must be controlled to reduce competition. Stand life is

about 8 years (Smith and Smith 1997).

Wildland seedings should be conducted in fall. Spring

plantings may be attempted, as emergence can occur

rather rapidly. However, spring plantings can be lost

when soil surfaces dry quickly in spring. Seed can be

planted by drilling if seeds are placed near the soil

surface. Seed may also be broadcast and covered or

seeded with a cultipacker. Mutton bluegrass seedlings

are generally quite small when they enter dormancy

during the first growing season, but they begin growth

early in the second growing season. Growth rate and

lifespan are both moderate (USDA Natural Resources

Conservation Service 2003). Established stands spread

slowly from seed, and generally do not interfere with

the establishment of other native species.

Uses and Management

Mutton bluegrass begins growth in early spring and

is grazed by livestock and wildlife. Palatability decreases as the plants mature, but they retain some

green leaves and are grazed late in the season when

other forage is not available. Plants normally green up

in the fall and offer good-quality forage at this period.

Mutton bluegrass persists well in openings and in the

understory with oak and other shrubs, furnishing

green forage over a long period. It is also an important

species within the sagebrush zone. Mutton bluegrass

is considered an excellent species for all classes of

domestic livestock. It provides forage for deer and elk
406 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

in spring, and is also used in winter when the foliage

is dry (Judd 1962). Pronghorns use the seedheads in

winter (Ferrel and Leach 1950). Mule deer and elk use

stands of this species in areas where pinyon and

juniper trees have been cleared. These stands can

draw game animals from nearby cultivated fields,

thus reducing crop damage. Stands can, however, be

reduced by excessive grazing. The seeds and foliage

are used by birds (Christensen 1958; Harper and

others 1958; Martin and others 1951).

Mutton bluegrass produces a fibrous root system

about 10 inches (25.4 cm) deep that provides good

surface erosion control. Like Sandberg bluegrass, this

species is drought tolerant. Its growth rate and lifespan

are both moderate (USDA Natural Resources Conservation Service 2003). Seed crops vary among years,

and natural spread can be slow. Good recovery has

been observed on sites protected from grazing.

Mutton bluegrass has not been widely used to restore disturbed areas, but it could be useful in efforts

to reestablish native diversity. The species could be

used in many areas where introduced grasses are now

planted. It is key in areas where pinyon and juniper

encroachment has depleted the understory and seasonal habitat for big game has been lost. It is also an

important species to develop for use in seeding big

sagebrush communities where the native understory

has been lost and annual weeds have become a major

problem.

Varieties and Ecotypes

There are no releases.

Poa pratensis

Kentucky Bluegrass _____________

Synonyms

Paneion pratense

Poa peckii

Poa agassizensis

Description

Kentucky bluegrass is considered a European introduction by many botanists, but its occurrence in remote areas throughout the Intermountain region suggests that there may be native populations. Some

publications treat Poa agassizensis as the native counterpart of Kentucky bluegrass. They are differentiated

as follows: Poa agassizensishas waxy-blue leaves with

blades 0.03 to 0.08 inch (0.8 to 2 mm) wide, spikelets

mostly two flowered, and lemmas only slightly webbed

as opposed to highly webbed at the base. Kentucky

bluegrass has bright green leaves, blades 0.08 to 0.12

inch (2 to 3 mm) wide, spikelets mostly three flowered,

and lemmas heavily webbed at the base.

Kentucky bluegrass is a leafy, long-lived perennial

that spreads by seed and especially by rhizomes. It can

form a dense sod, especially on moist, fertile soils, even

when grazed or clipped. Most roots are within 3 inches

(7.5 cm) of the soil surface. Leaf sheaths are smooth,

occasionally somewhat roughened, and closed for about

half their length. Blades are soft, usually green or

sometimes slightly bluish, flat or more commonly

folded, and about 0.04 to 0.18 inch (1 to 4.5 mm) wide

with roughened margins and boat-shaped tips. Ligules

are shorter than wide, squared, and mostly entire. The

inflorescence is an open, pyramidal panicle, 2 to 8

inches (5 to 20 cm) long. Panicle branches are spreading or ascending and whorled in groups of three to five.

Spikelets are 0.12 to 0.24 inch (3 to 6 mm) long, ovate,

congested, and two to four flowered. They are green to

purplish in color and strongly compressed. Glumes are

roughened on the keel; the first glume is one nerved

and the second is three nerved. Lemmas are three to

five nerved, with weak lateral nerves. They are rounded

or tapered to a point apically and there are cobweb-like

hairs at the base (Arnow 1987; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969;

USDA Forest Service 1937).

Ecological Relationships and Distribution

Kentucky bluegrass was one of the earliest grass

introductions to arrive in the United States. Plantings

in North America were reported as early as 1685

(USDA Forest Service 1937). Kentucky bluegrass is

now widely distributed throughout most of North

America, growing in every State and Canadian Province. It is adapted for growth in cool, humid climates,

and is most prevalent in the Northern United States

and Southern Canada. It is not common in the Gulf

States nor in the desert regions of the Southwest

(Wheeler and Hill 1957). Kentucky bluegrass is not

widely adapted to arid regions receiving less than16

inches (40 cm) of annual precipitation. The grass,

however, has spread into many wildland communities. Kentucky bluegrass can be found on prairies and

in cultivated fields, mountain grasslands, mountain

brush communities, mountain meadows, riparian

drainages, and open forests. It is common along roadsides and in waste places.

Kentucky bluegrass has proven well adapted to

many mountain brush, aspen and conifer communities, and wet or semiwet meadows in aspen and subalpine communities. It dominates riparian areas over a

wide range of elevations. Considerable variability has

been noted among collections of this species. Although

Kentucky bluegrass is generally best suited to areas

receiving more than 16 inches (40 cm) of annual
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 407

Chapter 18 Grasses

precipitation, it has sometimes persisted when seeded

in more arid sites. Vallentine (1961) noted that the

species is not shade tolerant, but numerous plantings

in forested communities have survived. Although Kentucky bluegrass grows best in full sunlight, it will

tolerate light shading if moisture and nutrient conditions are favorable (Hardy BBT Limited 1989; Smoliak

and others 1981).

Kentucky bluegrass does not produce vigorous seedlings. Plants persist once established, but stands are

often slow to develop. Once established, however, the

species can be aggressive, spreading by development

of strong rhizomes. It often invades and suppresses

populations of other useful species. Kentucky bluegrass produces abundant seed that establishes in

openings within the community. It often dominates in

heavily grazed meadows, pastures, riparian sites, and

other disturbances.

Kentucky bluegrass grows on a wide variety of soils,

but thrives on well-drained loams or clay loams rich in

humus (USDA Forest Service 1937), including soils

derived from limestone (Hardy BBT Limited 1989;

Smoliak and others 1981; USDA Forest Service 1937).

Optimal soil pH is between 5.8 and 8.2 (Smoliak and

others 1981). This species requires fertile soils and

exhibits a high nitrogen requirement at all stages of

growth.

Plant Culture

There are 2.1 to 2.2 million seeds per pound (4.6 to

4.8 million per kg). Germination is generally high,

ranging from 75 to 94 percent. Seed has been drilled or

broadcast seeded in fall, but it should not be planted

more than 0.20 inch (5 mm) deep because exposure to

light is required for germination (Fulbright and others 1982). If used, this species should be seeded in

limited amounts in mixtures with other species due

to its competitive ability. Kentucky bluegrass establishes well from broadcast seedings on exposed soils. It

has been widely seeded by aerial application on rangelands and over mountainous terrain. It is also broadcast seeded on loose soils, including mine spoils and

roadway disturbances. Mine wastes and other exposed

substrata must be fertilized to assure seedling establishment. Seeded stands are slow to develop. Once

established, however, Kentucky bluegrass is highly

persistent and spreads from seed and rhizomes.

Uses and Management

Because of its sodforming ability, Kentucky bluegrass has been widely used as a ground cover. It is one

of the first grasses to resume growth in late winter and

to green up in early spring. It is highly palatable in the

early stages of growth and provides nutritious forage

for all classes of livestock and wildlife. In the West, it

is often abundant in mountain grasslands, moist and

dry mountain meadows, aspen parklands, open ponderosa pine forests, and in riparian areas where it

receives heavy use by domestic sheep and cattle (Bowns

and Bagley 1986; Clary 1975a; Kauffman and others

1983). Mountain meadows dominated by Kentucky

bluegrass may be relatively limited in extent, but they

are highly productive and provide substantial amounts

of summer forage (McInnis and Vavra 1986). Plants

are highly resistant to grazing because their growing

points remain below ground throughout the growing

season, and they produce a low ratio of reproductive to

vegetative stems (Ehrenreich and Aikman 1963). Few

grasses withstand heavy grazing as well as Kentucky

bluegrass. It increases rapidly on overgrazed pastures

and ranges, and its presence is usually an indication of

poor grazing management. For livestock use, these

sites are best managed under a grazing system other

than season long.

Kentucky bluegrass is highly palatable to most large

ungulates in spring when it is green and succulent. In

moist mountain meadows, palatability remains moderately high during summer (Bellrose 1980). In drier

areas it becomes semidormant in summer and its

palatability is much reduced. Regionally, Kentucky

bluegrass is important in the diets of elk, mule deer,

and bighorn sheep (Dittberner and Olson 1983). Grasslands dominated by this grass provide habitat for

numerous small mammals (George and others 1978;

Medin and Clary 1989). Kentucky bluegrass meadows

found along mountain streams are often preferred

foraging areas for wild ungulates and grouse. Mueggler

and Campbell (1986), however, suggested that the

aspen/Kentucky bluegrass community type in Utah is

one of the poorest aspen types for wildlife habitat due

to its low diversity of plant species.

Kentucky bluegrass leaves and seeds are eaten by

small mammals and songbirds (Martin and others

1951). Prairie chickens eat small amounts of its seeds

(Crawford and others 1969). It is also an important

food for cottontail rabbits and wild turkeys (Crawford

and others 1969; George and others 1978). Sagegrouse use Kentucky bluegrass-dominated areas for

strutting and chick rearing.

The use of Kentucky bluegrass for revegetation on

rangelands is limited because of its slow establishment, lack of drought tolerance, and its high soil

fertility requirements (Vogel 1981; Wasser 1982). When

planted in seed mixtures, it often takes 2 to 3 years to

become established (fig. 45).

The value of Kentucky bluegrass on rangeland is

also limited because of its low productivity, summer

dormancy, and propensity to invade native grasslands. In the Intermountain West, Kentucky bluegrass is well adapted to meadows with seasonally high

water tables and midsummer drought (Volland 1985a).

It has become naturalized and dominates many
408 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

meadows once occupied by tufted hairgrass and sedges.

Reestablishment of the original natives is impractical

because of the competive ability of Kentucky bluegrass.

Because of its shallow root system, Kentucky bluegrass is not recommended as a soil stabilizer. It is less

effective than the native grasses and forbs it has

replaced, and it is not capable of stabilizing

streambanks. Erosion and channel downcutting may

occur when it dominates, especially in heavily grazed

areas (Hansen and others 1988a,b; Kovalchik 1987).

Kentucky bluegrass is intolerant of prolonged flooding, high water tables, and poor drainage (Wasser

1982).

Kentucky bluegrass is one of America’s most popular lawn grasses. It withstands considerable abuse

and it is often used as a sod grass on golf courses,

campgrounds, and ski slopes (Shaw and Cooper 1973).

It fails to control weed invasion and spread, particularly in riparian sites and meadows. Serious weed

problems occur where this species has displaced native grasslike species.

Varieties and Ecotypes

Many varieties of Kentucky bluegrass have been

selected and released for turf purposes. Most are not

adapted for rangeland use. The most popular variety

for wildland seeding is ‘Newport’, obtained from a

coastal site in Lincoln County, OR (Alderson and

Sharp 1994). Newport is seeded in the Intermountain

West to provide soil stabilization. It is adapted to

well-drained alluvial soils derived from limestone.

Newport is rhizomatous, moderately drought tolerant, and exhibits good growth compared to other

Kentucky bluegrass accessions. Newport is less competitive than some Kentucky bluegrass varieties, and

it can be seeded in mixtures with some species.

Poa secunda

Sandberg Bluegrass _____________

Synonyms

Paneion sandbergii

Poa canbyi

Poa confusa

Poa nevadensis

Poa sandbergii

Poa scabrella

Description

The Poa secunda complex occupies a wide range of

habitats, and its members exhibit many forms, several

having been given species status over the years.

Sandberg bluegrass is one of the more common earlyseason desert bunchgrasses in the Intermountain area.

It is a small, densely tufted perennial that often

produces extensive tufts of basal leaves that wither

and disappear early (fig. 46). Plants are often more or

less purple tinged throughout. Culms are 0.7 to 1.5 ft

(2 to 4.5 dm) tall and wiry. Sheaths are smooth to

roughened and open for three-quarters or more of

Figure 45—Two-year-old Kentucky bluegrass plants

spread rhizomatously and develop a dense ground

cover (RMRS photo).

Figure 46—A small, tufted cool-season perennial,

Sandberg bluegrass is one of the most common

bunchgrasses of the Intermountain area (photo courtesy of Loren St. John, USDA NRCS, Aberdeen Plant

Materials Center, Aberdeen, ID).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 409

Chapter 18 Grasses

their length. Leaf blades are soft, folded or involute,

rarely flat, 0.02 to 0.06 inch (0.5 to 1.5 mm) wide, and

1.2 to 1.95 inches (3 to 5 cm) long. The basal cluster of

leaves is 1.2 to 5.5 inches (3 to 14 cm) tall. The leaves

are roughened and their tips are prow shaped. Ligules

are tapered to a point and rather prominent. Those of

the basal leaves are mostly less than 0.06 inch (1.5

mm) long, and those of the culm leaves are usually

more than 0.10 inch (2.5 mm) long. Panicles are 0.8 to

7.9 inches (2 to 20 cm) long, narrow to rather open, but

usually not dense. Panicle branches are short and

usually ascending. Spikelets are 0.16 to 0.37 inch (4 to

9.5 mm) long, two to six flowered, slightly compressed

and rather cylindrical, narrow, and pointed. They are

usually strongly purple tinged. Glumes are roughened, translucent along the margins, and tapered to a

point. The first glume is 0.09 to 0.20 inch (2.2 to 5 mm)

long and one nerved; the second is 0.12 to 0.20 inch (3

to 5 mm) long and three nerved. Lemmas are 0.12 to

0.22 inch (3 to 5.5 mm) long, rounded below and only

slightly keeled above. They become increasingly curly

haired below, and are not at all webbed at the base

(Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969).

Ecological Relationships and Distribution

Sandberg bluegrass ranges from Alaska through

much of Canada and south into the Western United

States, Mexico, Nebraska, and Michigan. The species

is also found in South America. It grows at elevations

from 980 to 12,000 ft (300 to 3,660 m) on sites receiving

10 to 22 inches (254 to 559 mm) of annual precipitation

(Hitchcock 1950; Stubbendieck and others 1992; USDA

Forest Service 1937; USDA Natural Resources Conservation Service 2003). Sandberg bluegrass occurs on

dry to mesic plains, foothills, and ridgetops of salt

desert shrub, sagebrush, pinyon-juniper, oakbrush,

mountain brush, grassland, aspen, conifer, and meadow

communities (Arnow 1987).

Sandberg bluegrass occurs on soils of a wide range of

textures, but it is most productive on deep, welldrained sandy clay, silt, and sandy loam soils that

range from slightly acidic to basic. It is often a dominant species on dry, rocky sites and on shallow soils of

scablands. The species is tolerant of salinity and of

sites that are flooded in spring, but it does not tolerate

soils that are saturated for prolonged periods (Arnow

1987; Hafenrichter and others 1968; Plummer and

others 1968).

Sandberg bluegrass begins growth in early spring

prior to most other native bunchgrasses and sets seed

in late spring or early summer, depending upon elevation. Plants evade summer drought by entering dormancy. Growth may resume in fall if adequate precipitation occurs. The dense tufts of leaves and the fibrous

rooting system provide stabilization of surface soils

(Jensen and others 2001).

Sandberg bluegrass is generally not damaged by

wildfires. The small clumps, low amounts of litter, and

prolonged summer dormancy preclude extensive damage to the buds. Early fires during dry years may be

more damaging (Kearney and others 1960). Cover

provided by the species may increase for a time following fire if other species are lost or reduced (Blackburn

and others 1971). Reestablishment of sagebrush or

other taller species may eventually reduce the cover

provided by Sandberg bluegrass, as it is not shade

tolerant (Acker 1992; Tueller 1962).

Plant Culture

Regeneration of Sandberg bluegrass is by seed and

tillering with occasional production of a small number

of rhizomes (Arnow 1987). The species is self-fertile or

wind pollinated and facultatively apomictic

(Daubenmire 1970; Hitchcock and others 1969; Kellogg

1985a,b). Plants produce considerable seed in early

summer. Native stands are harvested by hand or with

mechanical strippers. Little seed is normally harvested from wildlands, and such seed is often of low

viability. Seed fields of the released varieties are

harvested by swathing (Smith and Smith 1997) or by

direct combining (Rose and others 1998). Some seed

may be lost through shattering. Seed is cleaned using

a hammermill and air-screen machine.

There are about 925,000 seeds per pound (2,039,255

per kg) (Hafenrichter and others 1968). Yields from

seed fields of the Canby bluegrass type average 150

pounds per acre (168 kg per ha) (Smith and Smith

1997). Wasser (1982) recommended 90 percent purity

and 65 percent germination as purchasing standards.

AOSA (1999) recommendations for germination are

pretreatment with 0.2 percent potassium nitrate followed by incubation at alternating temperatures of 68

and 86 ∞F (20 and 30 ∞C) (16 hrs/8 hrs) with exposure

to light during the high-temperature periods.

Seed fields and wildland seedings of Sandberg bluegrass should be planted on firm seedbeds in clay loam

to sandy loam soils at a 0.25 to 0.5 inch (0.6 to 1.3 cm)

depth in fall (Fulbright and others 1982). Germination

is slow and seedling vigor is low. Stands establish

slowly and there is little production during the first

season (Smith and Smith 1997; Wasser 1982). Stand

life for seed fields is 7 or 8 years. Use of local seed

sources is recommended for wildland plantings because of the wide distribution of the species.

Uses and Management

Sandberg bluegrass has been successfully seeded in

areas receiving as little as 12 inches (300 mm) of
410 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

annual precipitation (Arnow 1987). Seedlings are not

highly competitive. New seedings must be protected

from grazing and weed invasion. Established plants

are more competitive, but also compatible with other

native species. Plants are short lived; management

should permit periodic seed production and natural

seeding to perpetuate stands and reduce the risk of

cheatgrass invasion. Seedling establishment and stand

productivity fluctuate widely with weather conditions

(Daubenmire 1975b).

Sandberg bluegrass is one of the earliest grasses to

be grazed in spring. It provides high-quality and

palatable, but coarse, forage for livestock and wildlife

from spring to early summer. Both forage quality and

palatability decline rapidly as the plant matures, but

the species receives some use in summer and fall.

Summer regrowth is limited; fall regrowth may be

moderate, depending upon precipitation (Thornburg

1982). Plants are highly resistant to grazing and

trampling.

Sandberg bluegrass is an important component of

the Townsend ground squirrel’s diet (Johnson 1977).

It provides some cover for small mammals, songbirds

and waterfowl, and other small animals. Elk, deer,

and other wildlife species browse it (Vallentine 1967).

Sandberg bluegrass provides some surface soil stabilization, particularly on shallow soils where it may be

the dominant species.

Varieties and Ecotypes

‘Canbar’ originated in the Blue Mountains of

Columbia County, WA (Alderson and Sharp 1994). It

is adapted to basin and Wyoming big sagebrush and

mountain brush communities. Although it establishes slowly when seeded, it eventually fills the

interspaces between larger bunchgrasses, thus inhibiting the invasion of cheatgrass. Once established,

Canbar is a vigorous understory species with an abundance of basal leaves. It makes excellent early spring

growth, and it spreads well from seed. When planted

with other native species it is compatible and not

overly competitive.

‘High Plains’ Selected Germplasm is a composite of

materials from Montana and Wyoming (Englert and

others 2002). The variety grows to 2 ft (0.6 m) in

height. It was selected for use in native seedings to

provide wildlife habitat and restore native communities. It is recommended for use on adapted sites in

Montana and Wyoming.

‘Service’ originated near Whitehorse, Yukon Territory, Canada (Alderson and Sharp 1994). It has not

been seeded in the Intermountain West. It has, however, performed well in Alaska when seeded for erosion control and revegetation of disturbances on dry,

gravelly, and rocky soils.

Sitanion hystrix

Squirreltail, bottlebrush __________

Synonyms

Elymus elymoides

Aegilops hystrix

Sitanion elymoides

Sitanion californicum

Sitanion rigidum

Sitanion glabrum

Sitanion cinereum

Sitanion montanum

Sitanion brevifolium

Sitanion longifolium

Sitanion hordeoides

Description

Squirreltail is a short-lived, loosely to densely caespitose perennial that is nearly hairless to white hairy

throughout. Culms are erect to spreading and 0.3 to

2.1 ft (1 to 6.5 dm) tall, with hairless to silky haired leaf

sheaths. Leaf blades are flat to folded or involute, 0.4

to 0.24 inch (1 to 6 mm) wide, stiff haired to soft hairy

above, and sometimes hairless below. Auricles are

inconspicuous, up to 0.04 inch (1 mm) long, and often

purplish. Ligules are short, less than 0.2 inch (5 mm)

long, and squared at the margins. Spikes are compact

to loose and open, and 1.2 to 5.9 inches (3 to 15 cm)

long, excluding the awns. They are often included in

the upper leaf sheaths, or sometimes entirely exerted

in robust specimens. The rachis disarticulates as the

spikelets mature. Spikelets are mostly in pairs, sometimes in threes, or solitary when in the upper portion

of the spike. They are mostly one to six flowered,

sometimes all fertile, but more commonly with the

lowermost flower sterile and glumelike. Awns of the

glumes and lemmas are 0.8 to 4.7 inches (20 to 120

mm) long, varying from green to red or often purplish.

Glumes are 1.4 to 3.4 inches (35 to 85 mm) long,

subulate, and extend into slender, spreading, roughened awns. Lemmas are 0.3 to 0.4 inch (8 to 10 mm)

long, hairless to strongly short haired, and faintly

three to five nerved. The main nerve extends into a

slender to stout, flexuous awn that may be straight or

divergent, while two of the lateral nerves extend into

bristles that are about 0.4 inch (10 mm) long (Arnow

1987, 1993; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969). Plants are selfpollinating and allotetraploid (2n = 28).

Squirreltail is a member of the Tribe Triticeae, and

hybridizes with members of the Elymus, Hordeum,

and Pseudoroegneria complexes. It also produces sterile hybrids with Great Basin wildrye (Jones 1998).

Barkworth and others (1983) included the species in
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 411

Chapter 18 Grasses

Elymus elymoides. Wilson (1963) and Jones and Larson

(in press) describe squirreltail as a complex of five

taxa. Most widespread is E. e. var. elymoides. E. e. var.

californicus is native to the eastern Sierra Nevadas. E.

e. var. brevifolius are larger plants, while E. e. var.

hordeoidesare small and less common. Big squirreltail

occurs in the Northwest and southward into California and parts of Nevada. The taxa exhibit considerable

differences in morphological and phenological characteristics (Hironaka and Tisdale 1972; Jones and Larson,

in press) and occupy differing habitats.

Ecological Relationships and Distribution

Squirreltail ranges from British Columbia to

Saskatchewan and south throughout the Western and

Central United States and into Mexico. It grows in

plant communities ranging from salt desert shrub to

alpine meadows on sites receiving 8 inches (20 mm)or

more of annual precipitation (Horton 1989) at elevations ranging from 2,000 to 11,500 ft (610 to 3,500 m)

(Kearney and others 1960). It is a common understory

species in sagebrush steppe communities (Jones 1998).

Squirreltail commonly grows on dry, gravelly soils

that may be saline or alkaline, but some populations

also grow on fine-textured soils (Jensen and others

1990; Morris and others 1950). Squirreltail may be an

early successional species in overgrazed or otherwise

disturbed sites. It is also a mid to late successional

species (Young and others 1972). Jones (1998) suggests its seral status may be related to both genotype

and site conditions.

Bottlebrush squirreltail is competitive with the annual grasses cheatgrass and medusahead and has

been observed to persist on ranges invaded by these

weeds (Arredondo and others 1998; Beckstead 1994;

Hironaka 1994) (fig. 47). The ability to germinate and

produce roots at low temperatures permits it to establish in annual grass-infested areas (Hironaka 1994;

Hironaka and Sindelar 1975; Young and Evans 1977).

Facultative fall or spring germination, early spring

growth, extensive root development, and early seed

development contribute to the competitive ability of

the species (Reynolds and Fraley 1989). Plants are

drought tolerant and winter hardy.

Squirreltail is fairly tolerant of burning (Britton and

others 1990) due to its small size, limited leafy material, and coarse, solid stems. The low herbage volume

generally limits fire damage to the root crown (Wright

1971; Wright and Klemmedson 1965). Plants are more

susceptible to damage when surrounded by accumulations of litter and dead material or when fire frequencies increase (Wright and Klemmedson 1965). Damaged plants regenerate from surviving portions of the

root crown and from on- and offsite seed sources

Figure 47—Squirreltail is competitive with invasive

annual grasses, including cheatgrass and

medusahead (RMRS photo).

(Bradley and others 1992a,b). Squirreltail’s ability to

compete with cheatgrass may be favored by fire.

Plant Culture

Squirreltail seeds mature from June to September

depending on site conditions (Link 1993). Mature

seeds are dispersed following disarticulation of the

rachis, thus timing of harvest is critical. Doescher

(2001) found seed could be harvested when awns are

divergent and straw colored or about 1 week prior to

disarticulation of the rachis. Most viable seed falls

near the plant, but fruit morphology, particularly the

long awns, permits wind dispersal, and some seeds can

be carried as much as 130 ft (40 m) (Beckstead 1994;

Hironaka and Tisdale 1963; Marlett and Anderson

1986). The long awns, short seedstalks, and large

amounts of chaffy material make seed collection difficult. Seed fields and extensive wildland stands in

accessible areas are harvested by combining or by first

windrowing. Wildland stands are also hand harvested.

Awns may be removed with a brush machine, and

trash can be removed with an air-screen machine.

There are about 191,555 seeds per pound (422,302 per

kg) (Plummer and others 1968). Seed can be stored for

3 or 4 years (Link 1993).

Seed can be germinated at constant or alternating

temperatures of 59, 50/59, or 50/68 ∞F (15, 10/15, or 10/

20 ∞C) without prechilling (Young and Evans 1977). A

requirement for afterripening may vary among seed

lots and range from 0 to 4 months (Allen and others

1994; Beckstead and others 1993; Young and Evans

1977).
412 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Squirreltail seed may be direct seeded alone or in

mixtures at depths of 0.25 to 0.5 inch (0.6 to 1.2 cm).

Seed fields are short lived. Weed control is difficult

and plants are subject to rust. However, seed can be

harvested the first year (Archibald and Feigner 1995),

and production is generally good (Hironaka and Tisdale

1963; Young and Evans 1977).

Because of the great variability within this species

complex, seed sources for revegetation efforts must be

selected carefully. Seed should be planted in fall in the

Intermountain and Rocky Mountain areas and in late

winter to early spring in the Southern Great Plains.

Plummer and others (1968) ranked germination of

squirreltail seedings high, but they considered initial

establishment, growth rate, final establishment, and

persistence of squirreltail seedings only moderate.

Squirreltail has long been considered one of the

more competitive native grasses on cheatgrass rangelands. Its ability to germinate rapidly across a wide

temperature range is considered a factor in its ability

to establish on disturbed sites (Young and Evans

1977). Allen and others (1994) found that squirreltail

from a low-elevation site (4,100 ft [1,250 m]) was more

likely to germinate in fall than cheatgrass. In Nevada,

Tipton (1994) found that mature squirreltail plants

usually initiated growth before cheatgrass. Jones (1998)

described squirreltail as a potential species for “assisted succession,” a species capable of replacing invasive species following wildfire. He listed squirreltail

attributes suited to this role, including facultative fall

germination, rapid growth and maturation, root systems capable of mining nitrogen, early spring maturation in areas that dry rapidly, and efficient nitrate

assimilation.

Uses and Management

Squirreltail greens up early in spring and provides

fair amounts of forage where it grows in dense stands.

The bristlelike awns can injure grazing animals. Consequently, animals avoid these plants from seed ripening until seed dispersal. Late summer herbage and

especially fall regrowth following rainfall are palatable and used by livestock and wildlife. Squirreltail is

a valuable plant for providing energy in winter, as

some leaves remain green (Jensen and others 2001;

Stubbendieck and others 1992; Vallentine 1971). It is

particularly valuable to big game and small mammals

in winter (Anderson and Shumar 1986; Beale and

Smith 1970; Kufeld and others 1973). Palatability and

use by livestock and big game vary geographically

(USDA Forest Service 1937).

Squirreltail has considerable potential for revegetating degraded rangelands, particularly those invaded by annual weeds. The USDI Bureau of Land

Management considers development of squirreltail

seed sources a priority for revegetating Intermountain

rangelands (Hardegree 1994). Squirreltail provides a

persistent ground cover and can be used to control

erosion. Direct seeded squirreltail survived for 30

years on a big sagebrush/bluebunch wheatgrass site in

south-central Idaho (Monsen and Anderson 1993).

Stand management should emphasize protection from

excessive grazing and protection during heading so

adequate seed supplies will be produced and the stand

maintained (USDA Forest Service 1937; Sanders 1994).

Varieties and Ecotypes

Accessions 9040187 and 9040189 originated in Colorado and are being advanced toward release by the

Upper Colorado Environmental Plant Center in

Meeker, CO (Alderson and Sharp 1994; UCEPC 2002).

They were selected for their potential for revegetation

of mined lands and other disturbed rangeland sites.

Their forage quality is fair to moderate for livestock

and wildlife, especially in winter.

Fish Creek, a Sitanion hystrixvar. hystrix collection,

was recommended for release as a Selected Germplasm

by the Utah Agricultural Experiment Station in 2002.

It was collected from a site near Carey, Blaine County,

ID, at an elevation of 4,756 ft (1,400 m) that receives

about 15 inches (380 mm) of annual precipitation

(Jones and others 2002c). Fish Creek was selected for

its low proximal awn mass, a trait that may reduce

damage during awn removal. In addition, its spikes

disarticulate at the base, facilitating seed harvesting.

The intended area of use for Fish Creek is the upper

Snake River Plain.

Toe Jam Creek was collected from a site at 6,000 ft

(1,829 m) west of Tuscarora, Elko County, NV (Jones

and others 2002b). Tuscaraora receives 12.3 inches

(312 mm) of annual precipitation. Toe Jam Creek was

developed for use in the northern Great Basin and

lower Snake River Plain. It was selected for its large

seed size and low proximal awn mass. The latter trait

may reduce damage during awn removal, a problem

encountered in processing Sand Hollow. Toe Jam

Creek was identified as Sitanion hystrix var. californicus. The Utah Agricultural Experiment Station

recommended it for release as a Selected Germplasm

in 2002.

Sand Hollow (Sitanion jubatum or Elymus multisetus) big squirreltail Selected Germplasm originated

in Gem County, ID, on a loamy, coarse sand site at an

elevation of 2,690 ft (820 m) that receives 11 inches

(300 mm) of annual precipitation (Jones and others

1998b). It is adapted to sandy soils in southern Idaho

and adjacent regions in Oregon, Nevada, and Utah.

The variety was developed for revegetation of disturbed sites and rangelands dominated by exotic

annuals.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 413

Chapter 18 Grasses

Sporobolus airoides

Alkali Sacaton __________________

Synonyms

Agrostis airoides

Sporoblus wrightii

Description

Alkali sacaton is a long-lived, sodforming, native

perennial bunchgrass that produces dense clumps

that die out in the center over time. In the Southwest

this species acts as a warm-season grass, while in the

Intermountain area it responds more like a coolseason grass. Plants are stout and tufted forming

large clumps clothed with slick, shiny, cream-colored

sheaths at the base. Culms are 16 to 47 inches (4 to 12

dm) tall, hairless, round, and usually hollow, but

sometimes loosely pith filled. Leaf sheaths are smooth

and hairless to slightly silky haired at the upper edges

of the margins. Leaf blades are flat to strongly involute, 0.08 to 0.16 inch (2 to 4 mm) wide, smooth below,

finely roughened above, and stiff haired near the

throat. Ligules are very short with a dense band of

hairs. Panicles are 4.7 to 15.7 inches (12 to 40 cm) long,

0.16 to 0.80 inch (4 to 20 mm) wide, open and pyramidal, and often fully exerted or sometimes remaining

partially enclosed in the sheath (fig. 48). Spikelets are

borne mostly toward the tips of the branches. Glumes

are often deciduous. They are tapered to a point, one

nerved, roughened or often translucent throughout,

and glabrous. The first glume is 0.03 to 0.08 inch (0.7

to 2 mm) long, and the second is about 0.06 to 0.12 inch

(1.5 to 3 mm) long. Lemmas are 0.7 to 0.12 inch (1.8 to

3 mm) long and taper to a point; they are usually

longer than the second glume (Arnow 1987; Hitchcock

1950; Hitchcock and others 1969; Stubbendieck and

others 1986; USDA Forest Service 1937).

Ecological Relationships and Distribution

Alkali sacaton occurs throughout the Western United

States, the Great Plains, and Northern Mexico. It is

most abundant in saltgrass, salt desert shrub, desert

shrub, blackbrush, sagebrush-grass, and pinyon-juniper communities. It can also be found in ponderosa

pine, mountain brush, semiarid grasslands, and on

prairies. It inhabits lower, slightly moist, alkaline

flats, where it frequently develops almost pure stands.

Stands are also found scattered along drainages in

desert and semidesert areas. Throughout the northern portion of its range, it is scattered and less important, but in the Southwest, it occurs in sufficient

abundance to be a major forage species.

Alkali sacaton generally grows in areas receiving 12

to 14 inches (300 to 350 mm) of annual precipitation.

It grows on dry to moist sites in saline bottoms and in

areas with shallow water tables. It tolerates soils with

up to 3 percent total salts, as well as soils high in

carbonate and bicarbonate (Blaisdell and Holmgren

1984; Cook and others 1954; Roundy and others 1983;

Vallentine 1961). Although it can endure alkali, it is

not restricted to alkaline soils. It does poorly on sandy

soils, fair on silty soils, and good on clayey soils

(Vallentine 1971). Plants exhibit moderate fire tolerance, but little shade tolerance.

Plant Culture

Alkali sacaton reproduces from seeds and tillers.

Plants produce numerous seeds from late summer to

October. These may remain viable for many years

(Blaisdell and Holmgren 1984; Hitchcock 1950). Seeds

must undergo an afterripening period of up to 9 months

Figure 48—Alkali sacaton is a long-lived bunchgrass

that forms large, dense clumps and produces wide,

pyramidal panicles (Hitchcock 1950).
414 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

to relieve dormancy. Local flooding will move seeds

across flood plains to saturated sediments where they

germinate when favorable conditions are encountered

(Aldon 1981).

Seeds are small with an average of 1.75 million per

pound (3.85 million per kg). Purity can be high (98+

percent). Germination of 80 percent is acceptable for

seed purchases. Seed viability remains high for

more than 10 years when seeds are stored dry in a

warehouse.

Seed can be broadcast or drill seeded. Extensive

seedbed preparation is not required. Seeds should be

planted about 0.25 inch (6 mm) deep. Fall seeding is

recommended to permit afterripening and release of

dormancy. Within the same seed lot, some seeds germinate within a few days under favorable condtions,

but others are dormant and may not germinate for

weeks. Seedling establishment is, therefore, erratic

without some type of irrigation, flooding, or runoff.

Soils that remain moist for 5 to 10 days to support

germination can produce fair stands (Aldon 1975).

Difficulty in establishment is probably the factor that

most limits the wider use of alkali sacaton. Although

some plantings produce erratic stands, this species

does establish with a good degree of success in quite

arid communities. It can be seeded in situations where

few other species exist. Planting in mixtures with

other species, including some shrubs, does not prevent

its establishment.

Uses and Management

Alkali sacaton provides valuable forage for livestock

in arid and semiarid situations where it can produce a

large quantity of forage. As it matures and dries it can

become coarse and somewhat unpalatable. Cattle and

horses do, however, use it in winter months. Sheep will

use it when forced during drought and on overgrazed

ranges (Blaisdell and Holmgren 1984; Cook and others 1954; Vallentine 1961). Palatability has been rated

as good for cattle, good to fair for sheep and horses, and

fair to poor for antelope, elk, and deer (Dittberner and

Olson 1983).

Alkali sacaton is commonly seeded to stabilize disturbed lands, such as raw mine spoils in the semiarid

Southwest (Hassell 1982; Oaks 1982; Thornburg 1982).

It is also planted in riparian zones in a number of

major plant communities in the Intermountain area

(Monsen 1983). This species can be superior to western wheatgrass for seeding in drier portions of the

southern and northern desert shrub types (Alden

1981; Plummer 1977).

Alkali sacaton has been used in range and reclamation seedings in blackbrush, salt desert shrub, sagebrush, pinyon-juniper, and mountain brush types. It is

seeded along roadways and riparian corridors. It can

establish well and persist in lower elevation shrublands

that support few species. In these situations it is

important for controlling weed invasions and protecting unstable, erosive soils. This species could become

much more important for the restoration of arid sites.

Varieties and Ecotypes

Two varieties of alkali sacaton have been selected

and released (Alderson and Sharp 1994). Salado was

selected for use in conservation and range seedings.

Collected near Clauch, NM, at 5,800 ft elevation

(1,770 m) it is native to a site that receives 11 to 18

inches (280 to 460 mm) of annual precipitation. Salado

is seeded on valley bottoms, alkaline flats, rocky soil,

and open plains in the Intermountain West to provide

forage and soil stabilization. It is used on disturbed

sites where moisture accumulates, such as road rightof-ways, depressions, and gravel pits. Fall seeding is

preferred because germination may be erratic. Seedling vigor is generally good.

Saltalk, which originated near Erik, OK, has not

been seeded to any great extent in the Intermountain

West. It is seeded in the Southwest on disturbances in

salt-bearing soils around oil wells.

Sporobolus cryptandrus

Sand Dropseed _________________

Synonyms

Vilfa cryptandra

Agrostis cryptanda

Description

Sand dropseed is a tufted drought-tolerant,

nonrhizomatous native bunchgrass that sometimes

functions as an annual. It performs as a warm-season

grass in the Southwest and as a cool-season grass in

the Intermountain West and other northern areas. It

can be recognized by the large, distinctive, inflated

sheathes that subtend and partially enclose the inflorescence. When the sheaths wither, the panicle expands and the seeds are dispersed (fig. 49). Culms are

1.3 to 3.0 ft (4 to 9 dm) tall, erect or prostrate at the

base, hairless, and solid. Leaf sheaths are strongly

overlapping, ciliate at the summit, and sometimes

hairy along the upper margins. The hairs are 0.04 to

0.16 inch (1 to 4 mm) long and sometimes occur across

the collar. Blades are usually flat with roughened

margins and become involute toward the tip. They are

0.06 to 0.20 inch (1.5 to 5 mm) wide and 2 to 6 inches

(5 to 15 cm) long. Ligules consist of a ring of dense hairs

about 0.02 inch (0.5 mm) long. Panicles are open, 4 to

12 inches (1 to 3 dm) long, and 0.6 to 2.4 inches (1.5 to
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 415

Chapter 18 Grasses

6 cm) wide, with the lower part at least partially

enclosed in a sheath. Panicle branches are hairless,

spikelet bearing almost to the base, and usually rather

stiffly ascending to reflexed. Spikelets are cleistogamous, or self-fertilized, pale olive green, and hairless.

Glumes are tapered to a tip, one nerved, and deciduous

at maturity. The first is 0.03 to 0.04 inch (0.7 to 1 mm)

long and awl shaped. The second is 0.06 to 0.09 inch

(1.4 to 2.2 mm) long and broader than the first. Lemmas are about as long as the second glume, 0.06 to 0.10

inch (1.5 to 2.5 mm) long, and also tapered to a tip

(Arnow 1987; Cronquist and others 1977; Hitchcock

1950; Hitchcock and others 1969; USDA Forest Service

1937). Sand dropseed roots spread up to 2 ft (0.6 m)

horizontally and 3 ft (0.9 m) vertically (Weaver and

Albertson 1956).

Ecological Relationships and Distribution

Sand dropseed is an important grass in the Southwest, the Intermountain West, and parts of the Snake,

Salmon, and Clearwater drainages in Idaho and Oregon

(USDA Forest Service 1937). It is distributed from

Ontario, Canada, to Alberta and Washington, and

south to North Carolina, Indiana, southern California, and northern Mexico (Hitchcock 1950). In the

Intermountain West it is most abundant in deserts or

lowlands, and it is especially common on sandy soils.

This species can be a major component of sagebrushbunchgrass types, particularly in Idaho and Utah. It is

also important in juniper-pinyon, mountain brush,

salt desert shrub, blackbrush, chaparral, and ponderosa pine communities and short-grass prairies. Sand

dropseed will invade degenerated wheatgrass, fescue,

and bluegrass communities. It is able to persist with

heavy ungulate use because it has a protected root

crown, matures late, and is not a highly preferred species

(Johnson and Simon 1987). Sand dropseed responds

rapidly to grazing removal, becoming a major component of overgrazed communities (Blaisdell and

Holmgren 1984; Cook and others 1954). It can also be one

of the first grasses to regenerate on abandoned fields,

disturbed sites, waste places, and along roadsides.

Plant Culture

Sand dropseed is a prolific seed producer. It spreads

well from natural and artificial seeding, and it is often

one of the first perennial grasses to reestablish following drought. Seeds will lie dormant in the soil for many

years until favorable conditions for germination and

establishment occur (USDA Forest Service 1937;

Weaver and Albertson 1956). Seedling development is

rapid, with tillers appearing a few weeks after germination (Weaver and Albertson 1956).

Seeds are small with 5.6 million per pound (12.3

million per kg). Seed is often marketed with 98+

percent purity and total germination of 90 to 95 percent. However, some seed lots may contain as much as

50 percent hard seed. Seed can be stored for 20 years

and still be 75 percent germinable (USDA Forest

Service 1937). Normally, the older the seed, the better

the germination and establishment. The species is

widely distributed, and ecotypes differ in germination

requirements.

Sand dropseed requires minimal seedbed preparation. It can be successfully seeded by drilling or broadcasting and light harrowing on a variety of sites,

including open exposed soils and unstable disturbances. Seed should not be planted deeper than about

0.24 inch (6 mm), especially in sandy soils. The seed

coat of sand dropseed is very hard and requires overwinter prechilling for germination. Fall plantings can

provide natural prechilling and increase the probability of spring germination. Seed dormancy can be a

deterrent to the establishment of uniform seeded

stands. The growth rate of sand dropseed is considered

good on sandy and salty soils and fair on clayey soils

(Stubbendieck and others 1986). It is one of the few

Figure 49—Inflated sheaths that subtend and partially enclose the inflorescence of sand dropseed

wither when mature, permitting inflorescence expansion and seed dispersal (photo courtesy of

Kevin Jensen, USDA ARS, Logan, UT.
416 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

grasses that are well adapted to stabilizing sandy

sites. Once plants establish, they are long lived and

hardy.

Sand dropseed is compatible with many other species and can be seeded in mixtures in disturbed salt

desert shrub, blackbrush, big sagebrush, pinyon-juniper, mountain brush, chaparral, and short-grass prairie communities. In the Southwest, it does well in

areas that are too hot and dry for wheatgrasses. Sand

dropseed is also frequently planted alone.

Sand dropseed will establish on sites that receive

less than 9 inches (230 mm) of annual precipitation.

The survival of new plantings of sand dropseed is

dependent upon the availability of spring and early

summer soil moisture. Plantings that receive good

spring precipitation emerge, grow quickly, and become vigorous and capable of persisting during subsequent periods of summer drought.

Uses and Management

Sand dropseed is fairly productive and palatable to

all classes of livestock and many wildlife species,

especially from spring to midsummer (Daubenmire

1970; Hitchcock 1950; Stubbendieck and others 1986;

Vallentine 1961). Its palatability is rated from fair to

good depending upon associated plants. In many places,

this species has been removed due to continued excessive grazing. On ranges in poor to fair condition, sand

dropseed may still provide a major proportion of available forage (Alzerraca-Angelo and others 1998;

Stubbendieck and others 1986). It is also used in early

summer when plants are succulent. Cattle use it in fall

following rainfall. This pattern of use favors sand

dropseed growth (Johnson and Simon 1987). Plants

will respond to fall rains and provide succulent forage

into the winter months (Arnow 1987; Stubbendieck

and others 1986). In addition, the herbage cures well

and can provide fair to good winter forage (USDA

Forest Service 1937).

Wild turkeys feed on sand dropseed in south-central

South Dakota in the Missouri River Breaks region.

This grass is an important component in the summer

diet of Rocky Mountain bighorn sheep, and it is important to deer and elk during most seasons (Rominger

and others 1988).

Sand dropseed is an important species that can be

used to slow the spread of cheatgrass. Remnant stands

and established plants are competitive and can reduce

weed invasion. The species can often be seeded successfully on weedy sites if some initial control measures are used. Once established, it becomes highly

competitive. It has been useful when planted on big

sagebrush and mixed pinyon-juniper sites in central

and southern Utah where some cool-season bunchgrasses are less well adapted. This species should be

included in revegetation programs wherever it occurs

naturally. It will tend to dominate in some areas such

as sandy outcrops.

Varieties and Ecotypes

There are no released varieties. There is, however,

considerable variation within the species. Care should be

taken to insure that seed of adapted sources is seeded.

Stipa columbiana

Subalpine Needlegrass, Columbia

Needlegrass ____________________

Synonyms

Achnatherum nelsonii

Stipa viridula var. minor

Stipa nelsonii

Stipa occidentalis

Description

Subalpine needlegrass is an erect, long-lived, coolseason, native perennial bunchgrass. It produces fine,

densely bunched culms that are usually 12 to 24 inches

(30 to 60 cm) tall, but can be as much as 39 inches (1

m) tall. Leaf sheaths are strongly ribbed. Leaf blades

can be 4 to 10 inches (10 to 25 cm) long and 0.04 to 0.12

inch (1 to 3 mm) wide. Leaf edges are flat against the

stem and then rolled inward toward the midrib, especially on younger leaves; they are hairless or slightly

roughened to densely hairy and often strongly striated. Ligules are short, 0.01 to 0.07 inch long (0.2 to 2

mm), square, rather firm, and usually longest on the

side. Panicles are narrow, dense, and 2.8 to 8 inches (7

to 20 cm) long (fig. 50). Panicle branches are short and

remain close to each other. Spikelets produce only one

floret about 0.8 to 1 inch (2 to 2.5 cm) long. The awn is

short with the lower portion twisted. The basal end of

the awn is sharp pointed (Arnow 1987; Hitchcock

1950; Hitchcock and others 1969; USDA Forest Service 1937).

Ecological Relationships and Distribution

Subalpine needlegrass grows in all of the Western

States, particularly the Central Rocky Mountains. It

occurs mostly above 8,000 ft (2,440 m), often in dense

stands, on dry soils of open side hills. It also grows in

mountain parks, subalpine herblands, and openings

in aspen and conifer forests. It may occur as scattered

individuals in mountain brush and upper elevation

sagebrush areas. There are two recognized varieties:

S. c. var. columbiana occurs from the Yukon to South

Dakota and south to Texas and California; S. c. var.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 417

Chapter 18 Grasses

nelsonii occurs from Alberta to Washington and south

to Arizona and Colorado (Hitchcock 1950).

Subalpine needlegrass often increases on ranges

where wheatgrasses and bluegrasses have been removed by excessive use. It resprouts quickly after fires

and is able to persist, while other species are often

slower to respond. It recovers readily with rest from

grazing, often more rapidly than most associated

grasses. It will, however, suffer from use during periods of drought.

Subalpine needlegrass typically occurs as scattered

individuals in sagebrush and mountain-brush communities, but denser stands may be found in subalpine

grasslands (Vallentine 1961). It often exists on open

south and west slopes in association with spreading

rabbitbrush, western yarrow, and bluegrasses. Under

these conditions it furnishes important ground cover

and soil protection. This needlegrass is slightly to

moderately damaged by fire (Wright and others 1979).

It has moderate tolerance to salinity and will grow on

soils ranging from sandy loam to clay loam to dry,

rocky, infertile soils.

Plant Culture

Awns can be removed without damage to the seed,

and seeds can be cleaned to purities exceeding 90

percent. Seed lots contain between 150,000 to 200,000

seeds per pound (330,000 to 440,000 seeds per kg).

Considerable variation in seed weights, size, and

dormancy exist among and within separate wildland

collections. Afterripening of dry seeds does occur, and

most seeds maintained in storage for 6 months will

germinate. Seeds stored for 1 to 7 years usually germinate better than fresh seeds.

Cleaned seed lots can usually be planted with most

conventional seeders without damaging the seed. Seeds

are long and narrow and must be able to pass through

openings and regulators that can accommodate long

seeds. Seeds must be placed in the soil by drilling or

other methods that bury the seed at least 0.25 to 0.50

inch (6 to 12 mm) deep. Broadcasting seeds on the soil

surface without some means to cover them is not

recommended. Seeds may be dormant, but fall seeding

allows seeds to overwinter in the soil, which breaks

dormancy and facilitates spring germination. Most

species within this genus establish slowly the first

season. Although first-year seedlings are normally

small, survival may be quite high. Young plants can be

slow to reach maturity, especially if seeded in mixtures with other, more rapidly developing species. The

slow-developing plants furnish needed ground cover,

and are less competitive with establishing shrubs and

slower developing herbs. Once established, this perennial can persist with a number of other species in

rather harsh environments. It is able to exist in the

seedling stage with some degree of shade from mixed

overstory shrubs.

Uses and Management

Subalpine needlegrass provides valuable forage for

many species of wildlife and livestock. It begins growth

in early spring and remains green throughout a long

growing season, sometimes until early winter. Most

needlegrasses cure well and are grazed during fall and

winter (USDA Forest Service 1937). The pointed callus

of subalpine needlegrass sometimes works into the

mouth and ears of livestock (Stubbendieck and others

1992); this causes animals to often avoid the mature

seed.

Palatability of subalpine needlegrass is rated as

good for cattle and horses, fair to good for sheep and

deer, and fair for antelope. It is especially valuable for

sheep in the spring and for livestock in the summer.

Deer use it in summer, and many other wildlife species

consume it throughout the growing season (Vallentine

1961). It is particularly valuable for deer and elk in

late fall and winter, as some green remains on the

plant. Like most needlegrasses, subalpine needlegrass

is palatable early in the season, becoming less so as the

foliage becomes coarse and wiry (USDA Forest Service 1937). This species also provides cover for small

mammals and birds (Dittberner and Olson 1983).

Subalpine needlegrass is characterized by a slow to

moderate seedling growth rate, moderate salinity tolerance, and the ability to grow on a number of soil

Figure 50—Panicles of subalpine needlegrass are

narrow, compact, and up to 8 inches (20 cm) long;

spikelets produce a single floret (Hitchcock 1950).
418 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

types. It is an important species for revegetation of a

variety of plant communities that are often seeded to

intermediate wheatgrass and smooth brome. It establishes well and can be used to reestablish native

communities, especially where a mixture of species

and plant types are desired.

Varieties and Ecotypes

A number of accessions are undergoing development. Accession 9040137 from San Lewis Valley and

Accession 9024804 from western Colorado, are being

advanced toward release by the Upper Colorado Environmental Plant Center, in Meeker, CO. These are

adapted to areas at elevations of 5,000 to 12,000 ft

(1,525 to 3,660 m) receiving 15 inches (380 mm) or

more of annual precipitation. They have potential for

soil stabilization, rangeland seedings, and revegetation of disturbed sites in the West (Alderson and Sharp

1994).

Stipa comata

Needle-and-Thread ______________

Synonyms

Hesperostipa comata

Stipa juncea

Stipa tweedyi

Description

Needle-and-thread is an erect, long-lived native

perennial cool-season bunchgrass. It is named for the

exceptionally long, twisted, and tapering awns that

resemble threaded sewing needles. Plants are tufted

with culms that are 1.0 to 3.6 ft (3 to 11 dm) tall and

often slightly hairy at the nodes. Sheaths are smooth

to roughened, strongly ribbed, and usually longer than

the internodes. Leaf blades are involute, or if flat, up

to 0.12 inch (3 mm) wide and 4 to 12 inches (10 to 30 cm)

long, roughened on the upper surface, and smooth to

slightly roughened beneath. Ligules are 0.04 to 0.24

inch (1.1 to 6 mm) long, more or less minutely hairy,

and usually tapering to a tip. They may become lacerate with age. Panicles are 5 to 15.5 inches (13 to 40 cm)

long, narrow, and usually partly enclosed in an inflated sheath. Panicle branches are usually slender

and ascending, sometimes with drooping spikelets.

Glumes are long and narrow, tapering to a fine point.

They are five nerved, slightly convoluted, hairless,

and papery, with translucent margins and tips. The

first glume is 0.7 to 1.4 inches (1.8 to 3.5 cm) long, and

the second is 0.6 to 1.3 inches (1.5 to 3.4 cm) long.

Lemmas are 0.32 to 0.55 inch (8 to 14 mm) long, pale

green to yellowish or brownish, sparsely covered with

appressed silky hairs, and often hairless toward the

tip. The swollen node (callus) at the base of the floret

is very sharp, about 0.12 inch (3 mm) long. The first

joint in the awn is distinct; the second is rather

indistinct and more flexuous. The lower segment is

tightly twisted and roughened; the terminal segment

is not (Arnow 1987; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969; USDA

Forest Service 1937) (fig. 51).

Ecological Relationships and Distribution

Needle-and-thread is widely distributed throughout

the Western States and Great Plains. It commonly

occurs in sagebrush-grass, pinyon-juniper, and ponderosa pine communities of the Rocky Mountains, Intermountain West, and semidesert plains and foothills of

the Southwest. Needle-and-thread is most often found

on well-drained soils.

With overgrazing, needle-and-thread is one of the

first species to disappear from native pastures and

Figure 51—Needle-and-thread is named for the

florets with their extremely long awns that resemble

threaded sewing needles (photo courtesy of Loren

St. John, USDA NRCS, Aberdeen Plant Materials

Center, Aberdeen, ID).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 419

Chapter 18 Grasses

range sites. Likewise, it is one of the first species to

respond and reoccupy native communities following

rest from grazing.

Plant Culture

Needle-and-thread depends on seed for reproduction. In favorable areas, it reproduces well and it is

able to withstand grazing in spring and fall if plants

are allowed to mature seed in summer. On drier

ranges, however, it does not reproduce readily and it is

easily killed by overuse.

Needle-and-thread can be established in sagebrush

and some salt desert shrublands. It is drought tolerant

and requires well-drained soil. Seeds can be drilled or

broadcast, but they must be covered with at least 0.25

inch (6 mm) of soil. Needle-and-thread can be seeded

successfully with most native perennial grasses and

broadleaf herbs. It should not be seeded with species

that are vigorous competitors. Seedling development

of needle-and-thread can be slow, but even during lowprecipitation years some seedlings will generally establish. It may fail to develop dense or dominant

stands for the first 5 to 10 years after planting, even

when seeded alone. After this time, the stands develop

slowly, even during years of low annual rainfall. As the

stand becomes denser, persistent perennials may be

suppressed. Seeded stands of needle-and thread have

displaced intermediate wheatgrass on some pinyonjuniper planting sites in central Utah approximately

25 years after the initial seeding. This species has also

recovered naturally on sites where pinyon and juniper

trees have been removed and the native understory

has been allowed to recover. Heavy grazing by livestock or big game can delay or suppress natural recovery.

Needle-and-thread has not prevented shrub seedling establishment when seeded with big sagebrush,

antelope bitterbrush, Stansbury cliffrose, or curlleaf

mountain mahogany. It can be utilized to establish a

mixed array of native species without necessitating

separate planting operations. Stands of this species

provide sufficient competition to restrict the spread of

annual weeds. It recovers quickly after burning and

does not allow weeds to enter after fires.

Needle-and-thread seed is marketed with the awn

removed. There is an average of 115,000 seeds per

pound (253,000 seeds per kg). Some seed dormancy

exists, but seed with 75 to 85 percent germination and

80 to 85 percent purity is generally marketed. Seed

can be stored for a number of years with little loss of

viability. The long, tough awns limit commercial production and use of this species. Some growth forms

produce seeds with flexible awns that do not break

from the seeds. Growth forms with brittle awns can be

cleaned and seeded with much less difficulty. Techniques and equipment have been developed to de-awn

some seeds. Even when cleaned, the sharp tip on the

seed and awn fragments can cause handling and seeding problems. Mixtures of needle-and-thread and nonawned seed of other species can be planted with most

seeders.

Uses and Management

Needle-and-thread is an important range grass for

livestock and wildlife species because of its early

spring and late fall greenup, palatability, volume of

forage produced, and widespread distribution. The

forage value of needle-and-thread varies depending on

the region, season, and associated plants. Its palatability is rated very high in early spring before the

awns develop and again after the seeds are dispersed.

It is especially valuable because it begins development

in early spring when other grasses have not yet initiated growth. An abundance of herbage is produced and

will remain green through summer and fall if adequate moisture is available. Awns can become a

problem when the fruit matures because the sharp,

barbed lemmas can injure grazing animals. New growth

is produced in summer and fall if rainstorms occur.

Needle-and-thread cures well and is often utilized as

winter forage. This is one of the most important

grasses of semiarid and arid shrublands of the Intermountain region. Its loss due to grazing has resulted

in an increase of annual weeds, particularly cheatgrass.

Recovery of this perennial is critical to control weeds.

This species is an important element of the shrubbunch grass communities and pinyon-juniper woodlands, and reestablishment of functional communities

requires inclusion of this species.

Varieties and Ecotypes

There are no released varieties. Care should be

taken to ensure that seed of adapted populations is

planted. Great Plains collections do not do especially

well in the Intermountain West.

Stipa lettermanii

Letterman Needlegrass __________

Synonyms

Stipa viridula var. lettermanii

Description

Letterman needlegrass is an erect, fine-stemmed,

densely-tufted, native perennial bunchgrass that often forms large clumps. Stems are slender and 8 to 20

inches (20 to 50 cm) tall. Leaves are threadlike and

somewhat twisted. Culms are 10 to 33 inches (2.5 to

8.5 dm) tall. Culms and leaf sheaths are hairless or
420 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

minutely roughened. Leaf blades are involute, rounded,

rarely flat, and up to 0.08 inch (2 mm) wide and 4.0 to

8.0 inches (10 to 20 cm) long, stiff haired above and

hairless to minutely roughened below. Ligules are

0.01 to 0.08 inch (0.2 to 2 mm) long and rounded to

square. Panicles are narrow and about 2.8 to 9.5

inches (7 to 24 cm) long with erect branches and

spikelets. Glumes are subequal, 0.20 to 0.41 inch (5 to

10.5 mm) long, and taper gradually to a tip. They are

often purple with translucent margins, hairless or

sometimes minutely roughened, and three nerved.

Lemmas are only slightly hardened, 0.20 to 0.26 inch

(4.5 to 6.7 mm) long, and pale with soft, slender,

somewhat appressed hairs that become longer toward

the tip. The callus often has dense, white hairs less

than 0.04 inch (1 mm) in length. Awns are 0.40 to 0.87

inch (1.0 to 2.2 cm) long, slender, and twice bent with

the lower segment loosely twisted and minutely roughened; the terminal segment is 0.3 to 0.5 inch (7 to 12

mm) long and hairless. The basal end of the seed

tapers to a pointed tip (Arnow 1987; Cronquist and

others 1977; Hitchcock 1950; Hitchcock and others

1969; USDA Forest Service 1937).

Letterman needlegrass is often misidentified as

subalpine needlegrass because the two plants are

similar in size. They can, however, be differentiated by

the width and length of the palea in relation to the

length of the lemma. The palea of Letterman

needlegrass is about two-thirds the length of the

lemma, and it is often exposed. The palea of subalpine

needlegrass is less than half as long as the lemma. The

distribution of the two species is about the same, both

grow in all Western States, with the Central Rocky

Mountains as their center of distribution. Letterman

needlegrass is, however, less abundant at the northern and southern ends of its range, and it does not

extend to altitudes as high as does subalpine

needlegrass.

Ecological Relationships and Distribution

Letterman needlegrass is found on dry soils in open

mountain parks, mountain meadows, subalpine grasslands, and in openings in aspen and conifer forest from

Oregon and Montana, south to Arizona and New

Mexico (fig. 52). It is also found in midelevation pinyon-juniper, sagebrush, and mountain brush communities. It grows best on sandy loam, loam, or clay loam

soils 20 inches (51 cm) or more in depth (Dittberner

and Olson 1983). It tends to increase under heavy

grazing by sheep and decrease under heavy cattle use.

Letterman needlegrass is a climax or late seral

species in a number of grassland, sagebrush, mountain-shrub, and pinyon-juniper communities. It also

grows well in early successional assemblages, and is

well represented in many early to late seral plant

associations (Schott 1981).

Plant Culture

Seeds may be drilled or broadcast seeded, but they

must be covered with about 0.25 inch (0.60 cm) of soil.

Fall seeding is recommended because seed dormancy

can be overcome by overwinter stratification.

Letterman needlegrass has been seeded successfully on disturbed sites to control erosion, revegetate

mines, and restore lost stands. As is the case for other

needlegrass species, seedlings develop slowly.

Letterman needlegrass can be planted with most native species that are not highly competitive. Letterman

needlegrass usually produces seeds each year, and it

will spread into unoccupied sites.

Uses and Management

Letterman needlegrass is an important component

of many sagebrush, mountain brush, and aspen communities where it may vary in density and abundance.

It recovers well after fires and can persist with some

shade or competition. Although it is rated as only fair

for cattle and poor for sheep, it is an important forage

species on many sites (USDA Forest Service 1937).

Heavy grazing by cattle or sheep can cause a decrease,

or sometimes a short-term increase, in density.

Letterman needlegrass provides valuable forage for

wildlife and domestic livestock. Plants begin growth

early in the year, and they remain green throughout

the summer and late fall. This enables Letterman

needlegrass to be utilized when many other grasses

are less available or palatable. Most needlegrasses,

including Letterman needlegrass, cure well and can be

utilized in fall and winter. This species is particularly

Figure 52—Letterman needlegrass grows in mountain

meadows, forest openings, and subalpine grasslands

(RMRS photo).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 421

Chapter 18 Grasses

important as late fall and early winter forage for

big game; the presence of some green material within

the plant attracts these animals and influences their

distribution.

Varieties and Ecotypes

There are no releases.

Stipa thurberiana

Thurber Needlegrass ____________

Synonyms

Achnatherum thurberianum

Stipa occidentalis

Description

Thurber needlegrass is a densely tufted perennial

with culms 1.2 to 2.6 ft (3.5 to 8 dm) tall that are

minutely hairy, at least at the nodes. Leaf sheaths are

striated and usually hairless, but sometimes minutely

haired below. Blades are needlelike, involute, minutely roughened, and 3.9 to 9.8 inches (10 to 25 cm)

long. Ligules are 0.08 to 0.28 inch (2 to 7 mm) long, thin

and translucent, hairless to sparsely haired, and tapering at the tip. Panicles are narrow and about 2.8 to

9.5 inches (7 to 24 cm) long with erect branches that

are relatively few flowered. Glumes are subequal and

0.31 to 0.63 inch (8 to 16 mm) long with the second

slightly shorter than the first. They are three- to fivenerved, often purplish and translucent above, and

taper to a soft tip. Lemmas are 0.24 to 0.34 inch (6.2 to

8.7 mm) long and sparsely covered with silky, appressed hairs. The callus is about 0.04 inch (1 mm)

long, with dense, appressed hairs. Awns are 1.2 to 1.8

inches (30 to 45 mm) long and bent twice. The lower

segments are twisted with stiff, feather-like hairs up

to 0.08 inch (2 mm) long. The terminal segment is 0.6

to 1.2 inches (16 to 30 mm) long and roughened to

hairless. Thurber needlegrass flowers from late May

through July. It is known to hybridize with Oryzopsis

hymenoides, producing the hybrid Stipa bloomeri

(Arnow 1987, 1993; Cronquist and others 1977;

Hitchcock 1950; Hitchcock and others 1969; Johnson

1945).

Ecological Relationships and Distribution

Thurber needlegrass ranges from eastern Washington, Oregon, and California, to northern Nevada, southern Idaho, southwestern Montana, and northeastern

Wyoming (Arnow 1987, 1993; Cronquist and others

1977; Hitchcock 1950; Hitchcock and others 1969). It

may occur as a dominant or associated mid to late seral

species in grasslands, sagebrush deserts, and pinyonjuniper, oak, and ponderosa pine woodlands (fig. 53).

In sagebrush deserts it is most common in communities receiving 6 to 16 inches (152 to 406 mm) of annual

precipitation (USDA Natural Resources Conservation

Service 2003). It is fairly widespread in the northwestern portion of the Great Basin.

Thurber needlegrass is adapted to soils ranging

from well-drained clay loams to coarse-textured soils

that are fairly neutral (USDA Forest Service 1937;

USDA Natural Resources Conservation Service 2003).

Plants are tolerant of drought and cold, but they are

not salt or shade tolerant (USDA Natural Resources

Conservation Service 2003).

Response of Thurber needlegrass to fire is variable,

but it is generally less resistant to burning than other

bunchgrasses (Wright and others 1979). Burning is

more damaging early in the season than later (Britton

and others 1990; Ganskopp 1988; Uresk and others

Figure 53—Thurber needlegrass occurs as a

dominant or common associated species on dry

big sagebrush sites (RMRS photo).
422 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

1980). The presence of dry and sometimes abundant

leafy culms contributes to surface heating and burning of the root crown during wildfires (Wright and

Klemmedson 1965). Accumulations of dead vegetative

material contribute to fire damage at all seasons.

Plants may be killed or living tissue may survive near

the edges of the root crown. Plants then regenerate

from tillers if postfire weather conditions permit

(Britton and others 1990; Koniak 1985; Uresk and

others 1976, 1980). Surviving plants recover slowly

and exhibit reduced vigor and productivity for up to 3

years. The number of inflorescences and seed production, however, may be enhanced by burning (Uresk

and others 1980), contributing to postburn regeneration from seed (Hironaka and others 1983).

Thurber needlegrass seedlings compete poorly with

those of more competitive annual and perennial grasses

(Evans and Young 1978; Uresk and others 1980; Young

and Evans 1978). Dependence on residual or offsite

seed sources and low germination and seedling vigor

tend to slow recovery following burns. Consequently,

Thurber needlegrass often does not become abundant

until communities are mid to late successional (Koniak

1985).

Plant Culture

Thurber needlegrass ripens in mid to late summer

(Sampson 1924). Seed production is generally low. There

are about 225,000 seeds per pound (496,035 per kg)

(USDA Natural Resources Conservation Service 2003).

Haferkamp and McSwain (1951) found that germination was enhanced by incubation at a fluctuating (68/

86 ∞F) (20/30 ∞C) temperature (16 hrs/8 hrs) with

exposure to light during the high-temperature periods.

Thurber needlegrass can be seeded on adapted sites

receiving as little as 10 inches (254 mm) of annual

precipitation (Thornburg 1982). Litter has been found

to reduce germination and early growth of Thurber

needlegrass seedlings, but it was also associated with

improved growth of establishing seedlings 4 weeks

after germination (Schlatterer and Tisdale 1969). Seedling vigor is low to moderate (Hironaka and others

1983; USDA Natural Resources Conservation Service

2003), and spread of stands from seed is slow.

Uses and Management

Thurber needlegrass provides valuable forage for livestock and wildlife. Deer, wild horses, pronghorn, blacktailed jackrabbits, and other species graze the herbage

(Fagerstone and others 1980; McInnis and Vavra 1987;

Trainer and others 1983; Uresk and others 1976). Birds

and other small animals use its seeds. Thurber

needlegrass and associated species provide cover for

many small vertebrates and invertebrates (Stanton 1974).

Sage-grouse hens were found to use a Wyoming big

sagebrush/Thurber needlegrass cover type in eastern

Oregon prior to nesting (Barnett and Crawford 1994).

Thurber needlegrass is valuable for cattle and sheep

in early spring when other forage is not available.

Palatability and nutritive value of the species is greatest early in the season, declining as plants and seeds

mature and the forage becomes coarse (USDA Forest

Service 1937). Leaves are used to a greater degree

than stems. The species tends to be avoided as the

seeds mature because the awns can irritate or injure

the ears, eyes, and noses of grazing animals. Because

of this, considerable seed generally matures, even in

areas grazed by livestock (Jensen and others 2001;

USDA Forest Service 1937). Thurber needlegrass remains green after other grasses dry and provides

useful forage in fall.

Thurber needlegrass stands are damaged by excessive grazing and fire. Overgrazing in spring can be

particularly detrimental because defoliation, particularly during the boot stage, can weaken plants

(Ganskopp 1988). The species tends to increase with

protection from grazing (Robertson 1971). Thurber

needlegrass can be used for reclaiming mine sites and

other disturbances on dry sites where it is adapted.

Such uses, however, are limited by a lack of seed.

Varieties and Ecotypes

There are no releases.

Stipa viridula Trin.

Green Needlegrass ______________

Synonyms

Nassella viridula

Stipa parviflora

Stipa nuttalliana

Stipa sparta

Description

Green needlegrass is a long-lived, native, cool-season, perennial bunchgrass that is tolerant of drought,

cold winters, and a wide range of soil types. It is a

tufted perennial with culms 1.6 to 3.6 ft (5 to 11 dm)

tall. Leaf sheaths are persistent and hairless or hairy

along the margins. Blades are flat to involute, smooth

or roughened, and 0.80 to 0.24 inch (2 to 6 mm) wide.

Ligules are 0.02 to 0.12 inch (0.5 to 3 mm) long. Many

plants are functionally pistillate and bear only small,

sterile anthers; thus, the species may be semi-dioecious. Panicles are narrowly oblong, 4 to 10 inches (1 to

2.5 dm) long, and approximately 0.8 inch (2 cm) wide.

Panicle branches are erect or only slightly divergent.

Glumes are translucent throughout, or nearly so, equal

or subequal, and 0.32 to 0.51 inch (8 to 13 mm) long,
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 423

Chapter 18 Grasses

tapering to a long, hairlike tip. They are prominently

three to five nerved and hairless. The lemma is brown

and leathery at maturity, 0.20 to 0.26 inch (5 to 6.5

mm) long, and narrowed toward the tip into a hairless,

whitish “neck” approximately 0.04 inch (1 mm) long.

The awn is 0.8 to 1.4 inches (2 to 3.5 cm) long and twice

bent, with the two lower segments twisted and minutely roughened. The callus is up to 0.04 inch (1 mm)

long, with a very short, blunt, hairless tip (Arnow

1987; Cronquist and others 1977; Hitchcock 1950;

Hitchcock and others 1969; USDA Forest Service

1937). Green needlegrass occasionally hybridizes with

Indian ricegrass, but the offspring are sterile (Hitchcock

and others 1969).

Ecological Relationships and Distribution

Green needlegrass is a perennial bunchgrass distributed from British Columbia to Minnesota, Kansas,

New Mexico, Arizona, Nevada, and western Washington. It occurs more commonly in Montana, Wyoming,

and Colorado than in the other Western States. Green

needlegrass grows on dry slopes, plains, prairies, and

foothills at lower elevations and often may be found

growing along roadways. It is also common in mountain meadows, open woodlands, and on hillsides at

higher elevations. At the southern edge of its distribution, green needlegrass grows in dry parks and canyons and with ponderosa pine (USDA Forest Service

1937). It is a prominent species in many grassland and

sagebrush communities, and it grows well on abandoned agricultural lands (Wasser 1982).

Green needlegrass exists as an early or late seral

species. It is adapted to a wide range of soils. It grows

especially well on clay and moderately alkaline soils

derived from calcareous shales. It is somewhat less

adapted to loam and sand, and it is not very tolerant

of saline conditions (Wesser 1982). This species exhibits considerable drought tolerance, which is nearly

equal to that of western wheatgrass. It can be found

growing in areas with as little as 10 inches (250 mm)

of annual precipitation. Optimum growth occurs with

12 to 20 inches (310 to 510 mm) of precipitation. It

grows, however, in areas with as much as 30 inches

(760 mm) precipitation and in areas with water tables

only 1 to 2 ft (30 to 60 cm) deep (Wasser 1982). Green

needlegrass is moderately tolerant of flooding, only

weakly tolerant of shade, and somewhat tolerant of fire,

depending on the season of burn and site conditions.

Plant Culture

Green needlegrass flowers and disperses seeds from

July to September. Cleaning seeds to remove the awns

and other debris can be completed without damage to

the seed. There are an average of 181,000 seeds per

pound (400,000 per kg). Seed of most green needlegrass

collections exhibit relatively high levels of seed dormancy. It is not uncommon to purchase fresh seed

with less than 10 percent germination and 90 percent

hard seed. Considerable afterripening occurs during

dry storage with germination percentage increasing

up to 7 years after harvest (Rogler 1960). Consequently, better stands are normally obtained with

older seed. Fall seeding allows seeds to overwinter in

the soil. This can relieve dormancy and allow seeds to

germinate and seedlings to establish in spring. For

this reason spring seeding is not recommended. Seed

can be drilled or broadcast but must be covered because maximum germination occurs in darkness

(Fulbright and others 1982). Seed need not be seeded

deeper than about 0.25 inch (6 mm).

Germination potential and seed maturity are correlated. Immature seed, which is light gray to yellowish

green, has a lower average weight than does mature

seed, which is a uniform gray (Kinch and Wiesner

1963). Average viability of immature seed is approximately 22 percent compared with 71 percent for mature seed (Kinch and Wiesner 1963).

Within the Intermountain region, green needlegrass has performed well when planted in aspen,

mountain brush, and mountain big sagebrush communities. Sites that receive at least 14 inches (350 mm)

of annual precipitation will generally support this

grass. Vigor of green needlegrass seedlings is good

(Kinch and Wiesner 1963). However, when planted

with pubescent or intermediate wheatgrass, its initial performance has been suppressed by these

sodformers.

Mature green needlegrass plants are very competitive with annual weeds, but new seedlings are not.

When planted in areas with cheatgrass and other

annuals, the weedy species must be eliminated or

controlled to facilitate the establishment of this perennial. New plants will establish in areas where annual

kochia and mustards dominate. New seedlings are

also able to establish and survive in mixed grass-forb

seedings. Green needlegrass typically establishes well

but can decrease over time; it is often planted in

mixtures of cool-season, native or introduced grasses

or forbs that tend to persist.

Uses and Management

Many Stipa species produce awns or calluses that

cause injury to grazers. This is not the case with green

needlegrass. Green needlegrass provides valuable forage for many species of wildlife and all classes of

livestock (fig. 54). It is palatable to cattle and horses

season long, but is used by sheep primarily in spring

and early summer (USDA Forest Service 1937).

Green needlegrass begins growth in early spring

and remains green until late summer when most other

grasses have dried. In many areas, annual growth
424 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 18 Grasses

Figure 54—Most common in the northern Great Plains,

green needlegrass provides forage for many wildlife

species (photo courtesy of Tom Jones, USDA ARS,

Logan, UT).

begins as early as March (Wasser 1982), with most

growth occurring May and June. Late-season regrowth

can be good if fall precipitation is sufficient. This

species is generally found in mixed stands with other

grasses, although dense stands are common. As a

result, succulent forage is available over a longer

period of time where green needlegrass is part of the

community. Green needlegrass cures well and can be

used for fall and winter grazing.

Green needlegrass is moderately resistant to heavy

grazing (Wasser 1982), and generally recovers quickly

(Kinch and Wiesner 1963). It can be harmed by intense

early-season grazing (Wasser 1982). Wasser (1982)

recommends moderate grazing of stands and resting a

different unit for 2 months during the growing season

each year. When green needlegrass occurs with warmseason grasses, seeded stands should be grazed during

the spring and fall or in winter (Wasser 1982).

Green needlegrass has been successfully seeded

throughout the Intermountain area. It has done especially well on disturbed sites in the mountain brush

zone and lower elevation aspen zones. Green

needlegrass has been widely used for rehabilitating

disturbed sites. This species has a wide range of

adaptation and performs well in rangeland plantings

in the aspen and mountain brush communities throughout the Intermountain West. It has also performed

well on mine and rangeland seedings in eastern Wyoming, Montana, and western Colorado. Green

needlegrass is particularly useful for erosion control.

It has been successfully seeded on abandoned farmlands and pastures. Seedlings develop slowly, and

should not be grazed for up to 4 years following

seeding. This species can be planted in mixtures to

restore native communities. It does not compete to

exclude native shrubs and herbs.

Varieties and Ecotypes

‘Cucharas’ was collected near Walsenburg, CO, and

is recommended for use in the Central Great Plains

(Jones and Larson, in press; Jones and others 2002d,

2004b). It was selected for its high germination without a wet prechill treatment. Cucharas was recommended for release as a Selected Germplasm by the

Utah Agricultural Experiment Station in 2002.

‘Lodorm’ originated near Bismarck, ND (Alderson

and Sharp 1994), and is seeded for forage, soil stabilization, and revegetation of mined sites and other

disturbances. Lodorm was developed for the Northern

Great Plains, but it is also being seeded in the Intermountain West. It is adapted to areas receiving 14 or

more inches (355 mm) of annual precipitation and

elevations above 9,000 ft (2,745 m) in mountain brush,

ponderosa pine, and aspen openings. It is seeded on

bottomlands, flats, benches, and along streams. Lodorm

is adapted to silt and clay soils, and it is tolerant of

dense clays. It will also grow on loamy and sandy soils.

This variety exhibits rapid vegetative recovery, high

palatability, low seed dormancy, and rapid germination. It is compatible with other natives and can be

sown in mixtures. Fall seeding is recommended to

relieve seed dormancy. This grass is very winter hardy,

tolerant of fire and short-duration flooding, and weakly

shade tolerant.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 425

Richard Stevens

Stephen B. Monsen

19

Chapter

Forbs are abundant in all vegetative types throughout the

Intermountain West. Most are found intermixed in grasslands

and as understory plants in shrub and forest types. Forbs

provide ground cover, soil stability, community (flora and fauna)

diversity, nutritious forage, and are of aesthetic value.

Forbs should be seeded in most range, watershed, mine reclamation, highway, recreational site, restoration and revegetation

projects. Many commonly used forbs have been introduced to

the Intermountain West and are now under cultivation for seed

production. The principal introduced forbs planted on range and

wildlands are alfalfa, small burnet, and cicer milkvetch. Lewis

flax, Rocky Mountain penstemon, and Palmer penstemon are

the major native forbs being commercially grown and planted.

Additional species are being commercially grown in seed fields as

demand increases. Seeds of many native species are collected

from wildland stands but generally are in short supply.

Forbs for Seeding Range

and Wildlife Habitats
426 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Chapter Contents by Species

Achillea millefolium ssp. lanulosa (western yarrow) .......................................................................................... 430

Artemisia ludoviciana (Louisiana sage) ...............................................................................................................430

Aster chilensis (Pacific aster) ................................................................................................................................ 431

Aster glaucodes (blueleaf aster) ............................................................................................................................433

Astragalus cicer (cicer milkvetch) .........................................................................................................................433

Balsamorhiza sagittata (arrowleaf balsamroot) .................................................................................................. 435

Balsamorhiza macrophylla (cutleaf balsamroot) ................................................................................................. 436

Balsamorhiza hookeri var. neglecta (hairy balsamroot) ...................................................................................... 436

Coronilla varia (crownvetch) ................................................................................................................................ 437

Geranium richardsonii (Richardson geranium) .................................................................................................. 438

Geranium viscosissimum (sticky geranium) ........................................................................................................ 439

Hedysarum boreale (Utah sweetvetch) ................................................................................................................. 439

Helianthella uniflora (oneflower helianthella) .................................................................................................... 440

Heracleum lanatum (cow parsnip)........................................................................................................................441

Ligusticum porteri (Porter ligusticum)................................................................................................................. 442

Linum perenne spp. lewisii (Lewis flax) ...............................................................................................................443

Lomatium triternatum (nineleaf lomatium) ........................................................................................................ 444

Lomatium kingii (nuttall lomatium) .................................................................................................................... 445

Lupinus sericeus (silky lupine) .............................................................................................................................446

Lupinus argenteus (silvery lupine) ....................................................................................................................... 447

Medicago sativa (alfalfa) .......................................................................................................................................447

Medicago falcata (sicklepod alfalfa) .....................................................................................................................447

Melilotus officinalis (yellow sweetclover) ............................................................................................................. 451

Mertensia arizonica (tall bluebell) ........................................................................................................................452

Onobrychis viciaefolia (sainfoin) ...........................................................................................................................453

Osmorhiza occidentalis (sweetanise) .................................................................................................................... 454

Penstemon palmeri (Palmer penstemon) .............................................................................................................. 454

Penstemon cyananthus (Wasatch penstemon) ..................................................................................................... 456

Penstemon eatonii (Eaton penstemon) ................................................................................................................. 456

Penstemon humilis (low penstemon) .................................................................................................................... 457

Penstemon pachyphyllus (thickleaf penstemon) .................................................................................................. 457

Penstemon rydbergii (Rydberg penstemon) .......................................................................................................... 457

Penstemon strictus (Rocky Mountain penstemon) ............................................................................................... 457

Sanguisorba minor (small burnet) ....................................................................................................................... 457

Senecio serra (butterweed groundsel)...................................................................................................................459

Solidago canadensis (Canada goldenrod)............................................................................................................. 460

Sphaeralcea grossulariifolia (gooseberryleaf globemallow) ................................................................................ 461

Sphaeralcea coccinea (scarlet globemallow) .........................................................................................................462

Trifolium (clovers) .................................................................................................................................................463

Trifolium fragiferum (strawberry clover)............................................................................................................. 463

Trifolium hybridum (alsike clover)....................................................................................................................... 464

Trifolium pratense (red clover) .............................................................................................................................464

Trifolium repens (white clover) .............................................................................................................................464

Viguiera multiflora var. multiflora (Showy goldeneye)....................................................................................... 465

Viguiera miltiflora var. nevadensis (Nevada goldeneye) ..................................................................................... 466

Important characteristics of a number of forbs are

listed in table 1. Seeding recommendations for principal vegetative types and conditions are discussed in

chapter 17. Seed characteristics and information

about collection, cleaning, and storage are provided

in chapter 24. Forbs adapted to different vegetative

types and conditions are included in the seeding

recommendations.

The following forb species have the potential for

improving range and wildlife habitats in the Intermountain West. A brief description along with information regarding ecological relationships and distribution, cultural requirements, use, and management

of each species are included.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 427

Chapter 19 Forbs for Seeding Range and Wildlife Habitats Table 1—Characteristics of Selected Forbs. Ratingsb - Vegetative Early Compati- typea to Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of adapt- species is Species seed seeding planting nation lishment rate lishment tence spread yield bility bility grazing ness species tion stability ance ance ability adapted Alfalfa, range type 5 5 4 5 5 5 4 5 4 5 5 4 4 3 5 5 4 2 4 5 SA,A,PP,MB,JP, MS,BS,WS Alfileria 2 4 2 4 4 5 3 2 5 3 5 3 4 1 3 4 3 1 3 3 JP,MS,BS, WS,BB Angelica, small leaf 2 514 553 334 543 1 3 5 3 3 3 2 SA,A,MB Aster, blueleaf 2 2 5 4 4 4 5 5 5 4 4 5 5 2 2 3 5 4 3 4 A,WM,PP,MB, JP,BS Aster, Englemann 2 2 4 3 3 3 4 4 5 4 4 5 4 2 4 2 4 4 4 2 A,WM,PP,MB Aster, Pacific 2 2 5 4 3 3 4 5 5 3 5 5 5 2 3 3 5 4 3 4 A,WM,PP, MB,JP,MS,BS Balsamroot, arrowleaf 4 413 114 525 514 1 4 4 3 1 2 3 PP,MB,JP, MS,BS Balsamroot, cutleaf 4 413 114 423 4 1 4 1 4 3 3 1 2 2 PP,MB,MS Balsamroot, hairy 4 4 1 3 1 1 4 5 2 4 5 1 4 1 4 4 3 1 3 2 PP,MB,MS Bluebell, tall 3 3 4 3 2 4 3 4 3 5 5 5 4 1 2 3 4 4 3 2 SA,A,WM Bouncing-bet 5 5 5 4 4 4 4 5 5 4 3 3 4 2 3 4 5 4 3 3 PP,MB,MS Burnet, small 5 5 4 5 5 5 3 3 4 3 5 5 3 5 4 5 3 2 2 3 PP,MB,JP,MS, BS,WS Cinquefoil, gland 4 4 5 4 5 4 5 5 5 5 3 3 5 1 5 4 5 4 1 3 A,WM,PP,MB Clover, alsike 5 5 5 4 4 3 4 4 4 4 4 4 4 2 3 3 4 5 3 1 WM Clover, strawberry 5 5 5 5 4 3 4 5 5 2 4 4 4 2 4 3 4 5 3 2 WM,IS Columbine, Colorado 2 4 4 3 3 3 4 3 2 2 4 4 2 1 4 2 3 5 5 2 SA,A,WM Cow parsnip, common 3 3 2 3 2 3 4 3 2 5 4 4 2 1 4 4 3 4 4 2 SA,A,WM Crownvetch 5 5 5 4 4 4 4 5 5 4 4 4 4 2 2 4 5 3 4 3 A,PP,MB,MS Daisy, common oxeye 3 3 4 2 2 3 5 5 5 2 4 5 4 2 3 3 4 2 3 3 A,PP,MB,JP Deervetch, birdfoot 3 4 5 4 4 4 4 3 4 5 5 5 5 2 3 3 5 3 3 3 A,PP Eriogonum, cushion 4 424 434 432 354 4 3 4 323 4 PP,MB,JP, MS,BS Fireweed 1 2 4 5 5 5 3 2 5 4 4 3 5 1 4 4 4 3 4 3 SA,A,WM,PP, MB,JP,MS,BS Flax, Lewis 5 5 4 4 5 4 4 3 5 3 5 3 4 4 4 4 3 2 3 4 PP,MB,JP,MS, BS,WS Geranium, Richardson 2 3 4 3 2 4 4 4 3 3 4 4 4 2 4 2 3 4 4 2 SA,A,WM Geranium, sticky 2 3 4 3 2 4 4 4 3 3 4 4 4 2 4 2 3 4 4 2 SA,A,WM Giant hyssop, nettleleaf 2 433 233 334 444 2 4 4 333 2 SA,A,MB Globemallow, 3 5 3 3 3 3 4 4 4 3 3 2 4 2 4 3 4 2 2 3 JP,MS,BS,WS, gooseberryleaf SS,BB Globemallow, scarlet 4 4 4 2 3 4 4 5 3 3 3 3 4 2 4 3 4 3 3 3 MB,JP,MS,BS, WS,SS,BG,BB Globemallow, stream 4 5 4 3 2 3 4 4 4 4 3 3 3 3 4 3 4 3 2 3 JP,MS,BS,WS, SS,BB Goldeneye, Nevada 3 3 4 4 4 4 3 3 5 3 4 4 4 1 4 3 3 2 2 3 MS,BS,WS Goldeneye, showy 3 3 4 4 4 4 4 3 5 3 4 4 4 1 4 4 3 4 3 3 SA,A,PP,MB,JP, MS,BS Goldenrod, Canada 2 243 334 444 444 2 4 4 4 4 3 3 A,PP,MB Goldenrod, low 2 2 4 4 3 4 4 4 4 4 4 4 5 2 3 4 5 3 3 3 A,PP,MB Goldenrod, Parry 2 2 4 4 3 4 5 4 4 4 4 4 5 2 3 4 5 3 3 4 A,PP,MB Groundsel, butterweed 4 243 334 444 344 2 3 4 4 4 3 2 A,PP,MB Helianthella, oneflower 4 335 433 443 433 1 5 3 3 2 3 3 A,PP,MB,MS Iris, German 1 4 5 2 1 4 5 5 4 2 5 4 4 5 3 1 4 4 4 4 SA,A,PP,MB, (common iris) JP,MS,BS,WS Ligusticum, Porter 4 3 2 4 3 3 3 4 3 3 4 4 4 1 3 4 3 2 3 2 A,PP,MB(con.)
428 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 19 Forbs for Seeding Range and Wildlife Habitats Lomatium, nineleaf 3 343 344 433 434 143 4 3 3 3 PP,MB,MS Lomatium, nuttall 3 3 3 3 3 3 4 3 3 2 5 2 4 1 3 4 3 2 3 3 A,PP,MB Lupine, Nevada 5 5 2 3 3 3 4 4 3 4 4 4 4 1 4 3 3 2 3 4 JP,MS,BS,WS Lupine, silky 5 5 2 3 3 3 4 4 3 4 4 4 4 1 4 3 3 3 3 4 A,PP,MB,MS Lupine, silvery 5 5 2 3 4 3 4 4 3 4 4 4 4 1 4 3 3 3 3 4 A,PP,MB,MS Medick, black 4 5 5 5 5 4 4 4 4 3 4 4 4 2 4 5 5 4 3 2 A,WM,PP,MB, JP,MS,BS Milkvetch, cicer 4 5 5 3 3 3 4 5 5 5 4 4 5 3 4 5 5 3 4 4 SA,A,WM,PP, MB,JP,MS,BS Milkvetch, Snake River 4 5 2 3 3 3 4 5 4 4 4 4 4 3 4 3 4 3 4 3 PP,MB,JP,MS, plains BS,WS Milkvetch, tall 4 5 2 3 4 4 4 4 4 2 3 4 4 3 3 3 4 4 4 2 PP,MB,JP, MS,BS Painted-cup, 2 4 1 2 2 3 4 3 2 3 4 4 2 1 2 2 1 3 3 3 SA,A,WM,PP, northwestern MB,JP,MS,BS, WS Peavine, flat 4 5 4 3 3 3 3 4 3 4 4 4 3 3 5 2 5 4 4 3 SA,A,PP,MB Peavine, perennial 4 5 4 3 3 4 4 4 4 3 3 5 3 2 3 3 4 4 4 3 SA,A,PP,MB Peavine, thickleaf 4 5 4 3 2 3 4 5 3 4 4 4 4 2 4 2 5 4 4 4 SA,A,PP,MB Peavine, Utah 4 5 3 3 2 3 4 4 3 4 5 5 3 2 4 2 4 4 4 3 SA,A,PP,MB Penstemon, Eaton 5 5 5 3 5 4 4 3 5 3 4 4 3 3 3 3 3 3 3 4 PP,MB,JP,MS, BS,WS Penstemon, littlecup 5 5 4 4 3 2 4 4 4 4 3 4 3 3 3 3 4 3 3 3 PP,MB,JP,MS, BS,WS Penstemon, low 5 5 5 3 4 3 5 5 5 2 4 4 4 3 4 3 5 3 3 3 PP,MB,JP,MS, BS,WS Penstemon, Palmer 5 5 4 5 4 4 3 3 5 4 4 4 4 5 3 4 3 2 3 4 PP,MB,JP,MS, BS,WS,BB Penstemon, Rocky 5 5 4 5 4 3 4 3 4 3 4 4 4 4 4 5 4 3 3 3 A,PP,MB,JP, Mountain MS Penstemon, Rydberg 5 543 233 443 434 324 453 2 SA,A,WM, PP,MB Penstemon, sidehill 5 5 4 4 3 3 4 4 4 2 3 3 4 3 4 3 4 3 3 3 PP,MB,JP,MS, BS,WS Penstemon, thickleaf 5 543 334 342 334 343 4 3 3 3 PP,MB,JP, MS,BS Penstemon, toadflax 5 5 4 3 2 3 4 4 4 2 4 3 4 3 4 3 3 3 3 3 PP,MB,JP,MS, BS,WS Penstemon, Wasatch 5 543 443 354 3 2 4 344 443 4 A,PP,MB,JP, MS,BS Sage, Louisiana 2 3 5 4 2 4 4 4 5 3 2 3 5 3 4 4 5 4 3 4 SA,A,WM,PP MB,JP,MS, BS,WS Sage, tarragon 2 2 4 3 1 4 4 4 3 3 2 2 4 4 3 3 4 3 3 3 SA,A Sainfoin, common 5 5 5 5 4 4 4 3 4 5 5 4 4 3 3 4 3 3 4 3 PP,MB,JP, MS,BS Salsify, vegetable-oyster 3 234 553 343 343 3 4 4 3 4 4 4 PP,MB,JP, MS,BS Solomon-plume, fat 4 3 3 2 2 3 4 4 3 2 5 4 3 1 3 3 3 5 3 2 SA,A,WM, PP,MB Sweetanise 3 3 2 3 4 3 4 4 4 4 4 4 4 2 4 4 3 4 5 3 SA,A,WM, PP,MB Table 1 (Con.) Ratingsb

- Vegetative

Early Compati- typea to

Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the

handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of adapt- species is

 Species seed seeding planting nation lishment rate lishment tence spread yield bility bility grazing ness species tion stability ance ance ability adapted

(con.)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 429

Chapter 19 Forbs for Seeding Range and Wildlife Habitats Sweetclover, white 5 5 3 5 5 5 2 2 3 3 3 4 3 3 4 5 2 4 3 4 SA,A,WM, PP,MB Sweetclover, yellow 5 535 552 234 343 345 243 4 SA,A,WM, PP,MB, JP,MS,BS,WS, Sweetroot, spreading 3 432 334 322 443 244 344 2 SA,A,WM, PP,MB Sweetvetch, Utah 2 4 3 3 3 3 5 4 3 4 4 4 4 3 4 4 3 3 3 3 PP,MB,JP,MS, BS,WS Valerian, edible 3 3 2 2 2 2 3 5 2 3 4 4 4 1 4 3 3 4 4 3 SA,A,WM,PP, MB,JP,MS Vetch, American 4 5 4 4 2 4 4 5 3 4 4 4 4 3 5 4 4 3 4 3 SA,A,WM,PP, MB,JP,MS Vetch, bramble 4 5 4 5 3 2 4 5 3 3 4 4 4 3 5 4 4 3 4 3 BS,WS,SS, BG,BB Yarrow, western 4 4 5 4 4 4 5 5 5 3 4 4 5 4 4 5 5 5 4 5 SA,A,WM,PP, MB,JP,MS,BS aKey to vegetative type: SA = Subalpine; A = Aspen-conifer; WM = Wet and semiwet meadows; PP = Ponderosa pine; MB = Mountain brush; JP = Juniper-pinyon; MS = Mountain big sagebrush; BS = Basin big sagebrush;WS = Wyoming big sagebrush; SS = Shadscale saltbush; BG = Black greasewood; BB = Blackbrush; IS = Inland saltgrass. bKey to ratings: 1 = Poor; 2 = Fair; 3 = Medium; 4 = Good; 5 = Excellent. Table 1 (Con.) Ratingsb - Vegetative Early Compati- typea to Ease of Ease of Initial Seedling Final spring Summer Toler- bility with Seed Flood Shade Range which the handling Ease of trans- Germi- estab- growth estab- Persis- Natural Forage palata- palata- ance to Evergreen- other produc- Soil toler- toler- of adapt- species is Species seed seeding planting nation lishment rate lishment tence spread yield bility bility grazing ness species tion stability ance ance ability adapted
430 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Compositae

Achillea millefolium ssp. lanulosa__

Western Yarrow

In the Intermountain West the majority of the yarrows fall into three taxonomic units: Achillea

millefolium ssp. millefolium, which was introduced

from Europe and is widely cultivated in North America,

and A. millefolium ssp. lanulosa, a native to North

America, which is divided into two varieties—var.

lanulosa (western yarrow) that occurs from the valley

floor to above timberline and var. alpicola that is

generally found above timberline (Welsh and others

1987). All three taxonomic units intergrade. Western

yarrow, a perennial member of the sunflower family,

has flowers that are borne in flat-topped cymes (fig. 1)

that are white, grayish, and barely pink, all with

yellow centers (Hermann 1966). The leaves and flowers are aromatic.

Ecological Relationships and Distribution

Western yarrow is found in Southwestern Canada,

in all States west of North Dakota, south to Northern

Mexico, and from the low plains to subalpine communities. Greatest areas of occurrence are in mountain

brush, ponderosa pine, aspen, and open timber

Figure 1—Western yarrow in a mixed

grass forb community.

types (USDA Forest Service 1937). Western yarrow

has good fire tolerance, some shade tolerance, and is

fairly resistant to drought. It grows in sandy to loam

soils that range from weakly basic to weakly acidic

(USDA Forest Service 1937; Wasser 1982). Yarrow

spreads by rhizomes and seed, and is especially useful

on disturbed or misused sites. Tolerance to grazing is

good.

Plant Culture

Western yarrow can be seeded successfully by drilling or broadcasting. Seeds are small (4.1 million per lb

[9.0 million per kg] at 100 percent purity) and should

not be planted more than 0.25 inch (6 mm) deep. Seeds

germinate uniformly, and produce vigorous seedlings.

Seed is usually marketed with about 50 percent purity

and 80 percent germination. It is recommended that

seed be stored no more than 2 years, due to loss of

viability. The species is easily transplanted. Western

yarrow can be seeded successfully in mixtures with

other species (Stevens and others 1985c).

Uses and Management

Western yarrow has fair forage value for sheep,

somewhat less for cattle, and is used by small and

large game animals (Hermann 1966; Kufeld 1973;

Kufeld and others 1973; USDA Forest Service 1937;

Wasser 1982). It provides important habitat for upland game birds. This species has good potential for

revegetation and for stabilization of disturbed sites

from big sagebrush through the subalpine zones. Western yarrow establishes and spreads well in sites occupied by annual weeds. It persists with heavy grazing,

and often recovers well by natural seeding. Areas

seeded to western yarrow should not be grazed for at

least 1 to 2 years following seeding.

Varieties and Ecotypes

There is considerable variation, and locally adapted

ecotypes should be planted.

Family Compositae

Artemisia ludoviciana ____________

Louisiana Sage (wormwood)

There are a number of subspecies and numerous

varieties and ecotypes of Louisiana sage in the Intermountain West (Holmgren and Reveal 1966; Welsh

and others 1987). Considerable differences occur among

the various subspecies, varieties, and ecotypes. Louisiana sage is an herbaceous native perennial that is

aromatic and very rhizomatous, with stems that

grow from 1 to 3 ft (30 to 91 cm) tall. Leaves are

mainly cauline, entire, lobed or pinnately incised, and

borne on the stem.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution

Louisiana sage (fig. 2) occurs throughout North

America in all vegetation types from sagebrush

through alpine and tundra zones. This species does

well on shallow as well as on deep, slightly acidic to

basic soils. It prefers well-drained soil, but also grows

in riparian communities and on other moist sites.

Louisiana sage does well on disturbed exposed soils as

well as on sites supporting some cover. As a pioneer

species, it creates microenvironments suitable for the

invasion of other species.

Plant Culture

Flowering occurs in mid to late summer, depending

on location, with seed maturing in late August to

early November. Seed can be collected by hand stripping, beating, or combining. Most native stands do not

produce an abundance of seeds on a regular basis.

Seed is cleaned by debearding and screening. There

are about 2.5 million seeds per lb (5.5 million per kg)

at 100 percent purity. Seed is generally marketed at

10 to 15 percent purity and 80 percent germination.

Seed viability declines rapidly with more than 3 to

4 years of storage.

Seed can be drill seeded or broadcast seeded, followed by very shallow soil coverage. Drill seeding

requires a purity of greater than 40 percent for seed to

flow through most seeding units. Seed can be broadcast with as low as 10 percent purity, depending on the

type of broadcaster used. Seed that is fall broadcast on

disturbed soil generally does not require mechanical

coverage because natural soil sluff is sufficient to bury

the seed. Louisiana sage can be seeded as a single

species or as a component of a mixture. It also establishes easily from sprigs.

Uses and Management

Louisiana sage plays an important roll in providing

soil cover and stabilization in areas where it is adapted

(Hermann 1966; McCulloch 1973). It can be an early

invader on disturbed sites including mine spoils

(Monsen 1975; Monsen and Plummer 1978). Where it

is seeded or becomes established, it acts as a nursery

crop that allows establishment of other species. Ellison

(1951) considered it an important invader of overgrazed sheep range and as a soil stabilizer on the

Wasatch Plateau in central Utah. It can be used to

stabilize riparian disturbances including dry meadows and areas subjected to infrequent flooding.

Livestock, small and large game, and rodents use

this forb (Hermann 1966; Kufeld and others 1973;

McCulloch 1973). It has considerable tolerance to

grazing and trampling; however, when occurring on

shallow soils or disturbed sites, it should not receive

Figure 2—Louisiana sage.

extensive use or trampling from grazing animals.

Stand density fluctuates, particularly at higher elevations where the species may be suppressed by overstory shading. It recovers well after fires and can be

used to prevent the entry of weeds by providing ground

cover after a disturbance.

Varieties and Ecotypes

There is one variety. ‘Summit’ was selected for its

its ability (1) to establish from sprig planting and

direct seeding, (2) increase or spread rapidly by rooting, (3) provide quick cover and soil stabilization, (4)

grow on harsh sites, and (5) create microenvironments

suitable for the invasion of other species (Stranathan

and Monsen 1986). ‘Summit’ is particularly useful as

a ground cover or erosion control species for sites

above 5,000 ft (1,500 m) elevation.

Family Compositae

Aster chilensis__________________

Pacific Aster

Pacific aster is a native perennial of the compositae

family. It is described by Harrington (1964) as developing short to long root stocks and stems that are

usually less than 2.5 ft (0.8 m) tall, with long, linear

pubescent leaves. Few to numerous heads occur with

blue, violet, or rarely white flowers. Seeds or achenes

are usually hairy with many white pappus bristles.

Plants bloom from July to September. This is an

extremely diverse species. Welsh and others (1987)

consider Pacific aster a generalized taxon with no

well-defined diagnostic features. Hitchcock and others (1955) recognized three subspecies: spp. hallii,

which occurs in the Oregon Cascades, and in Washington west to the coast, spp. chilensis, which is found
432 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

throughout the Cascades and Sierras; and spp.

adscendens, which occurs in the Great Basin and

eastern Oregon, and on the Snake River plains.

Ecological Relationships and Distribution

Pacific aster (fig. 3) is a native perennial that can

spread rapidly by seed or rhizomes. This forb occurs

throughout the Intermountain region at elevations

between 5,000 and 9,500 ft (1,500 to 2,700 m). It

occupies areas from the sagebrush foothills to the

subalpine zone (Harrington 1964). It is most commonly found on dry, open sites in the sagebrush-grass

and pinyon-juniper types.

Plant Culture

Pacific aster can be established by direct seeding,

sprigging, and with nursery or field-grown transplants (Stevens and others 1985c). Seed is best planted

by broadcasting; however, if it is cleaned to at least

50 percent purity, it can be drilled. Picker or thimble

seeders can be used to plant the uncleaned, trashy

seed. Special care must be taken to ensure that seeds

are not planted more than 0.25 inch (6 mm) deep. To

ensure that viable seed is planted, seed should not be

stored more than 3 years. Harvested seed usually

consists of the achene and attached pappus. The

pappus is removed by gentle hammermilling, or

with a debearder. Seed is usually not cleaned to

more than 50 percent purity, and most seed is sold

at 10 to 14 percent purity. At 100 percent purity, a

pound of seed consists of just over 2.5 million seeds

(5.5 million per kg).

Seeds of Pacific aster ripen throughout the late

summer and fall. Not all seeds ripen uniformly; consequently, considerable debris and immature seeds are

often collected. Seeds can be separated from the debris

with special cleaning equipment. Seed should be purchased based on the percent of viable seed.

Seeds of Pacific aster germinate readily. Nearly all

viable seeds germinate within 15 days. Seedlings

appear early. They are able to compete with most

seeded herbs and persist under adverse conditions.

Young plants usually do not attain large stature the

first growing season, but they do form an extensive

root system.

Although the seeds are small, they establish well

with only minimum seedbed preparation if they are

seeded with the pappus attached. Seeding with the

pappus on, however, is very difficult.

Neither livestock grazing nor rodent and insect use

have been observed to damage new seedings. New

plantings are not heavily used by rabbits and can be

established where rabbit populations are high. The

plant can also be seeded on unstable slopes because

Figure 3—Pacific aster in full bloom.

small plants do not attract heavy use and provide

excellent ground cover.

Uses and Management

Pacific aster is one of the first forbs to “green up” in

the spring, causing it to be highly sought by livestock

and big game. Game and livestock do, however, make

some use of this species during all seasons because the

plants retain some green material most of the year

(Hermann 1966; Kufeld and others 1973; Plummer

and others 1968). Hermann (1966) and USDA Forest

Service (1937) reported that Engelmann aster is more

palatable than Pacific aster to most animals. Our

observations indicate similar differences, but recognize

that Pacific aster receives moderate to heavy seasonal grazing.

Pacific aster is one of the most widely distributed

native forbs within the Intermountain region. Selections obtained from the low foothill sagebrush communities are well adapted to semiarid situations. Pacific

aster is one of a few forbs that can be easily seeded

within the arid region occupied by big sagebrush. It

persists well, furnishes good midsummer forage, and

can be used to control cheatgrass. It recovers well from

wildfires and spreads naturally.

To date, Pacific aster has normally been seeded as

a minor component of most seed mixtures, usually at

0.25 lb seed per acre (0.3 kg per ha). However, it can be

seeded at higher rates of 2 to 5 lb per acre (2.2 to 5.6 kg

per ha) to provide more summer forage. Pacific aster

does not restrict seedling establishment of associated

herbs, and its own seedlings are not impaired by the

presence of other seeded species. Consequently, increasing the seeding rate of this plant usually results

in a corresponding increase in plant density.

The strong rhizomes and root system enables the

species to be very useful in stabilization of disturbed
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 433

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

areas. Pacific aster is a useful species for planting

exposed sites with shallow soils, particularly midwinter game ranges. It can be successfully seeded on

areas that are difficult to treat with mechanical equipment. It is well suited to infertile, rocky soils and sites

subject to erosion.

Pacific aster is a useful pioneer species and nurse

crop. The plant is able to occupy harsh sites and

improve surface stability, enhancing seedbed conditions for other species. New seedlings are able to

establish amid the seemingly dense ground cover

provided by this forb. It is not unusual to observe

seedlings of other plants establishing and surviving

within a clump of Pacific aster. Consequently, the

plant is used to furnish cover on steep slopes where

pinyon and juniper have been chained. The plant is

also valuable for seeding or transplanting roadways,

mines, and other major disturbances. Areas seeded to

Pacific aster should not be grazed for at least two

growing seasons following planting. Creeping rhizomes enable this species to withstand considerable

grazing and trampling once plants are established.

Pacific aster has only been sparingly used in controlled grazing studies, but it is apparent that this

species can be managed to provide midsummer and

winter forage. In addition, the plant can be seeded to

control cheatgrass and to furnish needed cover on

critical semiarid ranges.

Varieties and Ecotypes

There are no releases.

Family Compositae

Aster glaucodes ________________

Blueleaf Aster

Blueleaf aster (fig. 4), a native perennial, has about

the same forage value and seed characteristics as

Pacific aster. However, its seeds are more than three

times as large (500,000 per lb [1.1 million per kg]), and

the species is more widely distribted, commonly occurring in all vegetative types from salt desert shrublands

and saline seeps to spruce-fir forests. It furnishes an

open spreading clump, with short seed stalks and wide

leaves. It is well adapted to harsh sites (fig. 5) and

tends to exist alone. This species is better adapted to

calcareous soils and saline conditions than Pacific

aster. When seeded with principal forage grasses, it

does not provide a dominant cover. It is grazed by big

game and livestock, but it is not heavily used. Like

Pacific aster it persists amid heavy infestation of

grasshoppers when other succulent herbs are usually

consumed. If seed of both species were more available,

they would be recommended for planting range, watershed, and wildlife habitats.

Family Fabaceae

Astragalus cicer ________________

Cicer Milkvetch

A native of Eurasia, cicer milkvetch is a perennial

member of the pea or legume family. It has vigorous

creeping rhizomes and a short taproot. The leafy,

succulent, decumbent stems are fairly large in diameter. They grow to about 5.5 ft (1.7m) in length, and

originate from crown and rhizome buds. Flowers are

born in compact heads and are pale yellow to white.

Blooming occurs in midsummer, and seeds mature in

late fall. The inflated, bladder-shaped, leathery black

pods contain several seeds. Seed pods persist on the

plant into the winter.

Figure 4—Blueleaf aster at seed maturity. This

stand was established by broadcast seeding on a

raw roadfill in the mountain brush type.

Figure 5—Blueleaf aster stabilizing a severely

eroded cut.
434 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution

Cicer milkvetch (fig. 6) is an introduced, long-lived

perennial from northern Europe. It is bloat free and

does not accumulate toxic alkaloids (Davis 1982a;

Rumbaugh 1984; Rumbaugh and Townsend 1985;

USDA Soil Conservation Service 1968, 1972a; Williams

and others 1976). It is safe for grazing (Rumbaugh

1983).

This species has vigorous spreading rhizomes, good

frost and drought tolerance, and produces considerable succulent forage with a high protein content and

a low crude fiber level (Davis 1982a). It grows in

slightly acidic and basic soils, and does especially well

in soils derived from limestone. This forb establishes

and grows on thin, infertile soils and on disturbed sites.

Spring growth starts about 2 weeks later than alfalfa

and continues about 2 weeks longer in the fall. Foliage

is green and succulent throughout the summer and

fall months. Basal leaves remain green through winter. This species has more frost tolerance than alfalfa

and remarkable resistance to insects and disease.

Cicer milkvetch requires at least 13 inches (33 cm)

of annual precipitation, and does especially well with

16 inches (40 cm) or more. It is tolerant of water tables

within 3 ft (91 cm) of the surface and does fairly well

on wet sites. It has some shade tolerance, and is well

adapted to sagebrush-grass and pinyon-juniper sites

with sufficient precipitation. It also does well in the

mountain brush type (Plummer and others 1955;

Rumbaugh and Townsend 1985) and in openings in

aspen and subalpine areas.

Plant Culture

Cicer milkvetch can be broadcast or drilled seeded.

There are about 113,000 seeds per lb (249,000 per kg).

Figure 6—Cicer milkvetch growing in association with smooth brome, Gambel oak, and

mountain big sagebrush.

Seed should be inoculated with appropriate inoculant

before seeding (Carleton and others 1971; Hafenrichter

and others 1968), and should not be planted more than

0.25 inch (6.4 mm) deep. Because of its hard seedcoat

and dormancy, seeding should be done in the fall.

Scarification can improve seeding success. However,

once seeds are scarified they will not retain viability

over extended periods.

Seed of cicer milkvetch has been stored for up to 14

years without any loss of viability (USDA Soil Conservation Service 1968a). This species is easily transplanted. Cicer milkvetch has done well seeded in

mixtures and has been successfully interseeded into

various native and introduced grass communities

(Rumbaugh and others 1981; Stevens and others

1981b). Because of its lower seedling vigor (Rumbaugh

1984), cicer milkvetch does best when seeded in well

prepared seedbeds that are relatively free of competition. This species does not reach full productivity

until the third year following seeding. Once established, cicer milkvetch is very competitive and longlived. It has been demonstrated that cicer milkvetch

can stimulate and increase forage production of

associated species (Johnson and others 1983).

Uses and Management

Seed of cicer milkvetch is produced commercially

and is generally available. Acceptable purity and germination for commercial seed is 95 and 85 percent.

Cicer milkvetch is palatable to livestock and big game,

and seeds are eaten by small birds, deer, rabbits, sagegrouse, and pheasants (Plummer and others 1968;

USDA Soil Conservation Service 1968, 1972a; Wasser

1982). Once established, this species has excellent

grazing tolerance. This species can be classified as a

semievergreen because basal leaves are green all

year.

The rhizomatous characteristics of cicer milkvetch,

along with its semiprostrate nature and profuse flowering, make it ideal for use around summer homes, on

disturbed sites, in campgrounds and parks, and in

areas with high aesthetic value.

This species is useful as a forage and conservation

plant, providing excellent ground cover and stabilizing eroding sites. It can be seeded for intensive

pasture management. The plant has considerable value

for seeding semiwet sites that are used by upland

game birds for summer forage. It also provides winter

and nesting cover for upland game birds.

Varieties and Ecotypes

‘Lutana’ was released in 1970 and has the characteristics described for the species. ‘Monarch’ was selected

for its superior seedling emergence. ‘Oxley’ is a 1971

Canadian release (Rumbaugh and Townsend 1985).
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Compositae

Balsamorhiza sagittata ___________

Arrowleaf Balsamroot

Arrowleaf balsamroot (fig. 7) is a broadleaf perennial with a deep, woody taproot. It is the most abundant balsamroot in the Intermountain area. Simple

deltoid or sagittate leaves are entirely gray-green and

heavily pubescent. Flowering stems produce a few

reduced leaves. Flowers or heads are usually borne

solitary on extended seed stalks. Both disk and ray

flowers are produced, but they lack a pappus (Welsh

and others 1987). Flowering occurs in early April, and

seed matures in late May and early June (Harrington

1964). Leaves dry up between late June and the end of

July.

Ecological Relationships and Distribution

A native perennial forb, arrowleaf balsamroot can

be found growing on well-drained silty to loamy soils

in sagebrush, mountain brush, and ponderosa pine

communities, and open slopes in aspen and coniferous forests. This forb is usually found at elevations

between 1,000 to 9,600 ft (305 to 2,900 m) (Harrington

1964; Wasser 1982). It occurs in solid stands or mixed

communities. Arrowleaf balsamroot is the most widespread species of balsamroot. It ranges from the Sierras east to Colorado and north to southern Canada on

acidic and alkaline soils (Hitchcock and others 1955;

Plummer 1977). It dominates many extensive foothill

areas, often growing as nearly pure stands (USDA

Forest Service 1937).

Arrowleaf balsamroot is intolerant of shallow water

tables, but withstands brief periods of soil saturation.

It is strongly drought resistant, winter hardy, and

tolerant of semishade, open sunlight, and of grazing

(Wasser 1982).

Plant Culture

Arrowleaf balsamroot is one of the first plants to

initiate growth in the early spring. Plants flower in

April and usually attain maximum stature in late

April or May, and seeds ripen in May and early June.

Seed heads are formed individually on extended solitary peduncles, and if plants are not heavily grazed,

large quantities of seed can be produced. Seeds ripen

uniformly and are large, with just over 55,000 per lb

(120,000 per kg). Seed is easily harvested by hand

stripping or beating. Extensive areas of level terrain

and dense stands can be harvested with a combine or

reel-mounted harvester (Plummer and others 1968;

Stevens and others 1985c). Seed can also be produced

under cultivation (fig. 7).

Figure 7—Seed increase planting of arrowleaf

balsamroot.

Seeds lack a fluffy or bristly pappus common to

many other compositae. Seeds are long, smooth, and

easily cleaned. Wildland stands are often infested

with seed damaging insects, and entire seed crops can

be damaged. Seed should be carefully inspected prior

to collection. Plummer and others (1968) suggest treating seed production areas with insecticides to reduce

seed predation However, treatment must be timed to

avoid elimination of pollinators. Seedin storage should

be treated to prevent insect damage. Seed stored

properly in an open warehouse retained good viability

after 5 years (Stevens and Jorgensen 1994).

Mature seed will remain on the plant for a short

time, but is readily eaten by rodents (Everett and

others 1978b). Immature and insect-damaged seed

will persist on the plant for longer periods. Too often,

the persisting poor-quality seed is mistakenly harvested and marketed. Immature and damaged seed

may often be normal size. However, it is light weight

and can be separated from fully developed seed by

gravity, wind separation, or flotation seed cleaning

techniques. Good quality lots will have 95 percent

purity and at least 40 percent germination.

Arrowleaf balsamroot seeds can be drill seeded quite

easily with conventional equipment. Seeds should not

be placed more than 0.33 to 0.5 inch (8 to 13 mm) deep.

Seeds can be broadcast planted but only if they are

covered. Because of seed dormancy and rodent preference for this species, late fall seeding is recommended

(Eddleman 1978; Everett and others 1978b; McDonough

1976; Young and Evans 1979). Arrowleaf balsamroot

seeds are often planted alone or with slower developing herbs and shrubs. This species should not be

seeded directly with more aggressive developing

herbs, but should be planted in separate rows or
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

broadcast onto sites free of competition. If planted

with grasses or other broadleaf herbs, arrowleaf

balsamroot plants may survive, but would require

10 years or longer to reach maturity.

Although seedlings germinate and emerge quickly,

young plants develop slowly. Even under ideal conditions thick, dense stands usually do not develop

from artificial seedings. Causes for low return are

not fully known, but insects and rodents can cause

considerable damage. Seeded plants may produce

only two to three small leaves for a number of years

following planting. Plants are slow to increase in

stature, but they are extremely hardy and persistent.

Uses and Management

Arrowleaf balsamroot is an important forage plant

for deer, elk, cattle, and sheep, especially on spring

ranges where it greens up early (Harniss and Wright

1982; Hermann 1966; Holecheck and others 1982;

Kufeld 1973; Kufeld and others 1973; Mueggler and

Stewart 1980; Plummer and others 1968). The plant

is also grazed later in the season with considerable

use made of the heads during and following flowering. In many plant communities, a major portion of

total forage production is arrowleaf balsamroot.

Plantings of arrowleaf balsamroot do well when

seeded within areas of its natural occurrence. Attempts to extend or seed this species on areas outside

of its natural range or on mine or roadway disturbances have been unsuccessful. This forb is encountered in low, semiarid conditions with Wyoming big

sagebrush, but has been difficult to seed in these

circumstances, usually because of using unadapted

ecotypes, heavy grazing, and competitive effects of

associated species. The most successful seedings have

occurred in the lower elevation mountains in sagebrush and mountain brush communities.

Arrowleaf balsamroot can be used to improve forage

conditions on game and livestock spring and summer

ranges. It undoubtedly has been eliminated from some

situations, but is extremely hardy and persistent even

under heavy use.

Because of a deep fleshy taproot, arrowleaf balsamroot can withstand drought, trampling, fire and considerable grazing pressure. Natural reproduction will

occur if seeds are allowed to mature. Seeded areas

should not be grazed for at least two growing seasons

following planting. New plants are slow to mature,

requiring 3 to 4 years to flower on the most preferred

sites, and 7 to 8 or even 10 years on the lower

precipitation sites.

Varieties and Ecotypes

There are no releases.

Like most other natives, arrowleaf balsamroot

grows on many different soil types and in many precipitation zones. Consequently, various ecotypes likely

occur. Considerable differences are noted among populations in regard to palatability, seed production, and

drought tolerance. Seeds are normally produced each

year from upper elevation sites where precipitation is

more reliable; however, seeds should be acquired from

sites similar to the area proposed for treatment. The

genus also appears to lack genetic barriers to hybridization, and intergradation occurs when any two taxa

exist together (Welsh and others 1987).

Family Compositae

Balsamorhiza macrophylla________

Cutleaf Balsamroot

Cutleaf balsamroot has large leaves 12 to 24 inches

(30 to 61 cm) long that are divided into broad, entire,

or few-toothed segments 2 to 5 inches (5 to 13 cm) long

(Welsh and others 1987). The leaves have long, soft

hairs and a somewhat unpleasant odor.

Cutleaf balsamroot generally occurs at slightly

higher elevations and in more moist conditions than

arrowleaf balsamroot. It is not distributed as widely

as arrowleaf balsamroot and is less palatable to grazing animals. Cutleaf balsamroot ranges from northern

Utah and southeastern Idaho to western Wyoming

and southwestern Montana (Hitchcock and others

1955). Seed collection, cleaning, and storage as described for arrowleaf balsamroot. Seeds are somewhat

larger than those of arrowleaf balsamroot with

about 33,000 per lb (73,000 per kg). Seed can only be

stored for 3 years without significant loss of viability

(Stevens and Jorgensen 1994). Cutleaf balsamroot is

better adapted to areas of slightly higher elevations

and more moist conditions than arrowleaf balsamroot.

Neither species should be seeded in place of the other.

Site differences are not easy to differentiate.

Family Compositae

Balsamorhiza hookeri var.

neglecta _______________________

Hairy Balsamroot

Compared to arrowleaf balsamroot, hairy balsamroot has smaller leaves 4 to 12 inches (10 to 31 cm) long

that are divided into narrow segments 0.5 to

2 inches (1 to 5 cm) long (Welsh and others 1987).

These segments are often divided a second time into

even smaller segments. The leaves are covered with

coarse, short, stiff hairs.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

This species has a more limited distribution than

arrowleaf or cutleaf balsamroot, occurring from eastern Oregon to southwestern Nevada (Hitchcock and

others 1955). Welsh and others (1987) report two

varieties of Hooker balsamroot are found in Utah,

these occur in separate regions, but grow on a variety

of sites.

Seed collection, cleaning, and storage are as described

for arrowleaf balsamroot. Seed production is much lower

than for arrowleaf or cutleaf balsamroot. In addition,

plants usually do not exist in large dense patches, but are

scattered and intermixed with other herbs.

Plants usually grow as an understory with shrubs,

and have been useful in seeding pinyon-juniper and

mountain brush sites. Sources that grow in drier

circumstances offer opportunity to select and develop

materials adapted to sagebrush and salt desert

shrublands.

Family Fabaceae

Coronilla varia __________________

Crownvetch

Crownvetch (fig. 8) is an introduced perennial legume with strong rhizomes and a deep taproot (Leffel

1973). Varigated white to purple flowers start to blossom in early summer and blooming continues for 4 to

5 weeks. Seed matures in early fall in pencil-like pods

that break into segments as they dry. Each segment

contains one yellow to dark red, rod-shaped seed.

Flowers are arranged in an umbrella or crown-like

fashion. Basal leaves are green early in the spring and

remain green into the winter.

Ecological Relationships and Distribution

Crownvetch is a long-lived, cold-tolerant perennial

from the Mediterranean region. This species is not a

true vetch because it does not have tendrils for climbing. It does, however, tend to climb if supported by

shrubs like mountain big sagebrush, Gambel oak, or

serviceberry. This forb does fairly well with 21 inches

(530 mm) annual precipitation and thrives with over

30 inches (760 mm) of precipitation. Crownvetch exhibits some shade and fire tolerance. It prefers slightly

acidic soils but has also done well on slightly basic

calcareous soils. It does not persist on poorly drained

soils.

Crownvetch can be seeded in mountain brush, ponderosa pine, and aspen communities. It persists well

at lower elevations once established. Although it performs best when grown on fertile soils, it demonstrates

unusual ability to persist, spread, and remain productive when planted on mine disturbances and sites

lacking top soil. It is especially useful for stabilizing

erosive sites. This species is very competitive. Over time

it will dominate an area and exclude other species.

Figure 8—Crownvetch stabilizing a road cut.

Plant Culture

Seeds are produced in segmented pods. Cleaned

seeds are separated from the pod, and should be

inoculated with the proper rhizobium prior to seeding (Magness and others 1971; Wasser 1982). There

are about 135,000 seeds per lb (298,000 per kg) at

100 percent purity. Acceptable purity and germination for commercial seed is 95 and 75 percent. Seeding

is best accomplished in the fall on a well-prepared

seedbed. Stratification requirements can be overcome

with fall seeding. Seed can be drilled or broadcast, but

it should not be covered deeper than 0.25 inch (6 mm).

This forb can be seeded with other species. When

seeded with a grass mixture, it is best seeded separately

through the legume seedbox. Excellent establishment

can be obtained from transplanting or sprigging. Seeding at a rate of 2 to 4 lb per acre (2.2 to 4.5 kg per ha)

is satisfactory when seeded in a mixture.

Uses and Management

Crownvetch produces strong, spreading, fleshy rhizomes. This extensive system enables crownvetch to

be an excellent soil stabilizer and an aggressive

spreader. It has been used to stabilize watershed and

mine disturbances at locations receiving over 16 inches

(40 cm) of annual precipitation. The plant is characterized as having a succulent dark green appearance,

profuse flowering, and excellent ground cover characteristics. These features make crownvetch a prime

landscape species for summer home developments,

roadways, campgrounds, and areas of high recreational use.

Palatable forage is produced by crownvetch for all

classes of livestock and wildlife (Wasser 1982), with

little or no bloat hazard (Leffel 1973; USDA Soil
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Conservation Service 1978). Deer, elk, and livestock

will paw through snow to find and use its semievergreen forage. Birds and rabbits use crownvetch for

food and cover. Crownvetch has some poisonous characteristics (Shenk and others 1976). However, when

used in conjunction with other species, poisoning has

not occurred (Shultz 1984). Crownvetch is normally

seeded as a component of a mixture to enhance forage

production and quality. This species establishes

slowly, and production is low the first or second year

following seeding. Once established, it can withstand

considerable grazing and trampling. It recovers from

close grazing, but does not have the regrowth capabilities of alfalfa. When seeded in mixtures with other

species, it will spread vegetatively, and will tend to

dominate a community.

Varieties and Ecotypes

‘Emerald’, ‘Penngift’, and ‘Chemung’ have done well

in mountain big sagebrush, mountain brush, and

aspen communities. Emerald is the smallest in stature and produces less forage, but it is a more aggressive spreader.

Family Geraniceae

Geranium richardsonii ___________

Richardson Geranium

A member of the geranium family, Richardson geranium (fig. 9) is a perennial native forb, with flowers

that are generally white with pinkish or dark veins.

Stems are single or few and somewhat hairy and

viscid. Leaves are large but thin. Flowering occurs in

July and early August, and seed matures in September. The foliage gives off a distinctive geranium odor

when crushed (USDA Forest Service 1937).

Figure 9—Richardson geranium.

Ecological Relationships and Distribution

Richardson geranium occurs in openings throughout the coniferous forest as an understory species in

aspen and subalpine meadows (Judd 1962; USDA

Forest Service 1937). It is the most widely distributed

geranium in North America (Hermann 1966). The

species is not restricted to any particular soil type, but

usually occurs on fairly moist gravelly or sandy loams.

However, it can be found on drier granitic soils and

heavy clay loams (Judd 1962; USDA Forest Service

1937).

Plant Culture

Seed is generally produced yearly if plants are not

grazed. Seed can be collected by beating it into a

hopper or with a mechanical seed beater mounted on

a vehicle. Seed is extracted from the collected material

by hammermilling or with a debearder (Stevens and

others 1985c). Seed can be stored for at least 5 years

without substantial loss of viability. Acceptable germination is 60 percent. There are about 65,000 seeds

per lb (143,000 per kg) at 100 percent purity. Seed can

be cleaned to 60 percent or higher purity. Seed can be

drilled or broadcast seeded on a prepared seedbed,

preferably in the fall. Seeding depth should not exceed

0.25 inch (6 mm). This species can be seeded singly or

as a component of a mixture of other forbs and grasses.

It demonstrates excellent ability to spread by natural

seeding. Seeds are small and can establish on a loose

seedbed or on sites where litter has accumulated.

Uses and Management

Forage value for cattle and sheep is rated good to

excellent during early growth stages, and poor to good

later in the season. Forage value for elk (Kufeld 1973)

and deer (Kufeld and others 1973), is greater than for

livestock (Hermann 1966; Judd 1962; USDA Forest

Service 1937).

The species develops a stout, woody root system that

enables it to stabilize soil and withstand considerable

grazing, trampling, and some drought. Seedings on

fertile soils have been most successful. Attempting to

seed or use this species on sites where it does not

normally occur is not recommended. However, this

forb spreads naturally and has successfully invaded

roadway disturbances. Richardson geranium can persist with moderate to heavy grazing. Plants appear to

be long-lived and spread well from natural seeding.

Newly seeded stands should not be grazed for at least

two growing seasons following planting. This species

frequently increases with proper management. It persists well with seeded grasses, including smooth brome

and intermediate wheatgrass. It is useful for adding

diverity to mixtures, especially where shade from

overstory species exists. Limited seed supplies restrict

the use of this species.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Varieties and Ecotypes

There are no releases.

Family Geraniceae

Geranium viscosissimum_________

Sticky Geranium

This species occurs in the same climatic and edaphic

areas as Richardson geranium. Its forage value for

cattle is rated from fair to good and from good to

excellent for sheep, deer, and elk (Buchanan and

others 1972; Kufeld 1973; Kufeld and others 1973;

Mueggler and Stewart 1980). Seed is collected, cleaned,

stored, and seeded in the same manner as described

for Richardson geranium. Seeds of sticky geranium

are a little larger with about 52,000 per lb (115,000 per

kg). These two species hybridize on the Wasatch Plateau of central Utah (fig. 10) (USDA Forest Service

1937).

Sticky geranium (fig. 11) can be distinguished from

Richardson geranium by its multiple, erect stems

bearing deep purple to pink colored flowers, and its

usually sticky and glandular herbage.

Varieties and Ecotypes

There are no releases.

Family Fabaceae

Hedysarum boreale ______________

Utah Sweetvetch

Utah or northern sweetvetch (fig. 12) is a native

perennial member of the legume or pea family that

grows 10 to 30 inches (25 to 76 cm) tall and 30 to 80

Figure 10—Hybrid of Richardson and sticky

geranium.

Figure 11—Sticky geranium.

inches (76 to 203 cm) wide. Blossoms are bright rosepink to purple and very showy. Leaves are green

above and grayish on the underside. Growth begins

early in spring and provides a considerable amount of

early spring forage that is highly palatable to both big

game and domestic livestock. An abundance of forage is provided with some basal leaves remaining

green throughout the winter (Rumbaugh 1983).

This species produces a tap root (Welsh and others

1987). Some strains are rhizomatous (Ford and others

1984). This species has nitrogen fixing capabilities

(Ford 1988; Ford and others 1989; Redente and Reeves

1981). Abundant seed is produced most years. Considerable variation occurs within the species, and there

are a number of subspecies and varieties (Ford 1988;

Northstrom and Welsh 1970, 1979; Rollins 1940).

Utah sweetvetch is the most abundant sweetvetch in

the Intermountain West.

Figure 12—Utah sweetvetch.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Ecological Relationships and Distribution

Utah sweetvetch occurs in most Western States

(Ford 1988). It has fair shade tolerance (Rumbaugh

and Townsend 1985; Wasser 1982) and can be found

in subalpine, aspen, mountain brush, ponderosa pine,

pinyon-juniper, and big sagebrush types. In northern

Utah it has been found in the shadscale saltbush type

(Plummer and others 1968). Sweetvetch grows with

10 to 36 inches (25 to 91 cm) of annual precipitation

on acidic or basic soils ranging from sands to heavy

clays (Plummer 1977), making it well adapted to

semiarid areas of the Intermountain West (Redente

1980).

This species has a stout, branched, woody taproot,

thus it can take advantage of deep soil moisture,

resulting in considerable drought resistance and winter hardiness. Rhizomatous accessions have been reported (Ford and others 1984, 1989; McKell and others

1979; Plummer and others 1968; Rumbaugh 1984).

Plant Culture

Flowering occurs in late May and June. Seeds mature in late July and early August. A segmented

loment or pod is formed (fig. 13) that can be harvested

by hand or with a combine. Seed has been successfully

produced under cultivation. Seeds can be stored

and seeded in or out of the pod. Seeds should be

treated with an insecticide to control insect damage.

There are about 33,000 seeds per lb (73,000 per kg) at

100 percent purity. Acceptable purity and germination in the commercial market is 90 and 60 percent. By

extracting the seed from the pod, unfilled, damaged,

and immature seed can be removed. Seed is extracted

from the pod with a debearder or Dybvig. Heat and

mechanical breakage resulting from improper use of

Figure 13—Segmented loment of Utah sweetvetch.

either machine can reduce viability of the seed lot.

Seed can be stored up to 5 years without any significant loss in viability (Stevens and Jorgensen 1994).

Good results can be expected from direct seeding.

Seeding should occur in the fall or early winter, with

the seed covered at least 0.13 inch (3 mm) but not more

than 0.38 inch (10 mm) of soil. Seeds should be inoculated with the appropriate inoculant (Redente and

Reeves 1981; Rumbaugh and Johnson 1984). Seed can

be drilled or broadcast. Utah sweetvetch should not be

seeded where its seedlings are in direct competition

with seedlings of aggressive species. If drilled, this

forb should be seeded in rows separate from grasses.

Utah sweetvetch does not produce strong seedlings.

Germination is slow and seedling emergence follows

that of many other species. Seedlings and young plants

also develop slowly. Flowering does not occur until the

third or fourth year following planting. Once established, this species is very persistent, particularly

when seeded with other herbs. Plants can be established by transplanting in the early spring (Institute

for Land Rehabilitation 1979).

Uses and Management

Livestock and big game make considerable use of

Utah sweetvetch (Hermann 1966; Plummer and others 1968; Rumbaugh 1984; USDA Forest Service

1937; Wasser 1982). Spring growth provides succulent

forage. Considerable green herbage is retained throughout the growing season, and basal leaves remain green

through the winter (Redente 1980; Rumbaugh 1984).

This species is thought to be a bloat-safe legume

(Rumbaugh and Townsend 1985). It has considerable

potential for improving big game and livestock ranges,

and for stabilization of disturbed and eroding areas

(Plummer and others 1968; Redente and Reeves 1981).

Seeded areas should not be grazed for at least 2 years

following seeding. Once established, this species can

withstand considerable use; however, continual use

has reduced its density in some areas.

Varieties and Ecotypes

‘Timp’ exhibits excellent growth rate, seed production, and nitrogen fixing capacity. This variety is fairly

widely adapted.

Family Compositae

Helianthella uniflora _____________

Oneflower Helianthella

There are eight species of Helianthella in North

America (Weber 1952), with oneflower helianthella

having the widest distribution. Plants are upright

with erect heads. Seeds are flat and covered with
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

hair, whereas most sunflower seeds are thick and

lack pubescence. Oneflower helianthella produces yellow flowers in late May and early June, and seeds

mature in July (Hermann 1966).

Ecological Relationships and Distribution

Oneflower helianthella (fig. 14) ranges from Montana

to Oregon and south to New Mexico, mainly in mountain brush, aspen, spruce-fir, and ponderosa pine

types (Hermann 1966; USDA Forest Service 1937). It

does well in full sunlight and partial shade, growing on

dry hillsides, open woods, and intermixed in various

shrub communities.

Plant Culture

Seed matures in July and can be collected by hand,

stripping, mechanical beating, or combining. Seed is

easily cleaned, by screening to remove debris. Seeds

do not remain viable for more than 2 years (Stevens

and Jorgensen 1994). There are just over 52,000 seeds

per lb (115,000 per kg) at 100 percent purity.

Direct seeding by drilling or broadcasting has worked

well. When seeding oneflower helianthella, seed should

be covered no more than 0.38 inch (10 mm). Germination is rapid, and strong seedlings develop quickly.

This species does well seeded in mixtures with other

herbs. Reproduction from new seedings usually

occurs within 3 to 4 years after the initial planting.

Uses and Management

The leaves, flowers, and more tender portions of the

stems of oneflower helianthella are eaten by big game

and livestock. Palatability is rated as poor to fair for

cattle and elk, and fair to good for sheep and deer

(Kufeld and others 1973; Mueggler and Stewart 1980).

This perennial provides considerable forage in the

spring and through the summer.

Oneflower helianthella can be seeded in mixtures

with other herbs to furnish diversity and forage. The

plant grows on a variety of sites, occupying harsh open

conditions, yet also doing well on moist, fertile soils in

some shade. When seeded with native and introduced

grasses, it persists well and provides useful forage. It

will spread to colonize open disturbances, and can be

used to beautify recreational sites and roadways. It

usually does not provide a dense ground cover, but

mixtures of oneflower helianthella with other herbs

are useful for controlling erosion.

New seedlings should receive at least two growing

seasons of nonuse following planting. Selective grazing of this species does not occur, and it can be easily

maintained in a mixture with other herbs on summer

ranges.

Figure 14—Oneflower helianthella in full bloom.

Varieties and Ecotypes

There are no releases.

Family Umbelliferae

Heracleum lanatum ______________

Cow Parsnip

There are about 60 species of Heracleum throughout the world. However, only one, cow parsnip, occurs

in North America (Hitchcock and others 1961). Cow

parsnip forms a compound umbel (fig. 15), with flowers occurring at the ends of long, hollow stalks. Plants

are tall, often reaching 7 ft (2.1 m). The white to light

yellow flowers bloom in July and early August. Seed

matures in late August and early September. Roots

are woody, short, jointed, and aromatic. Leaves are

divided into three very large, irregularly toothed

leaflets, each 4 to 10 inches (10 to 25 cm) long.

Ecological Relationships and Distribution

Cow parsnip is found from Alaska, eastward across

Canada, and south through the Western States and to

Georgia. In the Intermountain West, cow parsnip

occurs in aspen, spruce-fir, and subalpine areas, and

in meadows, with rich, loamy soils. This species prefer

some shade and exists in open woodlands.

Plant Culture

Cow parsnip seeds are flat, obovate in shape, and

fairly large, with just over 44,000 per lb (97,000 per kg).

Collection of seed is best accomplished by hand stripping or beating. If allowed to mature, good seed crops

are generally produced yearly. Seeds are easily

cleaned by screening to separate them from debris.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Germination is generally low, and viability is not

maintained for much more than 2 years (Stevens and

Jorgensen 1994).

Seed can be broadcast or drill seeded, preferably in

autumn before snowfall. Seed should not be covered

more than 0.5 inch (1.3 cm) deep. Seeds are rather

large and are difficult to seed in mixtures with smaller

seeds. Seeds must be placed in seedboxes separate

from smaller seeded species so that planting rates can

be properly controlled. Seeds can be easily fractured

during cleaning or mechanical planting. Seedlings

usually establish and grow rapidly, and can persist

when seeded with most herbs. Plants normally require 3 to 4 years to attain maximum size, but once

established they remain highly productive.

Uses and Management

Cow parsnip is highly palatable to livestock and big

game (Ellison 1951; Hermann 1966; USDA Forest

Service 1937). It has considerable potential as an

ornamental plant for summer homes, administration

buildings, and recreational areas.

Figure 15—Cow parsnip in full bloom.

Reproduction is by seed only. Continual annual

grazing will prevent seed production, reduce vigor,

and result in elimination of this forb. Cow parsnip is a

highly preferred species that requires deferred grazing. Seeded areas should not be grazed for at least two

growing seasons following planting.

Varieties and Ecotypes

There are no releases.

Family Umbelliferae

Ligusticum porteri_______________

Porter Ligusticum

Porter ligusticum (fig. 16) a North American native,

is a member of the umbel or carrot family (Welsh and

others 1987). Plants are perennial with a characteristic aromatic taproot. White flowers are borne in a

terminal umbel which is often subtended by a whorl of

three to eight lateral umbels. Flower stalks are hollow

and may attain a height of 4 ft (1.2 m). Flowering

occurs in July and early August. Seed matures in

August and September.

Ecological Relationships and Distribution

Porter ligusticum is the most widespread ligusticum

in the Intermountain West. It can be found in openings

in aspen, spruce-fir, and subalpine types and as a

component of the understory in aspen stands. Porter

ligusticum prefers well-drained soils. It has fairly

good drought tolerance and good winter hardiness,

and can withstand considerable grazing (USDA Forest Service 1937).

Figure 16—Porter ligusticum growing in a

subalpine forb community.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 443

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture

If allowed to mature, good seed crops are generally

produced yearly. Seed is collected by hand stripping or

beating. Seed is flat, winged, and relatively easy to

separate from debris by screening. Dried seeds are

brittle and can be fractured or damaged by improper

cleaning or mechanical seeding. There are about

70,000 seeds per lb (154,000 per kg).

Seed can be stored for up to 5 years without

appreciable loss of viability (Stevens and Jorgensen

1994). Seed can be drilled or broadcast seeded. Fall

seeding onto a disturbed seedbed has worked well if

the seed is covered with soil. Seeds are heavy, irregularly shaped and do not flow well through most conventional seedboxes. However, when mixed with other

seeds of the same size, uniform plantings can be

achieved. Seeds are relatively large, and the gates of

most seedboxes must be opened wide enough to assure

passage. When this is done, seeds of smaller species

may be planted too heavily. Consequently, this species is often planted separately from other seeds, and

a carrier is added to aid in uniform planting.

Drill seeding at rates of 2 to 4 lb per acre (2.2 to

4.5 kg per ha) in separate rows from seeded grasses

is recommended. Broadcast seeding is possible if

followed by light harrowing or chaining. Although

studies have not identified specific ecotypic differences among populations, survival and growth rate

differences strongly indicate distinct ecotypes exist.

Seed lots should not be planted in different elevational

zones than the collection site. Moving seed from aspen

communities to sagebrush types is not advised.

Good germination and seedling establishment usually result from field plantings, but subsequent survival is often low. Plants are suited to grow with other

species and persist well in mixtures once established.

Some plantings have been conducted in the aspen and

oakbrush communities, but this forb also exhibits

usefulness in the upper sagebrush communities. It is

often encountered in openings and on shallow soils,

and can be seeded to improve forage conditions on

somewhat harsh situations.

Uses and Management

Livestock, particularly sheep, and big game make

considerable use of this species in spring and summer. It is highly sought out at all dates (Hermann

1966; Kufeld and others 1973; USDA Forest Service

1937). Seeded areas should not be grazed for at least 2

years following planting. Established stands should

be allowed to produce seed periodically, as reproduction is entirely from seed.

Varieties and Ecotypes

There are no releases.

Family Linaceae

Linum lewisii ssp. lewisii _________

Lewis Flax

There are approximately 100 Linum species in the

temperate regions of the world (Hitchcock and others

1961), with about 20 species occurring in the western

United States. Lewis flax is a delicate, erect perennial

sometimes reaching 3 ft (0.9 m) in height. Numerous

glabrous stems grow from a single crown with a woody

taproot. Flowers are generally sky blue (fig. 17), but

also range from white to dark blue. The petals are

caducous, dropping within 1 day of flowering. Flowers

open in the morning and close by mid-afternoon

(Howard and Jorgensen 1980). Flowering begins in

May and extends through June. Spherical capsules

that produce up to 10 seeds each mature in late July

and August.

Ecological Relationships and Distribution

Lewis flax ranges from Alaska south to California

and Texas (Harrington 1964). It is a perennial semievergreen that grows on well-drained soils ranging from

moderately basic to weakly acidic (Wasser 1982). Lewis

flax can be found in shadscale-saltbush, sagebrush

grass, and pinyon-juniper communities, and in openings in mountain brush, aspen, and conifer types,

especially on warmer south and west exposures. This

species is intolerant of poor drainage, flooding, and

high water tables (Wasser 1982).

Figure 17—Lewis flax in full bloom.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture

Lewis flax can be established easily from direct

seeding. Seeding can be accomplished by aerial and

hand broadcasting, drilling, and interseeding. Care

should be taken to ensure that seeds are not covered by

more than 0.13 inch (3 mm) of soil. Lewis flax has been

successfully seeded in range, wildlife, roadway, mine

disturbance, and landscape plantings. It establishes

well when included in mixtures (McKenzie and others

1980; Plummer and others 1970a), and grows and

reproduces well in association with other species

(Stevens and others 1981b). However, young plants

can be suppressed by competitive perennial grasses.

This species does well when seeded in mixtures with

shrubs. Generally, 0.25 to 0.5 lb per acre (0.3 to 0.6 kg

per ha) is seeded in mixtures. Reproduction is entirely

from seed. The species is well adapted to sagebrush,

pinyon-juniper, and mountain brush communities,

and is one of the few forbs that can be seeded successfully in the salt desert types (Monsen and Plummer

1978). Everett (1982) reported that Lewis flax was the

superior forb in a number of pinyon-juniper seedings.

It has been one of the most successful native forbs

seeded in the sagebrush and pinyon-juniper communities in Utah. Seeding is best accomplished in fall or

winter (Everett 1982; Howard and Jorgensen 1980).

Seeds require an afterripening period to obtain maximum germination (Eddleman 1977; Stevens and

Jorgensen 1994). Seed can be held in storage up to 10

years without any appreciable loss of viability (Stevens

and Jorgensen 1994).

Seed can be collected by hand, with power-driven

beaters, or with a combine. Seed production is generally high. Seed is also being successfully produced

under cultivation. Natural reproduction is usually

good, resulting in stable numbers and production

even though individual plants are relatively shortlived (4 to 7 years). Flowering occurs over a 6 week

period, causing seed to mature unevenly over an extended period of time. There are about 280,000 seeds

per lb (617,000 per kg) at 100 percent purity.

Uses and Management

Lewis flax produces basal foliage from a woody taproot (Plummer and others 1968). This species can be

classified as a semievergreen because some basal

foliage remains green throughout the winter. Animals

paw through the snow to reach the green growth in

winter. Early spring growth is grazed by big game,

upland game birds, and livestock. Forage value for

livestock and big game is moderate to high, and birds

and rodents seek out the seed (Everett and others

1978; Howard and Jorgensen 1980; Plummer and

others 1970a). Lewis flax responds well to late summer and fall storms by producing new basal leaves.

Because it maintains green basal foliage year round,

this species does not burn readily, and it can be used

as a fire suppressant species. Lewis flax has the ability

to compete with and spread in some cheatgrass communities. When seeded in mixtures, Lewis flax establishes and spreads well. It quickly occupies open areas

and slowly suppresses annual weeds.

Flower color ranges from nearly white to dark blue.

Profuse flowering occurs for about 6 weeks (Addicott

1977), beginning in mid-May. The flowering characteristics of this species make it a prime ornamental

candidate. When seeded in a mixture, Lewis flax can

help make a seeding more aesthetically pleasing.

Seeded areas should not be grazed for at least 2 years

on better sites and 3 years on drier, poorer sites. Seed

should be allowed to mature periodically. Individual

plants are relatively short-lived (4 to 7 years). Abundant seed is produced when plants are not excessively

grazed. Lewis flax will reproduce and remain in most

seedings if properly managed.

Varieties and Ecotypes

‘Appar’ Lewis flax was released by the Utah Division

of Wildlife Resources, the USDA Forest Service Intermountain Forest and Range Experiment Station, and

the USDA Soil Conservation Service Aberdeen Plant

Materials Center in 1980. This cultivar was named for

A. Perry Plummer. It was initially selected for its

appearance and competitiveness with native grasses.

It is easily grown under agricultural conditions and

has produced up to 700 lb of seed per acre (785 kg per

ha) with irrigation. ‘Appar’ exhibits a wide adaptability to sites in the Intermountain area. It is recommended for sites receiving 10 to 23 inches (25 to 58 cm)

of annual precipitation (Howard and Jorgensen 1980;

Shaw and Monsen 1983a). Pendleton and others (1993)

report that the origin of the cultivar appears to be

European and that it is derived from a naturalized

population of Linum perenne.

Maple Grove Lewis flax was released by the USDA

Forest Service, Rocky Mountain Research Station,

and the USDA NRCS Aberdeen Plant Materials

Center. It is a selected germplasm of Linum lewisii

that originated near Maple Grove, UT.

Family Umbelliferae

Lomatium triternatum ____________

Nineleaf Lomatium

Nineleaf lomatium (fig. 18) is a member of the

umbel or carrot family (Welsh and others 1987). It

was formerly identified as Lomatium simplex. It has

now been divided into two subspecies, L. triternatum

spp. platycarpum and L. triternatum spp. triternatum

(Hitchcock and others 1961; Welsh and others 1987).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 445

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

This aromatic perennial forb, with yellow flowers and

green leaves, grows to about 14 inches (36 cm) in

height. It blooms in April and May, and seeds mature

in July and August.

Ecological Relationships and Distribution

Nineleaf lomatium occurs in central and southwestern Colorado, Montana, southwestern Canada, and

south from California, to Colorado (Hermann 1966;

Hitchcock and others 1961; Welsh and others 1987).

Nineleaf lomatium prefers well-drained or dry

rocky soils on flats, sunny open ridges, and slopes.

L. triternatum ssp. triternatum occurs in mountain

brush, ponderosa pine and aspen forests, and dry

valleys from 5,200 to 8,500 ft (1,600 to 2,600 m). L. triternatum ssp. platycarpum is present in sagebrushgrass types, pinyon-juniper, mountain brush, ponderosa pine, lodgepole pine, and dry meadow communities

from 4,300 to 9,500 ft (1,300 to 2,900 m) (Welsh and

others 1987).

Plant Culture

Seed matures in July and August. It can be hand

collected. Screening to remove debris is all that is

generally required to clean field-harvested seed.

Viability of freshly harvested seed is generally good

and remains high for 3 to 4 years. There are about

42,000 seeds per lb (93,000 per kg) at 100 percent

purity. Seed can be drilled or broadcast seeded. Seeding is best accomplished in the fall, with seed being

covered not more than 0.25 inch (6.4 mm) deep. This

species does well when seeded in mixtures with other

herbs.

Figure 18—Lomatium in full bloom.

Uses and Management

Nineleaf lomatium is one of the first species to

begin growth following snowmelt. Extremely early

growth and development takes advantage of spring

soil moisture and provides much needed spring succulence for big game and livestock. Deer, elk, antelope,

cattle, and sheep make use of the foliage (Hermann

1966; Mueggler and Stewart 1980; USDA Forest Service 1937).

Large, deep taproots enable nineleaf lomatium to

withstand considerable drought, grazing, and trampling. Newly seeded areas should be given at least two

seasons of growth before they are grazed.

Varieties and Ecotypes

There are no releases.

Family Umbelliferae

Lomatium kingii_________________

Nuttall Lomatium

Nuttall lomatium, (fig. 19) formerly referred to as

Lomatium nuttallii (Welsh and others 1987), is a

native perennial forb that can be found growing from

sagebrush-grass up through the subalpine communities. It exists on basic and acidic soils (Plummer 1977;

Welsh and others 1987). Nuttall lomatium is not as

widely distributed as nineleaf lomatium. It does,

however, occur on drier sites and may have more

drought tolerance. Seed is best collected by hand.

Viability is generally high. Seeds are large with approximately 12,500 seeds per lb (28,000 per kg). More

Figure 19—Nuttall lomatium setting seed.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

than 70 percent germination can be expected even

after 5 years of storage (Stevens and Jorgensen 1994).

Seed is collected and cleaned in the same manner as

described for nineleaf lomatium.

This species is readily grazed by big game during

spring months. It is one of the first forbs to green up,

thus providing much sought after spring and early

summer succulence.

Family Fabaceae

Lupinus sericeus________________

Silky Lupine

Silky lupine (fig. 20) is a highly variable native

perennial legume. Plants are usually 12 to 47 inches

(30 to 120 cm) tall, growing from a branching caudex.

Flowers and foliage are usually quite abundant.

Flowers are large, blue, blue-purple, or white (Welsh

and others 1987). Flowers emit a distinctive pungent

odor. The leaves are large, palmately compound, often

pubescent, and gray-green (Welsh and others 1987).

Individual seedpods usually contain 3 to 5 seeds.

Ecological Relationships and Distribution

Silky lupine ranges from southern Canada south to

New Mexico (Hermann 1966). It is common in the

Great Basin and can be found in sagebrush-grass,

pinyon-juniper, mountain brush, ponderosa pine, and

aspen types. Welsh and others (1987) reported that

silky lupine is one of the most widely distributed

species in Utah, with four different varieties reported.

It is most abundant in upper mountain brush and

aspen communities. This species grows on basic,

neutral, and slightly acid soils. It prefers open sunlight, but possesses some shade tolerance.

There are a number of perennial lupines in the

Intermountain West. Most tend to intergrade with

each other. Two common and abundant species are

silky lupine and silvery lupine.

Plant Culture

Seed is usually ready to harvest in early September.

Seed pods must be collected before they dry and seeds

are shattered. Consequently, seed is collected by hand

stripping of the complete inflorescence and, if in pure

stands, by mechanical seed beaters or combines. Following collection, the pods need to be further dried

before seeds can be extracted. During drying, collected

material must be covered with a screen to prevent seed

loss. As pods dry, they open, twist, and propel the

seed a distance of 3 to 10 ft (0.9 to 3.0 m). Lupine

seeds have considerable longevity. Silky lupine seeds

have been stored for 20 years and still retained 72

percent germination (Stevens and Jorgensen 1994).

Figure 20—Silky lupine.

Seeds are rather large with only 13,000 per lb (29,000

per kg). Most seeds imbibe water when planted but

some fail to germinate. Some seeds that fail to germinate are lighter in color than healthy seeds. Silky

lupine should be seeded in the fall to meet stratification requirements. Seed can be drilled or broadcast,

with seed covered not more than 0.25 inch (6.4 mm)

deep.

Silky lupine establishes well when seeded on disturbed or depleted sites in acid or basic soils (Plummer

1977). Fairly strong seedlings are produced; however,

full plant development can be somewhat slow, with

flowers not being produced for 3 to 5 years following

seeding.

Good numbers of seedlings appear from most

plantings. However, plant numbers are generally reduced quite dramatically the first growing season.

Plants that persist the first year are usually very

hardy and will persist even under adverse conditions.

Young plants are quite competitive and persist with

some competition. To date, most range seedings have

been conducted using seed collected from aspen communities. These sources are not suited to lower, drier

elevations.

Uses and Management

Silky lupine is one native legume that has good

potential for revegetation of ranges in upper sagebrush-grass, mountain brush, ponderosa pine, and

aspen types. This species provides early spring succulence and is grazed by cattle, sheep, deer, rodents,

and small mammals throughout the year (Hermann

1966; Holechek and others 1982; Kufeld and others 1973;

Plummer and others 1968). It can be seeded in mixtures or in separate drill rows from grasses and other

broadleaf herbs. It is extremely productive and provides not only useful forage but also soil protection
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 447

Chapter 19 Forbs for Seeding Range and Wildlife Habitats

and erosion control. It persists with grazing, and does

well as an understory species. It can be used in riparian disturbances and on dry meadows that are not

flooded.

Silky lupine often recovers well when disturbed

sites are protected for a number of years. Rodents

gather and forage on lupine seeds and can interfere

with seedings and natural spread. Seeded areas should

not be grazed for at least 2 years, and in some cases,

3 years following planting. Once established, this

species has excellent grazing and drought tolerance.

Varieties and Ecotypes

There are no releases.

Family Fabaceae

Lupinus argenteus ______________

Silvery Lupine

A major difference between silvery lupine and silky

lupine is the presence of less pubescence on the leaves

and stems of silvery lupine. Its flowers are predominantly blue-purple, blue, and white. Meeuwig (1960)

reported that silvery lupine has outstanding forage,

ground cover, and revegetation potential at higher

elevations. Most of the perennial lupines readily hybridize (Hermann 1966). Lupinus alpestris is now

included as part of the silvery lupine complex (Welsh

and others 1987). Seed production, seed handling,

seeding, and seedling establishment characteristics of

silvery lupine are similar to those of silky lupine.

Silvery lupine occurs at higher elevations than does

silky lupine. Major areas of occurrence are aspen,

spruce-fir, and subalpine herblands (fig. 21).

Figure 21—Silvery lupine in a subalpine community.

Poisonous Characteristics

Some lupines, including silky and silvery, have

poisonous characteristics and can cause death if used

improperly during some seasons of the year (Davis

1982b; Davis and Stout 1986; James and others 1980;

USDA Soil Conservation Service 1968). Care should

be taken with livestock where substantial quantities

of lupine are available. Livestock that obtain forage of

other species in addition to lupine are generally not

adversely affected (USDA Soil Conservation Service

1972b). When planting lupine, care should be taken to

ensure that it is seeded in mixtures with other species, and that large, solid stands are not established.

Family Fabaceae

Medicago sativa

Medicago falcata ________________

Alfalfa

Sicklepod Alfalfa

Two species, Medicago sativa(alfalfa) and M. falcata

(sicklepod alfalfa), were introduced to North America

(Hansen 1913). Alfalfa is generally blue-flowered (fig.

22) and nonrhizomatous, whereas sicklepod alfalfa is

generally yellow-flowered and somewhat rhizomatous.

Both species are perennials. Stems normally reach

11 to 40 inches (30 to 100 cm), depending on growing

conditions. Stems are branched; some are ascending.

Most range varieties form decumbent stems at the

base of the plant. Varieties cultivated for hay production or irrigated pastures are normally more erect.

Figure 22—‘Nomad’ alfalfa. Yellow- and blue-flowered

alfalfa growing together on a pinyon-juniper site.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Leaves of alfalfa are glabrous to hairy, alternate,

and pinnately trifoliolate; the leaflets are serrate

distally and oblanceolate to oblong. Flowers are small,

usually less than 7 to 10 mm long. Fruits consist of

several-seeded spiral pods (Harrington 1964). Plants

produce an abundance of herbage, with growth beginning early in the spring.

Alfalfa is a complex taxon with nine recognized

subspecies (Gunn and others 1978). These express

intergrading morphologies and hybridize quite freely.

The genus evolved in the Mediterranean region. The

perennial mesophytic subspecies arose in Western

and Central Asia (Lesins and Lesins 1979). Diploid

(2N = 16) and tetraploid (2N = 32) forms commonly

occur. Most cultivars are tetraploid (Rumbaugh and

Townsend 1985). Lesins and Lesins (1979) report that

the diploid populations of alfalfa originate from steep

mountainous regions.

Miller and Melton (1983) described over 400 cultivars and strains of alfalfa in North America. Today

there are many more. The origin of the 15 most

commonly grazed populations are described by

Rumbaugh (1982a). All dryland forage types are interspecific hybrids of alfalfa and sicklepod alfalfa

(Rumbaugh and Townsend 1985) (fig. 23). These

authors reported that sicklepod alfalfa contributes

genes that increase drought tolerance, winter hardiness, and tolerance to grazing. Alfalfa germplasm

enhances resistance to disease and insects and improves forage and seed production.

Most range and field varieties of alfalfa used in

Western range plantings arose from the early efforts

of Dr. N. E. Hansen (1913) of South Dakota. Early

testing conducted by Dr. Hansen, led to identification

of population with spreading root systems. This characteristic has subsequently been bred into field and

range varieties (Rumbaugh and Pedersen 1979).

Canadian scientists later advanced the usefulness of

this species by breeding varieties with better grazing and cold tolerance (Heinricks 1963). Alfalfas

were entered into extensive field plantings within

the Intermountain region in the 1930’s. Numerous

field and range varieties have subsequently been

developed.

Ecological Relationships and Distribution

Rangeland varieties of alfalfa are well adapted to

areas that receive 10 inches (25 cm) or more of annual

precipitation. They are especially well-adapted to sagebrush/grass, pinyon-juniper, and mountain brush

(fig. 24) communities. They have been seeded and

have performed fairly well in aspen, spruce-fir, and

subalpine types. This forb is semievergreen, having

green basal leaves throughout the winter. Growth

starts in the spring when the plant is exposed from

underneath the snow. Once grazed, regrowth of rangeland types is limited compared to that of irrigated

alfalfas.

Figure 23—‘Rambler’ alfalfa seeded into rootplow

strips in a Gambel oak-mountain big sagebrush

community.

Seedling establishment is rapid, and seeding success rate can be high. Grazing stimulates rhizome

production, which results in increased crown size and

ground cover (Rosenstock and Stevens 1989).

Alfalfa is well suited to harsh sites and infertile

soils. Under these conditions seedlings of few other

herbs survive as well as alfalfa. Once established,

alfalfa serves as an excellent nurse crop. Although

plants begin growth early in the spring, they do not

prevent the entry of other species. Alfalfa may produce

considerable annual litter that provides a good cover

for the seedbed. Once established, alfalfa is very persistent (Rumbaugh 1982b) (fig. 25), but little reproduction has occurred from established stands on

rangeland sites (Rosenstock and Stevens 1989). Rhizomatous forms do spread vegetatively even under

arid conditions.

Figure 24—‘Ladak’ alfalfa on a 27-year-old

pinyon-juniper chained site.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture

Alfalfa is the most widely seeded rangeland forb in

the Intermountain West (Rumbaugh 1984; Rumbaugh

and Townsend 1985). Depending on the rangeland

sites and seeding technique, it can be seeded in the fall

or spring. Planting early in the spring usually produces good stands, but plantings in late fall or winter

have been satisfactory in large restoration programs.

Late winter plantings may prevent precocious germination (Plummer and others 1968), but alfalfa

germinates quickly in early spring and may succumb

to spring frosts.

Seed can be drilled or broadcast seeded, but it must

be covered. Aerial seeding followed by anchor chaining

or pipe harrowing and aerial seeding on snow over

disturbed soil has produced excellent rangeland stands.

Broadcast planting has often been more successful

than drilling (Plummer and others 1968). Fall aerial

seeding on a rough seedbed has resulted in excellent

stands (fig. 25). Whether broadcast or drilled, seed

should be covered 0.25 to 0.5 inch (6 to 13 mm) deep

(Plummer and others 1968; USDA Soil Conservation

Service 1971a).

Alfalfa is particularly adapted to drill or broadcast

seeding in mixtures with other species. Seeds are

rather small, round, and smooth. When mixed with

grasses, seed sorting and separation can occur in the

seedbox. Alfalfa seed usually does not separate when

mixed with grasses if planted at a ratio of 1 to 5.

Separate legume seedboxes are mounted on most

drills. The legume boxes are designed to dispense the

small alfalfa seed more accurately than the larger

seedboxes and eliminate seed separation when mixtures are planted.

Although alfalfa seeds should be planted slightly

shallower than large seeded grasses, adequate stands

normally appear when they are drilled together. Alfalfa seedings develop rapidly and compete favorably

with many grasses and other broadleaf herbs. Alfalfa

seedlings, however, do not compete well with seedlings of crested and intermediate wheatgrass and

smooth brome.

Because of the possibility of bloat, alfalfa should be

seeded as a component of a mixture when planted in a

pasture (Lorenz 1982). Up to 40 percent by weight of

a grass-legume mixture has been seeded to alfalfa

without causing bloat problems to livestock. Bloat has

not been observed as a problem on mixed rangeland

seedings.

Because alfalfa is highly utilized by livestock, big

and small game, birds, rodents, and rabbits, a sufficient number of plants must be established to ensure

the species is not eliminated by overuse. On most

ranges, at least 2 lb of seed per acre (2.2 kg per ha)

should be planted in the seed mix. Seed costs are

generally not prohibitive, and under some conditions

alfalfa should be planted at even higher rates. As

seeding rates are increased, stand density also increases. Seeding alfalfa at rates exceeding 6 to 8 lb per

acre (6.7 to 9.0 kg per ha) is unnecessary. There are

about 220,000 seeds per lb (485,000 per kg). Purity is

generally over 95 percent with germination at least

85 percent. Under warehouse storage seed longevity is

excellent and little viability is lost after 25 years

(Stevens and Jorgensen 1994).

Alfalfa is able to fix nitrogen, thereby improving

soil fertility. Johnson and Rumbaugh (1981) reported that alfalfa appears able to fix nitrogen even

during periods of drought when other legumes do not

nodulate.

Uses and Management

Alfalfa is heavily grazed by livestock, big and small

game, upland game birds, rodents, and rabbits during

all seasons (fig. 26). The inclusion of alfalfa in a

rangeland seeding can have a number of positive

effects. It can (1) increase total herbage production, (2)

increase production and protein content of associated

species (Johnson and others 1983; Rumbaugh and

others 1981, 1982), (3) extend the grazing season, both

during spring and fall, (4) increase diversity (flora and

fauna) of seeded communities, and (5) improve soil

stability. Species growing in association with alfalfa

are generally positively affected by its ability to fix

nitrogen (Johnson and others 1983; Rumbaugh and

others 1981, 1982). Consequently, alfalfa can be

interseeded into established grass stands to improve

vigor and herbage production.

Figure 25—Rehabilitated range 21 years following chaining of juniper-pinyon and aerial

seeding. Thirteen grasses, shrubs, and forbs

were seeded. After 21 years alfalfa is the

predominant forb.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Alfafa is compatible with native species in seedings

in mountain brush, pinyon-juniper and big sagebrush communities. The forb has not noticeably diminished or eliminated native broadleaf herbs or

grasses.

Regrowth capacity varies among varieties. Deeprooted types possess better regrowth capabilities than

rhizomatous range types. Alfalfa initiates growth early

in the spring and remains green and productive as

long as soil moisture permits. It can recover following

grazing or clipping. It normally attains maximum

production after most cool-season grasses cease

growth. Certain strains of grass, including ‘Latar’

orchardgrass and ‘Greenar’ intermediate wheatgrass,

mature when alfalfa is at the optimum stage of growth

for harvesting as hay (Hafenrichter and others 1968).

These species can be planted in mixtures to significantly extend the season of grazing and enhance total

herbage production. Alfalfa also furnishes excellent

fall green-up, and standing crops cure quite well.

Consequently, this species has been used in wildlife

seedings to attract and keep game animals from trespassing in agricultural fields. Areas seeded to alfalfa

should not be grazed for a minimum of two growing

seasons following seeding. Early use can result in poor

survival and loss of vigor (Berdahl and others 1986).

Drier areas may require additional time before stands

fully establish.

It is not unusual for seedlings and young plants to

succumb during the first 1 to 3 years following planting (Rosenstock and Stevens 1989). Once established,

however, alfalfa has considerable longevity, resistance to grazing and trampling, and moderate shade

tolerance. Kilcher and Heinrichs (1966b), Pearse (1965),

Rosenstock and Stevens (1989), and Rumbaugh and

Figure 26—Alfalfa seeding in a chained pinyon-juniper basin big sagebrush area. Deer

use only on left, rabbit and deer use on right.

Peterson (1979) reported excellent stands persist

23 to 28 years after seeding on sites with 8 to 11.5 inches

(20 to 29 cm) of precipitation. The Utah Division of

Wildlife Resources has aerial seeded alfalfa on over

150,000 acres (60,700 ha) of game range during the

past 30 years. At a few sites, there has been a major

loss in plant density. Where a decrease in density has

occurred, it has been a result of continuous heavy

rabbit use and continual spring use by cattle. Similar

results have been reported by Rumbaugh and Pedersen

(1979) and USDA Soil Conservation Service (1971a).

Rumbaugh (1982b) reported that stand density of

eight different varieties was adequately maintained

during a 25-year period, with proper spring grazing by

cattle. Where insufficient seed is planted and plant

density is low, considerable grazing pressure will

reduce the stand At least 2 lb. (0.9 kg) of seed per acre

should be seeded to ensure sufficient density to sustain grazing.

Alfalfa generally receives more grazing pressure than

other associated species and withstands greater use

than other broadleaf herbs. Weakened stands recover

readily when grazing pressures are removed. The deep

taproot, large crowns, and extensive rhizomes enable

the plant to persist over extended periods of drought.

New seedings can be damaged by continuous grazing

by rabbits, rodents, livestock, or big game animals.

Insects, including grasshoppers, have seriously defoliated alfalfa, often within a 2 to 5 day period. Attempting to seed alfalfa and palatable grasses in areas with

high rabbit populations can be risky. Once established, mature plants can usually recover from cyclic

periods of heavy use, but young seedlings cannot.

Alfalfa can be used to sustain big game animals

during the early spring and summer months. Plantings established on pinyon-juniper sites in central

Utah clearly attract deer and provide a substantial

part of their diet, beginning in early March and continuing until early June (Rosenstock and others 1989).

Wildland sites seeded to support big game should be

grazed only lightly by livestock. If plants are subjected

to stress by drought or excessive use by insects or

rabbits, livestock grazing should be curtailed.

Alfalfa has been seeded on some rather arid sites,

including sites dominated by Wyoming big sagebrush

and shadscale saltbush. When seeded during years of

average precipitation, plants usually establish. However, failures can be expected if seeding occurs during

years with low precipitation. Plantings can be maintained under arid circumstances if grazed judiciously.

Varieties and Ecotypes

A number of strains of alfalfa—‘Ladak’, ‘Rambler’,

and ‘Nomad’—are adapted to ranges where annual

precipitation exceeds 10 inches (25 cm) (Plummer and
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

others 1968; Rumbaugh 1982b). These strains produce much larger crowns than the usual field strains

and have more complex root systems. In addition, they

survive better from underground attacks by gophers

and other rodents. Some plants also spread by stem

layering. Consequently, these strains are superior for

seeding rangelands (Plummer and others 1968).

‘Ladak’ is the best known and most widely used

rangeland variety. If seeds of several strains are

available, a mixture may be used. The varieties mentioned above have shown unusual persistence and

production under heavy grazing and lower precipitation (Plummer and others 1968). Other cultivars—‘Spreader’, ‘Spreader II’, ‘Rhizoma’, ‘Runner’,

‘Travois’, ‘Teton’, and ‘Drylander’—have also demonstrated good adaptability to wildland sites.

Additional varieties are currently being developed

for rangelands. A major concern is to improve drought

tolerance so the forb may be planted in more arid

environments.

Family Fabaceae

Melilotus officinalis ______________

Yellow Sweetclover

The two most abundant sweetclovers on Western

ranges are yellow sweetclover (yellow-flowered), and

white sweetclover (white-flowered). Neither are native to North America. There are annual and biennial

forms of each; however, most populations are biennials (Rumbaugh 1984; Smith and Gorz 1965). During

the first season, only vegetative growth occurs; during

the second season, long, well-branched stems 1 to 6 ft

(0.3 to 1.8 m) tall (fig. 27) develop. Leaves are trifoliolate, and dentate margined, with small narrow

stipules. Flowering occurs on long narrow racemes,

and single seeds are produced in straight smooth

pods (Hermann 1966). After flowering and seed

maturation, plants usually die.

Ecological Relationships and Distribution

Yellow sweetclover originated in Europe and Asia

and is now distributed over most of the United States

and Southern Canada. The species is especially adapted

to disturbed sites. It has a strong seedling, fairly good

drought resistance, and good winter hardiness (USDA

Forest Service 1937; USDA Soil Conservation Service

1971b). It grows well with over 10 inches (25 cm)

annual precipitation in aspen, mountain brush,

ponderosa pine, big and black sagebrush, and pinyon-juniper types. Yellow sweetclover also does

well in saline and alkaline soils and on many riparian

sites (Plummer 1977; USDA Soil Conservation Service 1968, 1971b; Wasser 1982).

Figure 27—Yellow sweetclover.

Plant Culture

Seed production of yellow sweetclover is generally high. There are more than 250,000 seeds per lb

(550,000 per kg). Germination is generally high, and

good viability is retained for at least 2 years with

warehouse storage. Adequate, inexpensive seed is

generally available. Like most legumes, sweetclover

seeds have a hard coat and are best seeded in the fall

or winter. Seeds can lie dormant in the soil for years

and then germinate when temperatures and moisture

conditions are favorable. In central Utah, seedings

more than 35 years old still retain some yellow

sweetclover. Nichols and Johnson (1969) report that

with 15 inches (38 cm) of precipitation, yellow

sweetclover will reseed successfully. Seeds can be

drilled or broadcast on relatively unprepared seedbeds. Best stand establishment occurs when seeds are

planted 0.13 to 0.25 inch (3 to 6 mm) deep, preferably

on a fairly firm seedbed.

Uses and Management

Yellow sweetclover is frequently seeded on Western

rangelands (Rumbaugh and Townsend 1985). It is an

effective erosion control species because it has an

extensive, deep taproot system and a strong vigorous
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

seedling. It also produces a dense ground cover in 1 or

2 months and fixes nitrogen. Up to maturity, palatability is high for big and small game (Graham 1942;

Kufeld 1973; Kufeld and others 1973; Mueggler and

Stewart 1980; Plummer and others 1968; Rumbaugh

1984; USDA 1971b) and for livestock (Hermann 1966;

USDA Forest Service 1937; USDA Soil Conservation

Service 1971b), with forage quality reportedly higher

than that of alfalfa (Magness and others 1971;

Rumbaugh 1984). Waterfowl, upland game birds, and

bees also use yellow sweetclover (Autenrieth and others 1982; Rumbaugh 1984; USDA Soil Conservation

Service 1971b). The species has a very strong seedling

that emerges in the spring, providing early succulence. Yellow sweetclover provides much needed live

ground cover and forage during the critical first and

second years. It can be seeded at 0.5 to 2 lb per acre

(0.56 to 2.2 kg per ha), (Nichols and Johnson 1969;

Plummer and others 1968; USDA Soil Conservation

Service 1971b). Yellow sweetclover is generally seeded

with perennials on disturbed sites. It furnishes initial

cover and forage as slower developing herbs attain

mature stature. It withstands heavy use and can

reseed following fires or other disturbances.

Varieties and Ecotypes

There are three released varieties—‘Goldtop’,

‘Madrid’, and ‘Yukon’ (Wasser 1982). These varieties

have not performed any better than the common seed

sources. Most seed marketed is from uncertified

common lots.

Family Boraginaceae

Mertensia arizonica ______________

Tall Bluebell

Tall bluebell is the most widely distributed Mertensia

in the Intermountain West (Higgins 1972; Mathews

1968). There are, however, other bluebells that are

locally important (Hermann 1966). Different varieties

of tall bluebell have been described, particularly from

collections obtained in Utah (Cronquist and others

1984; Welsh and others 1987). Tall bluebell is a rhizomatous native perennial with a thick, woody root. It

grows to a height of 2 ft (61 cm) and produces an

abundance of pale blue to very dark blue flowers

(fig. 28). Leaves are abundant and are borne on erect

stems. Fruits are hard and consist of four nutlets

(Cronquist and others 1984).

Ecological Relationships and Distribution

Tall bluebell occurs in fairly moist to well-drained

soils in aspen, spruce-fir, ponderosa pine, and subalpine types, usually at elevations from 6,000 to

11,000 ft (1,800 to 3,400 m). The species can be found

along streams in canyon bottoms, meadows, parks,

and in scattered timber stands. It grows well in shade

and open situations in pure stands as well as in mixed

communities. Tall bluebell is present in all mountain

regions of Utah, southwestern Wyoming, and northwestern Colorado (Cronquist and others 1984).

Plant Culture

Flowering occurs during midsummer. Seeds mature

in September and can be collected by hand stripping,

beating, and in pure stands with various types of

combines. Germination is generally high (60 to 70

percent), with good viability being maintained for at

least 5 years. Seeding is best accomplished in the fall

on disturbed soil, with seed covered 0.25 to 0.5 inch

(6 to 13 mm) deep. Tall bluebell can be seeded in

mixtures with other herbs. It usually grows on fertile

soils. When planted on disturbed, degraded sites it has

not done well. Under good conditions, plants establish

quickly and reach maturity in 3 to 5 years. Plants

spread slowly by rhizomes, yet are very persistent and

compatible with other native species when seeded in

mixtures.

Uses and Management

Sheep, elk (Kufeld 1973), and deer (Kufeld and

others 1973) seek tall bluebell and gain weight exceptionally well on the herbage. Cattle prefer the flower

stalks but will make use of all plant parts (USDA

Forest Service 1937). Sheep graze the herb whenever

available. The bluebells are generally very showy and

fairly easily seeded or transplanted, and are good soil

stabilizers. Consequently, the plant is a good candidate for use in recreational areas, summer home sites,

and areas with high aesthetic value and use. The

plant furnishes such excellent forage that it should

Figure 28—Tall bluebell in full bloom.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

be included in most seedings on adapted sites. However, the lack of sufficient seed currently curtails the

use of this forb. This species has not done well if

planted off site, particularly at lower elevations where

it normally does not occur.

Newly seeded areas should not be grazed for at least

2 years. Large fibrous root systems, once established,

allow tall bluebell to withstand periods of drought,

grazing, and trampling. Concentrated use can occur if

seedings are not properly managed.

Varieties and Ecotypes

There are no releases.

Family Fabaceae

Onobrychis viciaefolia

Sainfoin

Sainfoin (fig. 29) is a perennial, nonbloating, introduced legume that grows 1 to 3 ft (30 to 91 cm) tall, and

produces white or pink to purplish pea-like flowers.

Stems are semierect, originating from a branched root

crown attached to a deep main taproot. Leaves are

alternate, odd-pinnate and oblong to elliptic or oblanceolate (Rumbaugh and Townsend 1985; Welsh and

others 1987).

Ecological Relationships and Distribution

Sainfoin was introduced into North America from

Europe. It has been used primarily as a hay or pasture

crop. However, the species demonstrates considerable

usefulness for rangelands.

Sainfoin is adapted to sagebrush, pinyon-juniper,

and mountain brush areas that receive at least 14

Figure 29—Sainfoin growing with crested wheatgrass and basin big sagebrush.

inches (36 cm) of annual precipitation. Sainfoin prefers well-drained, deep calcareous soils (Eslick 1968),

exhibits fairly good salt tolerance (Jensen and Sharp

1968), and grows well in soils low in phosphorus

(Rumbaugh and Townsend 1985).

Plant Culture

Single seeds are produced in pods and are marketed

unshelled. There are about 26,000 unshelled seeds

per lb (57,000 per kg). Unshelled seeds are best drilled,

but can be broadcast if covered sufficiently. Seed

should be covered at least 0.25 inch (6 mm) and not

more than 0.75 inch (19 mm) deep (Jensen and Sharp

1968). Seeding is best done on a firm seedbed that is

relatively weed free. For range seeding, it is best to

seed in the fall. Seed should be inoculated with the

proper rhizobium (Dubbs 1968). Seed is marketed

with about 70 percent germination and 95 percent

purity. When seeded in a mixture, about 2 to 5 lb per

acre (2.2 to 5.6 kg per ha) is recommended. Seed can

be stored up to 4 years without much loss of viability.

This species can be seeded individually or as a

component of a mixture. A strong seedling is usually

produced. Rate of growth following seeding is only fair

on mesic sites. However, plants are quite persistent

and hardy. Two years are required for flowering to

occur under favorable conditions.

Uses and Management

Sainfoin is nonbloating and palatable to livestock,

big game, and sage-grouse (Autenrieth and others

1982; Holder 1968; Rumbaugh 1984; Shaw and Cooper 1973; USDA Soil Conservation Service 1968a).

Mule deer in Utah have been observed to prefer sainfoin hay over alfalfa hay. Sainfoin has a deep taproot

and has wilt and drought hardiness equal to or slightly

less than does ‘Ladak’ alfalfa (Dubbs 1968; USDA Soil

Conservation Service 1968a). Some basal leaves remain green throughout most of the winter. Sainfoin

starts growth early in the spring, usually prior to

alfalfa. It continues to grow or remains green

throughout the summer period. Once grazed or clipped,

regrowth is limited. Characteristics that give sainfoin

considerable potential in the Intermountain area as a

forage crop include early greenup, cold and drought

tolerance, nonbloating characteristics, high forage production potential, ability to grow with other species,

and adaptation to calcareous soil.

Areas seeded to sainfoin should not be grazed for at

least the first two growing seasons. Every 2 to 3 years,

seed should be allowed to mature and seed naturally.

Plants are persistent and withstand heavy clipping or

grazing. Once established, this species also competes

fairly well in mixed communities.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Varieties and Ecotypes

A number of improved varieties have been released,

including ‘Eski’, ‘Melrose’, ‘Nova’, ‘Onar’, ‘Remont’,

and ‘Runemex’. All were released for cultivated pastures and for hay. ‘Eski’ is the most commonly seeded

variety on ranges. This variety does well when seeded

in upper pinyon-juniper, mountain big sagebrush, and

mountain brush areas.

Family Umbelliferae

Osmorhiza occidentalis __________

Sweetanise

Sweetanise is an erect, perennial, aromatic native

that grows up to 4 ft (1.2 m) tall (Welsh and others

1987) (fig. 30). Crushed leaves and seed give off a

heavy licorice odor (Hermann 1966). It is a member of

the umbel or carrot family and produces small umbels

of yellowish-green flowers. Flowering occurs in May

and June, and seed matures in September. Fruits are

slender and sharp-pointed.

Ecological Relationships and Distribution

Sweetanise occurs from Western Canada south

through Utah and Colorado. It grows well in shade,

open areas, mixed communities, and in solid stands.

Sweetanise prefers cool, moist woods, moist hillsides,

valleys, and forest openings in aspen, spruce-fir, and

subalpine types. It grows on basic, neutral, and slightly

acidic soils.

Plant Culture

This species can be successfully established by direct seeding. Lack of a consistent seed source reduces

seeding of this species. It establishes easily and spreads

quickly from seed. Seeding is best completed in the

fall. Seed should be covered, but not more than 0.25

inch (6 cm). Sweetanise does well seeded in mixtures.

Seed is usually collected by hand, but dense stands can

be mechanically harvested. Seeds are brittle and can

be damaged during cleaning and seeding. Seeds can

generally be cleaned by screening from harvested

debris. There are about 30,000 seeds per lb (73,000 per

kg). Seed should not be stored more than 2 years, or

considerable loss in viability can occur. Thirteen year

old seed is reported to exhibit 44 percent germination

(Stevens and Jorgensen 1994). Seeds are large and

usually must be drilled seeded to ensure proper soil

coverage. Seeds are approximately the same length as

some grass seed and can be planted in a mixture,

depending on the species sown.

Uses and Management

Cattle, sheep, and big game show a particular fondness for the foliage and for developing seeds of

sweetanise (Hermann 1966; Kufeld 1973; Kufeld and

others 1973; Plummer and others 1955; USDA Forest

Service 1937). The plant is palatable and remains

green throughout the grazing season. Consistent seed

crops are produced yearly if seeds are allowed to

mature before being grazed. Large, thick, woody roots

allow the plant to withstand considerable use during

dry periods. This species does not furnish a dense

ground cover and is not recommended for erosion

control. Heavy grazing can eliminate or reduce its

density.

Seeded areas should not be grazed for at least 2

years following planting. A fairly strong seedling is

produced, and the rate of growth is good. Seeds are

generally produced the second year following planting. Maturing seeds are readily eaten by grazing

animals. Some type of deferred grazing should occur to

allow seeds to mature at least every 2 to 3 years.

Varieties and Ecotypes

There are no releases.

Family Scrophulariaceae

Penstemon palmeri ______________

Palmer Penstemon

In North America there are over 200 native species of Penstemon (Hermann 1966; USDA Forest Service 1937). Many are short-lived herbaceous species

Figure 30—Sweetanise with mature seed.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

that spread readily by seed. A number are rhizomatous. Some penstemon are semievergreen, producing

basal leaves that persist the entire year. They can be

found in most vegetative types of Western North

America.

Palmer penstemon (fig. 31) has long flowering stalks

20 to 55 inches (50 to 139 cm) tall with large showy

pink blossoms that are produced in late spring and

early summer. Seed matures in September. Flowers

give off a pleasant fragrance that is unique among the

penstemons, as they usually have little or no scent

(Stevens and others 1985b).

Ecological Relationships and Distribution

Palmer penstemon is an evergreen native perennial

that occurs in blackbrush, sagebrush-grass, pinyonjuniper, mountain brush, and ponderosa pine types

of the Intermountain West. It is a fairly short-lived

(4 to 5 years) pioneering species. Plants spread by

seed. This species can be found growing on basic and

acidic soils (Plummer 1977). It produces long seed

stalks and an abundance of basal evergreen leaves

that usually remain gray-green throughout the year.

A large fibrous root system is produced. Palmer penstemon has considerable grazing tolerance and good

drought tolerance.

Plant Culture

Good results can be expected from drilled or broadcast seedings on disturbed soils and on raw and

eroding sites (fig. 32) where it occurs naturally as an

early invader. Seed can be drilled or broadcast onto

disturbed soil, with covering not to exceed 0.15 inch

(3 mm). Natural reproduction generally compensates

for the short-life span of individual plants. Seed is

generally produced in abundance. Some dormancy

exists that allows seeds to accumulate in the soil

until favorable conditions occur for germination and

establishment.

Seeds of most penstemons, particularly Palmer penstemon, can be seeded in mixtures with most other

herbs. Small vigorous seedlings appear early in the

spring. They compete well and usually are not eliminated by competition from other species. When seeded

heavily, considerable thinning or dieoff of young

seedlings normally results. However, sufficient survival usually occurs to provide adequate stands. Palmer

penstemon is particularly adapted to harsh situations and tends to dominate or establish on areas

where other species fail. It is one of the few native

broadleaf herbs that most consistently establishes

from seeding. It is particularly important for its ability

to establish and enhance the establishment of other

plants on harsh sites.

All species of penstemon have small seeds that are

easily cleaned to a purity that exceeds 90 percent.

Palmer penstemon has 350,000 seeds per lb

(770,000 per kg) at 100 percent purity. Seeds are so

small that they are difficult to meter through most

conventional drills. If seeded alone, a carrier is often

used to regulate planting rates. Seeds can be mixed

with seeds of most other species although some separation can occur in the seedbox. When seeded in

mixtures, 1 to 3 lb per acre (1.1 to 3.4 kg per ha) are

Figure 31—Palmer penstemon in full bloom. adequate.

Figure 32—Broadcast seeding of Palmer penstemon on a roadside disturbance.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Seeds can be collected by hand stripping or beating;

pure stands can be combined. Seeds are cleaned by

hammermilling and screening. Storage of up to 5 years

has resulted in no significant loss in germination

(Stevens and Jorgensen 1994). Seeds are being produced commercially.

Uses and Management

Palmer penstemon is eaten readily by deer and elk,

especially during winter and spring months (Kufeld

and others 1973; Plummer and others 1968). Basal

leaves are green and succulent all year long. Cattle

and sheep also make use of this species. The large,

fibrous root system and large, succulent basal leaves

provide considerable soil protection. This species is an

early invader and does especially well on disturbed raw

soils. Pink blossoms in late spring and early summer

make this plant useful for beautification projects.

Because of seed dormancy, seeding should occur in

the fall. Once established, stands are self-perpetuating. Seeded areas should not be grazed for at least two

growing seasons following planting. Flowering and

seed production often occur the second year following

planting. This species has little shade tolerance, but

grows as an understory with sagebrush. It is somewhat resistant to fire and trampling, but can be eliminated by excessive grazing.

Varieties and Ecotypes

‘Cedar’ was selected and released in 1985 (Stevens

and others 1985b) for its wide area of adaptation,

winter succulence, forage and seed production, and

grazing preference by livestock and wildlife. ‘Cedar’

does well in blackbrush, salt desert shrub, sagebrushgrass, pinyon-juniper, and mountain brush types

within the Intermountain region. Commercial seed is

readily available and recommended for use.

Family Scrophulariaceae

Penstemon cyananthus __________

Wasatch Penstemon

Wasatch penstemon (fig. 33) is a native perennial

with bright semievergreen basal leaves. Plants develop from a primary taproot and produce several 2 to

3 ft (61.0 to 91.4 cm) long stems with deep blue to

nearly purple flowers. Wasatch penstemon can be

found on rocky soils, but is usually associated with

more fertile, deeper soils that may be slightly acidic or

basic. It occurs from the mountain brush type to

higher elevations, including the subalpine zone.

Flowering occurs from May through July. Seeds

are formed in September. Seed processing and

planting features are similar to those of Palmer

penstemon and most other penstemons. Seeds are,

however, somewhat larger with about 230,000 per lb

(510,000 per kg) at 100 percent purity. This species is

particularly important as a forage and cover plant,

and establishes readily from artificial seeding. It persists under heavy grazing and can be seeded in mixtures with other herbs.

Family Scrophulariaceae

Penstemon eatonii ______________

Eaton or Firecracker Penstemon

Eaton or firecracker penstemon (fig. 34) is a bright

red to scarlet-flowered perennial semievergreen native that occurs primarily on rocky soils. It occurs in

Figure 33—Wasatch penstemon in a mixed subalpine forb community.

Figure 34—Eaton or firecracker

penstemon.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

mixed desert shrubs, sagebrush-grass, mountain

brush, and aspen communities. This species forms a

taproot. It is generally found on harsh, rocky sites; but

it also occurs on more favorable sites. Seed collection,

handling, and seeding techniques are similar to

those described for Palmer penstemon. Seeds can be

stored up to 12 years without much loss of viability

(Stevens and Jorgensen 1994). Eaton penstemon is particularly useful for seeding in pinyon-juniper, mountain

brush, and big sagebrush sites. It is not an abundant

seed producer but establishes easily and is a useful

forage species. Its brilliant flowers are extremely attractive and useful in landscape plantings. ‘Richfield’,

a source-identified variety, has been released.

Family Scrophulariaceae

Penstemon humilis ______________

Low Penstemon

This semievergreen native penstemon is a lowgrowing, 12 inch (30 cm) tall perennial forb with a

matted, much-branched root system. Flowers are

borne on short flower stalks and range in color from

deep-blue to violet-blue. Foliage is light green. Low

penstemon can be found in sagebrush-grass, pinyonjuniper, mountain brush, aspen, and subalpine communities, generally on gravelly, well-drained soils.

Planting procedures and seed collection are similar

to those of other penstemons. This penstemon is eaten

by wildlife and livestock and can be used to stabilize

erosive sites.

Family Scrophulariaceae

Penstemon pachyphyllus _________

Thickleaf Penstemon

Thickleaf penstemon, as its name indicates, has

broad, fleshy, thickened, bluish-green leaves covered

with wax. Flowers are light purple to bluish-violet.

Seeds are small with 335,000 per lb (739,000 per kg) at

100 percent purity. Seeds can be stored for up to 14

years without appreciable loss of viability (Stevens

and Jorgensen 1994). Seeding requirements are similar to those of Palmer penstemon. This species occurs

in eastern and northern Utah in salt desert shrub,

sagebrush-grass, pinyon-juniper, mountain brush, and

conifer communities. It is eaten by wildlife and livestock, and provides excellent soil protection.

Family Scrophulariaceae

Penstemon rydbergii ____________

Rydberg Penstemon

This species occurs in mountain brush, aspen, coniferous forests, and open parklands on basic and acid

soils. Rydberg penstemon has a dark violet-purple

flower and dark green basal leaves that are present

throughout the year. It is highly sought by sheep and

big game animals. Seed and seeding characteristics

are similar to those of Palmer penstemon.

Family Scrophulariaceae

Penstemon strictus ______________

Rocky Mountain Penstemon

This is a semievergreen penstemon that has an

abundance of dark, shiny green leaves and blue to

violet flowers borne on stalks 1 to 2.5 ft (30 to 76 cm)

tall. Rocky Mountain penstemon (fig. 35) does well on

rocky and sandy loam soils that range from weakly

acidic to alkaline. This species does best with 15 to

20 inches (38 to 50 cm) of annual precipitation. Seeding and seed handling requirements are similar to

those of Palmer penstemon. One variety, ‘Bandera’,

has been released, and seed is being produced commercially. ‘Bandera’ was selected for its longevity and

excellent forage and seed production. It is well adapted

to mountain brush, ponderosa pine, and spruce-fir

areas.

Family Rosaceae

Sanguisorba minor ______________

Small Burnet

A member of the rose family, small burnet (fig. 36),

is an introduced perennial with a basal rosette of

pinnately compound leaves arising from a caudex and

taproot. Numerous flowering stalks may grow from

2 to 20 inches (5 to 51 cm) in height. Flowers are

Figure 35—Rocky Mountain penstemon.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

formed in dense, terminal heads. Basal leaves remain

green nearly the entire year. Growth starts in early

spring, flowers appear in May and June, and seed

matures in August and early September.

Ecological Relationships and Distribution

There are approximately 30 species of burnet,

occurring primarily in Europe and the Middle East

(Hermann 1966). Two species, western burnet and

Alaskan burnet, are native to Western North America.

They are locally valued as forage plants (Hermann

1966; Hitchcock and others 1961). Selections of

small burnet from Mediterranean and Middle Eastern countries have been more widely adapted to sites

in the Intermountain region than any of the Western

North American species. Small burnet has proven to

be well adapted to pinyon-juniper, basin big sagebrush, and mountain big sagebrush communities, and

to drier exposures in the mountain brush and aspen

types. Small burnet will grows well on acid or alkaline

soils (Plummer 1977). The species is adapted to sandy

and clay soils, with silty and loamy soils the most

preferred (Wasser 1982). Small burnet has good winter hardiness and some fire and shade tolerance.

Introduction of S. magnollii, S. dictyocarpum, and

S. muricata have exhibited characteristics similar to

those of S. minor when grown on southwestern Idaho

rangelands (Shaw and Monsen 1983a).

Plant Culture

Excellent seed production, high seed quality, and

ease of seed processing and planting have contributed

to the widespread use of small burnet. Aerial or hand

broadcasting, drilling, and dribbling have all proven

to be successful means of seeding. Seed needs to be

Figure 36—Small burnet.

covered, but not more than 0.25 inch (6 cm) deep. Fall

and winter seedings have proven to be the most successful. Because of the rapid rate of germination and

seedling development, small burnet can usually be

successfully established by spring seedings. Small

burnet has been seeded widely throughout the West

(Plummer 1977; Plummer and others 1970a,b; Stevens

and others 1977). Following germination, growth is

rapid, resulting in good ground cover within 1 or 2

years. A large amount of foliage and seed is usually

produced, even during the first year of growth. Small

burnet is a excellent species to seed in mixtures.

Seed lots of small burnet germinate between 80 and

95 percent, and retains high viability when stored for

over 25 years (Stevens and Jorgensen 1994). Seeds

undergo afterripening, with germination improving

yearly up to 3 years following harvest (Stevens and

Jorgensen 1994). There are 55,000 seeds per lb (120,000

per kg) at 100 percent purity. Seed is marketed at

about 95 percent purity and 90 percent germination.

Seedling emergence and establishment for small burnet is high (5 to 8 percent) compared to many other

range species (Everett 1982).

Most plants appear to have a lifespan of 7 to 12 years,

but some have persisted to 20 years (Plummer and

others 1968; Stevens and others 1977b). Because seeds

are readily eaten by rodents (Everett and others 1978b)

and rabbits, little natural increase has occurred on

many areas. Where seeds are allowed to mature and

rodent and rabbit populations are low, reproduction

will take place. Dense stands of small burnet have remained in some seedings for as long as 25 years. Vegetative propagation can be accomplished by dividing

and transplanting the somewhat rhizomatous plants.

Uses and Management

Small burnet forage and seeds are highly preferred

at all seasons by livestock, big game, rodents, and

upland game birds (Autenrieth and others 1982;

Wasser 1982). Birds and rodents also make considerable use of the seeds. Seed caches and resulting plants

are common. Small burnet is particularly important

in late winter, early spring, and late summer when

other species provide less green forage. Plants produce an abundance of basal leaves that remain green

and persist throughout the dry summer and winter

months. Stand vigor and density may be reduced by

selective grazing during these periods.

Once established, small burnet can compete fairly

well with cheatgrass. This forb has done well on

mountain brush, pinyon-juniper, and upper sagebrush sites. Although it exhibits useful attributes

for the drier sagebrush and desert shrublands, it is

not adapted to areas receiving less than 10 inches

(25 cm) of moisture. Under favorable circumstances,
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

seedlings will establish when planted on dry sites.

However, seedlings often fail if seeded when spring

moisture is low. Mature plants are hardy and competitive and will persist, once established, on areas

receiving 8 to 10 inches (20 to 25 cm) of moisture.

Currently, some ecotypes express much better drought

tolerance and longevity than most cultivated strains.

These selections offer opportunity to expand the use

of this forb to drier environments.

Seeded areas should not be grazed for at least 2 years

following seeding. Because plants live from 7 to 12

years, every effort should be made to ensure that seed

crops are not grazed and are allowed to mature every

3 to 4 years.

Small burnet can be seeded, grazed, and maintained

in mixed grass and broadleaf herb plantings. Too

often, the species is heavily grazed with livestock in

the spring and early summer, and later grazed by

game animals in the fall and winter periods. Use can

be expected during any season as the plant retains

some green foliage throughout the entire year. Rodents, insects, and game animals will forage on the

plant whenever it is available. Consequently, livestock grazing must be regulated to compensate for

wildlife uses. The plant should be seeded heavy enough

to lessen concentrated use and satisfy animal needs.

If seeded in mixtures with perennial grasses, the

species can diminish in density because it is shortlived. Competitive grasses and heavy grazing can

prevent natural seeding. Sufficient plants have persisted on most wildlife plantings where livestock

grazing has been controlled.

Varieties and Ecotypes

‘Delar’ small burnet was released in 1979. It is

recommended for sites in the Intermountain region

that receive at least 12 inches (30 cm) of annual

precipitation. It will, however, establish and do fairly

well with as little as 10 inches (25 cm) of precipitation.

Forage production is excellent, even the first growing

season. Under irrigation ‘Delar’ has produced up to

1,050 lb seed per acre (1,177 kg per ha) (Howard 1981).

Additional selections are being investigated.

There are some selections that do well with as little

as 7 inches (18 cm) of annual precipitation, but these

strains are not commercially available.

Family Compositae

Senecio serra___________________

Butterweed Groundsel

This native perennial is a member of the sunflower

family. It grows in clumps of several stems 2 to 6 ft

(0.6 to 1.8 m) tall and has a woody base. It produces

numerous yellow heads about 0.25 inch (1.0 cm) in

diameter with black-tipped bracts (Welsh and others

1987). Flowering occurs in May to July.

Ecological Relationships and Distribution

A native in all Western States, butterweed groundsel (fig. 37) can be found in plant communities ranging

from sagebrush to subalpine. Welsh and others (1987)

describe two varieties with somewhat separate areas

of occurrence. Senecio serra var. serra exists in the

sagebrush, pinyon-juniper, mountain brush, lodgepole

pine, and spruce-fir communities at 6,000 to 10,000 ft

(1,800 to 3,000 m). Senecio serra var. admirabilies

grows in ponderosa pine and subalpine meadow communities (Welsh and others 1987). Both varieties are

early colonizers of disturbed sites and grow on sites

ranging from open, dry slopes with sandy to gravelly

loam soils to forest openings. Both varieties grow well

in association with wheatgrasses, bromes, wildryes,

and a large number of forbs.

Figure 37—Butterweed groundsel in full bloom.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Plant Culture

Seeds usually mature in September, and can be

collected by hand stripping or by using small paddles

to beat them into collection trays. Seeds are attached

to a white pappus that should be removed to facilitate

seeding. This can be accomplished by mechanical

debearding and screening. Seeds are very small, with

just under 3.5 million per lb (7.7 million per kg) at 100

percent purity and with the pappus removed. Seed is

best broadcast planted unless the pappus is removed,

after which seed can be drilled. Acceptable purity and

germination is 50 to 70 percent.

This species does well when seeded in mixtures with

other herbs because it has a fairly strong seedling. Fall

seeding is preferred. Seed should not be covered more

than 0.13 inch (3 mm).

Uses and Management

Butterweed groundsel is grazed extensively by sheep,

cattle, and big game (USDA Forest Service 1937). It

greens up early in the spring and retains a large

amount of green growth late into the summer. It can

tolerate considerable grazing, trampling, and drought.

It persists well in partial shade and frequently remains green late in the season. Deer and elk (Kufeld

1973; Kufeld and others 1973) make good use of it in

the spring and early summer. Cattle and sheep may

forage on the entire plant late in the season when the

leaves are partially dry. Because of its woody base and

stout, fibrous, rhizome system, this species does an

exceptionally good job of stabilizing disturbed soil. It

can be used to plant both dry and wet sites, although

it does not tolerate prolonged flooding. This species

can be seeded in disturbed riparian sites where the

water level has been lowered by down cutting of the

channel. It establishes well from broadcast seeding

and can be planted with a grass-forb mixture. Young

plants persist well amid competition, but the plants

are slow to attain maximum stature. Because it is slow

to achieve dominance, it can be seeded with slower

developing shrubs and trees without limiting their

survival.

Young plantings are usually not adversely effected

by grazing. Plants normally require 2 to 3 years to

fully establish. When planted with a grass mixture,

this forb persists equally well and does not require

special management. Established stands should be

allowed to set seed periodically.

Varieties and Ecotypes

There are no releases.

Family Compositae

Solidago canadensis ____________

Canada Goldenrod

A native perennial composite, Canada goldenrod

(fig. 38) produces stems 1 to 6 ft (0.3 to 1.9 m) tall that

grow from creeping rhizomes. Flowering occurs from

July to October. Pale golden-yellow flowers occur

along only one side of the curved flowering stalk.

Basal leaves are deciduous or become withered at

the time of anthesis (Welsh and others 1987).

Ecological Relationships and Distribution

Canada goldenrod occurs from southwestern

Canada to New Mexico and California in pinyonjuniper, mountain brush, ponderosa pine, and aspenspruce-fir types. It occurs along riparian areas, in

moist places and forest openings, and on dry slopes. It

grows on basic, neutral, and slightly acidic soils.

Plant Culture

Flowering occurs during midsummer, and seeds

mature in October. Generally, an abundance of seeds

are produced each year. Seeds are best collected by

hand stripping or beating. The pappus can be removed

from the seed using a debearder. Seeds are then

separated from the debris with a screening type

cleaner. The seed pappus is generally removed to

facilitate seeding. Cleaned seeds can be drilled or

broadcast planted. Uncleaned seeds can only be hand

broadcast. Seeds should not be covered more than

0.13 inch (3 mm) deep. Good results can be expected

Figure 38—Canada goldenrod.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

from broadcast seeding on prepared seedbeds and on

disturbed or eroding soils. Once established, plants

spread well from creeping rhizomes. Wildland collected seed lots are generally highly viable. Only fresh

seed should be planted because viability declines with

1 year of storage. This species does well when seeded

in mixtures with other herbs because it has fairly

vigorous seedlings. More successful stands usually

establish from fall seedings.

Uses and Management

Canada goldenrod is palatable to livestock and big

game. Some use is made at most seasons. Plants green

up early in the spring and remain green longer in the

fall than most associated species. Canada goldenrod

invades disturbed areas and provides an effective

ground cover. It can be used to control erosion on

extremely disturbed areas and is often used in watershed plantings. This species can be used to beautify

recreational areas, summer home sites, administrative areas, and other areas with high aesthetic value.

It is adapted to semiwet situations and can be used to

treat riparian disturbances where dry and semiwet

sites are closely aligned. Canada goldenrod establishes well by transplanting and can be used to provide

immediate ground cover. It persists with heavy trampling and grazing disturbances.

Canada goldenrod usually occurs in aspen and conifer forest communities. It recovers well following burning, logging, and grazing. Established stands should

be allowed to set seed periodically. Newly seeded

stands should not be grazed for at least 2 years

following planting.

Varieties and Ecotypes

There are no releases.

Family Malvaceae

Sphaeralcea grossulariifolia ______

Gooseberryleaf Globemallow

There are approximately 200 species of globemallow; about 20 are native to Western North America

(Hermann 1966). A member of mallow family, the

genus is variable and complex, and its taxonomy has

been extensively revised (Jefferies 1972; Kearney

1935; Welsh and others 1987). Gooseberryleaf

globemallow (fig. 39) is a densely, gray-green, pubescent perennial. Leaves are gooseberry-like. Floral

stalks grow from 1 to 2.5 ft (30 to 76 cm) tall. Plants

develop a branched taproot, and a near-surface fibrous root system. Flowers have showy orange to

reddish petals (Hitchcock and others 1961). Flowering

occurs from May to July, and seed matures unevenly

from June to August. Figure 39—Gooseberryleaf globemallow.

Ecological Relationships and Distribution

This species is a drought-tolerant native forb that

occurs in blackbrush, shadscale, rabbitbrush, sagebrush, and pinyon-juniper communities, and occasionally in the mountain brush zone on sites receiving

8 to 12 inches (200 to 300 mm) of annual precipitation

(Stevens and others 1985c). It is common on disturbed

and burned pinyon-juniper sites (Everett 1982;

Plummer and others 1968). It ranges throughout the

West from Washington to Arizona (Welsh and others

1987).

Gooseberryleaf globemallow prefers full sunlight. It

exhibits good winter hardiness, grazing tolerance,

resistant to burning, and considerable drought resistance. Gooseberryleaf globemallow is found in alkaline soils and tolerates moderate salinity, but not sodic

soils (Pendery and Rumbaugh 1986).

Plant Culture

Gooseberryleaf globemallow can be established on

harsh sites. It is one native forb that has been successfully seeded in blackbrush, shadscale, pinyon-juniper,

and sagebrush communities and in disturbances with

basic soils (Monsen and Plummer 1978; Plummer

1966, 1977).
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Seeds can be collected by hand or with mechanical

strippers. They are easily cleaned. There are just over

500,000 seeds per lb (1.1 million per kg) at 100 percent

purity. Seeds should be dusted with an appropriate

insecticide to prevent destruction by weevils (Pendery

and Rumbaugh 1986). Seed viability is, however, maintained for up to 15 years without any significant loss

(Stevens and Jorgensen 1994). Page and others (1966)

reported that low germination results from a combination of low fill and a hard seedcoat impregnated with

nonwetable substances. Mechanical or acid scarification can increase germination. It can be seeded on

disturbed and burned pinyon-juniper areas. Because

the species has fairly strong, competitive seedlings

and a moderate growth rate, it can be seeded in

mixtures with other species. Fall and winter seeding is

recommended (Monsen and Plummer 1978). Seeds

can be drill seeded and should be planted at depths of

0.25 inch (6 mm). Seeds may lie in the soil for years and

germinate when conditions are suitable (Plummer

and others 1968). This species can be successfully

transplanted and will reproduce from stem cuttings

(Everett 1982).

Uses and Management

Forage value of gooseberryleaf globemallow is rated

as good for sheep and cattle (Hermann 1966; Wasser

1982). Preference or palatability is rated as none to

fair for cattle and sheep (Hermann 1966), and fair to

excellent for antelope (Hancock 1966; Smith and Beale

1980), elk, and deer (Kufeld 1973; Kufeld and others

1973; Urness and McCullock 1973). Antelope especially prefer this herb when it is flowering (Pendery

and Rumbaugh 1986). Protein content during winter

and early spring has been found to be high, ranging

from 10 to 20 percent (Pendery and Rumbaugh 1986;

Urness and McCullock 1973). Gooseberryleaf

globemallow greens up early in spring and will green

up again in autumn after fall storms.

This species establishes well on disturbed sites from

seed or from transplanting. Because of its drought

resistance, continuous, colorful flowering, and ease of

establishment, this species has been recommended for

use in dryland ornamental landscaping (Natural Vegetation Committee 1973). Gooseberry globemallow is

quite competitive and can be used to suppress

cheatgrass and other annuals. It is one of only a few

native herbs that can be seeded on low elevation

Wyoming big sagebrush and shadscale sites.

Following germination gooseberry globemallow produces strong and persistent seedlings. It has a deep

taproot. Plants are tolerant of grazing, but can be

weakened by close grazing in late spring. Drought

tolerance is excellent, but plants should not be grazed

following extended periods of drought. Plant density

tends to fluctuate from year to year.

Varieties and Ecotypes

There are no releases.

Family Malvaceae

Sphaeralcea coccinea____________

Scarlet Globemallow

Scarlet globemallow (fig. 40) is a low-spreading, native

perennial with creeping rhizomes (Arnow 1971; McKell

and others 1979; Welsh and others 1987). It prefers

clay soils and can be found on hills and dry plains in

blackbrush, shadscale-greasewood, sagebrush, pinyonjuniper, mountain brush, and ponderosa pine communities from Southern Canada to Texas and Arizona

(Hermann 1966; Welsh and others 1987).

Scarlet globemallow exhibits considerable drought

resistance, and establishes especially well on disturbed sites. This species can be successfully seeded or

transplanted in blackbrush, shadscale, black greasewood, sagebrush, and pinyon-juniper types. It is particularly useful for planting on disturbed sites, as it

is drought tolerant and can spread by rhizomes. It is

well suited to adapted sites where wildfires frequently

occur.

Palatability for livestock and big game is rated from

poor to good (Hermann 1966; Kufeld 1973; Kufeld and

others 1973; McKell and others 1979; Wasser 1982).

Varieties and Ecotypes

There is one released germplasm, ‘ARS 2936’.

Figure 40—Scarlet globemallow.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Fabaceae

Trifolium _______________________

Clovers

Taylor (1985) reported that about 250 species of true

clovers occur throughout the world. All species possess

the typical papilionaceous legume flower with 10 stamens. Leaves usually consist of three leaflets. Most

have simple taproots; some produce stolons or rhizomes. About one-third are perennials and the others

are annuals. About one-third are self-pollinated; the

others are cross-pollinated and require bees for pollination (Taylor and others 1980). All species are able

to fix nitrogen if nodulated with strains of Rhizobium

(Allen and Allen 1981).

Within the contiguous United States, clovers are

most important as range plants from the Rocky Mountains west to the Pacific Coast (Crampton 1985).

Approximately 65 species occur in the Western States.

Most species, particularly many annuals, are abundant in the central valley of California (Crampton

1985; Küchler 1977).

Although native perennial clovers are found abundantly throughout the mountains, meadows and wetlands of the Intermountain area, and to a lesser extent

in the drier sagebrush habitats (Crampton 1985), few

natives have been utilized in range revegetation efforts. Most clovers of agricultural interest were introduced from Europe. There are about 15 species that

are used in North America (Hermann 1953; Hollowell

1960). Most are adapted to temperate climates and are

used to improve irrigated or native meadow pastures.

Because they are legumes, clovers are particularly

useful in fixing atmospheric nitrogen and enhancing

soil fertility. A cool, moist climate is required for the

best growth (Taylor 1985). Spring and fall periods are

favorable for adequate growth in the Intermountain

area.Various species demonstrate usefulness on a

variety of sites, and future trials will undoubtedly

result in more widespread use of different clovers.

Most clovers are quite palatable, shade tolerant, and

tolerant of flooding for short periods. Consequently,

they are useful for seeding semiwet meadows, and as

understorys in conifer forests and aspen stands. All

species attract use by wildlife and can be planted to

supply forage for extended periods. Most species demonstrate good regrowth and, thus, withstand heavy

grazing.

Seeds are small and include a high percentage of

hard seeds. Seeds should be treated with an inoculum

to assure establishment.

Family Fabaceae

Trifolium fragiferum _____________

Strawberry Clover

Strawberry clover was introduced to the West from

Eurasia and the Mediterranean region (Forde and

others 1981). It is not known when it was introduced

into the United States, but specimens were collected

in 1878 in Pennsylvania.

Strawberry clover is a low-growing stoloniferous

perennial used primarily as a pasture legume

(Townsend 1985). Stems are decumbent to creeping

and root at the nodes. Plants are less than 10 inches

(25 cm) tall. White to pink flower heads are borne at

the ends of each seed stalk (Gillett 1985; Harrington

1964; Hermann 1966). The flower and seed head

resemble a strawberry—hence the name. Plants are

pubescent or glabrous. The leaves are congested on

auxiliary peduncles that are curved and ascending

(Bendixen and others 1960).

Ecological Relationships and Distribution

Strawberry clover has spread or been seeded throughout North America and has become naturalized in

some locations (Townsend 1985). It is noted for its

ability to grow on wet saline or alkaline soils and has

demonstrated adaptability to wet meadows, streams,

and seeps. Bendixen and Peterson (1962) report that

strawberry clover withstands flooding because the

stolons exhibit a tropic response that causes the tips to

be elevated above the water level. Light, oxygen deficiency, and atmospheric conditions also influence the

tropic response.

Strawberry clover is not hindered by wet-saline

conditions; consequently, it is very useful on many

seeps and salty sites at a wide range of elevations.

Seedlings are not suppressed by high osmotic conditions that depress the growth of white clover (George

and Williams 1964).

Plant Culture

Strawberry clover can be drill or broadcast seeded

on a prepared seedbed. Seed coverage should be shallow, less than 0.25 inch (6.4 mm). When seeding irrigated pastures, a planting depth of about 0.50 inch

(13 mm) is recommended (Townsend 1985). Seeds are

small with about 290,000 seeds per lb (650,000 per kg)

at 100 percent purity. They require planting into a

firm seedbed. Evers (1982) found that drill seeding
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

produced only slightly higher yields than broadcasting. Seeding rates vary depending on seeding methods, but acceptable stands are achieved with 6 to 8 lb

per acre (6.7 to 9.0 kg per ha) when seeded alone, and

2 to 4 lb per acre (2.2 to 4.5 kg per ha) when planted in

a grass mixture.

Strawberry clover seedlings are easily established

and grow rapidly. The plant establishes well with

minimal seedbed preparation. Successful stands establish from broadcast seeding and natural spread.

This clover does well when seeded in mixtures with

grasses. When planted in mixtures under irrigation,

herbage production can be increased and better distributed throughout the growing season (Peterson

and others 1962).

Uses and Management

Forage and seeds of strawberry clover are used

extensively by livestock, big and small game, and

upland game birds.

Strawberry clover is especially well suited to close

and continuous grazing (Raguse and other 1971), and

when seeded on most rangeland sites, tends to attract

heavy use. In vitro digestibility compares favorably

with white clover, red clover, and alfalfa (Reed and

others 1980).

This forb can be successfully seeded onto depleted

riparian habitats, degraded meadows, and inland

saltgrass areas. It can be used to stabilize streambanks,

erodible soils, and related disturbances. It is particularly useful for seeding disturbances on saline soils

and sites subjected to frequent periods of flooding.

Varieties and Ecotypes

No selection or cultivar has been developed for

rangeland uses, although various cultivars have been

developed for pasture seedings in the United States

(Peterson and others 1962).

Family Fabaceae

Trifolium hybridum ______________

Alsike Clover

Alsike clover is a native of northern Europe that was

introduced into the United States in about 1839

(Townsend 1985). It was thought to be a hybrid of red

and white clover—hence its species name “hybridum”.

It is well adapted to high elevations within the Intermountain region where conditions are cool and moist.

It is a short-lived perennial, but can be managed to

reseed and maintain itself. Alsike clover is an excellent hay and pasture species (Townsend 1985), but is

also useful as a range forage species. It is adapted to

sites that are too wet or acidic for red clover or alfalfa

(Townsend 1985). Plants establish well from either

spring or fall plantings. Drilling or broadcast seeding

on a firm seedbed is recommended. Planting with

grasses to improve forage productivity is a common

practice (Grable and others 1965).

Seeding in mixtures at a rate of 2 to 4 lb per acre

(2.2 to 4.5 kg per ha) produces excellent stands. There

are approximately 68,000 seeds per lb (150,000 per kg).

Seeds have a hard seedcoat, but this does not prevent

good establishment even from spring seedings.

No cultivars have been developed for rangeland

conditions in the United States, but two Canadian

cultivars, ‘Aurora’ and ‘Dawn’, have been developed

for pasture plantings.

Family Fabaceae

Trifolium pratense _______________

Red Clover

Red clover is perhaps the most widely used pasture

clover in the United States. It is generally grown in

mixtures with grasses for intense pasture management. It is also grown for hay or silage production, or

as a soil improvement crop. Plants produce nitrogen

that favors growth of companion species and improves soil conditions (Smith and others 1985).

Red clover can be used as a short-term perennial or

winter annual to furnish immediate forage and improve soil fertility. It requires 20 or more inches (51

cm) of annual precipitation, but it is well suited to cut

over timber lands and moderately acid soils

(Hafenrichter and others 1968). It can be used to

improve meadows and degraded riparian sites.

Hafenrichter and others (1968) reported that the

commercial varieties—‘Kenland’, ‘Pennscott’, ‘Dollard’,

and ‘Lakeland’ are well adapted to conservation plantings in the Northwest. None have been extensively

evaluated for range sites in the Intermountain area.

Family Fabaceae

Trifolium repens ________________

White Clover

White clover is a shallow rooted perennial with

creeping stems that root at the joints. This introduced

European species has become naturalized in North

America in fairly moist areas on medium and high

elevation mountain ranges. White clover is winter

hardy and can withstand more extreme temperatures

than either red or alsike clover (USDA Forest Service

1937). Livestock and wildlife make good use of this

species. Areas of adaptation include sites with fertile

and well-drained soils that are moist throughout the

growing season, and along streams. It has good soil

stabilization characteristics. White clover is generally

seeded in a mixture with perennial grasses.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Family Compositae

Viguiera multiflora var. multiflora __

Showy Goldeneye

There are about 150 species of Viguiera. Most species occur in the new world (USDA Forest Service

1937). Two varieties of goldeneye, showy goldeneye

and Nevada goldeneye, occur within the Intermountain region (Welsh and others 1987).

Showy goldeneye (fig. 41), a member of the sunflower or compositae family, is a perennial native

forb growing from a short, branched, woody taproot.

It is usually 1 to 3 ft (30 to 91 cm) high with leaves

commonly more than 0.25 inch (6 mm) wide and 0.50

to 0.75 inch (12 to 19 mm) long. Two or more showy,

yellow, sunflower-like blossoms (heads) are borne on

each plant. Flowering begins in July. Seed matures in

late August and September. Basal leaves elongate and

green up shortly following snowmelt (Welsh and others 1987).

Figure 41—Showy goldeneye.

Ecological Relationships and Distribution

Showy goldeneye is a very attractive perennial, that

occurs throughout the Intermountain West at elevations between 3,500 to 11,000 ft (1,100 to 3,400 m)

(Stevens and others 1985c). Showy goldeneye can be

found from moderately moist habitats to dry, open,

rocky slopes in sagebrush-grass, pinyon-juniper, mountain brush, aspen (fig. 42), spruce-fir, subalpine, and

often in riparian communities (Welsh and others 1987).

This species is not restricted to full sunlight, but can

be found in partial shade and in densely wooded areas

(Arnow 1971; Plummer and others 1955). Showy goldeneye can be found on a wide variety of soils ranging

from heavy clays to gravel (USDA Forest Service

1937), and from acidic to basic conditions (Plummer

1977). It establishes quickly on disturbed sites, has a

rapidly developing seedling, and competes well with

annuals and perennials.

Plant Culture

Each showy goldeneye plant can support numerous,

golden yellow, sunflower-like heads that generally produce an abundance of seeds. Seeds are collected by

hand, mechanical beating, or stripping. Viability of wildland lots is only fair. Seed can be stored up to 7 years

without any major loss in viability (Stevens and

Jorgensen 1994). There are just over 1,000,000 seeds

per lb (2.2 million per kg) at 100 percent purity.

Showy goldeneye develops strong seedlings. Plants

establish quickly, and can spread from seed. Showy

goldeneye will invade disturbed sites and may quickly

become the dominant species. Seeds can be broadcast

or drilled with equal success. Seeds should not be

planted more than 0.25 inch (6 mm) deep. Seeding is

best accomplished in the fall or winter. The species

does well when planted as a component of a mixture

(Stevens and others 1981b).

Figure 42—Showy goldeneye in an aspen opening.
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Chapter 19 Forbs for Seeding Range and Wildlife Habitats

Principal Areas of Use

Showy goldeneye is an early spring-greening forb

that is sought by big game (Kufeld 1973) and livestock.

Foliage and flower heads are readily consumed by

cattle, sheep, deer, and elk. Birds also make considerable use of the seeds.

Strong seedlings and a rapid rate of growth enable

showy goldeneye to be seeded in conjunction with

other species. These features also contribute to its

establishment and spread into annual communities.

Profuse flowering of this species makes it a prime

candidate for summer home sites, campgrounds, administration sites, and other areas with high aesthetic

values. This species has potential as an erosion control

species on disturbed and burned sites.

Newly seeded areas should not be grazed for at least

2 years following planting. Established stands should

be allowed to set seed every 3 to 4 years.

Varieties and Ecotypes

There are no releases.

Family Compositae

Viguiera multiflora var.

nevadensis_____________________

Nevada Goldeneye

Compared to showy goldeneye, Nevada goldeneye

has narrower leaves (usually less than 5 mm wide) and

can be found on drier sites in southwestern Utah,

Nevada, and California. It occurs with pinyon and

juniper, Wyoming and basin big sagebrush, black

sage, and various saltbushes. Plummer (1966) recommends it as a candidate species to be used in seeding

areas in the salt desert shrublands. Coles (1982)

reported that Nevada goldeneye rapidly invaded Wyoming and basin big sagebrush burns in southwestern

Utah. He further reported that mule deer tend to seek

out and concentrate on areas with high densities of

Nevada showy goldeneye. The species invades roadsides and other disturbed areas. Seed production and

seeding procedures are similar to those described for

showy goldeneye.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 467

E. Durant McArthur

Stephen B. Monsen

20

Chapter

Chenopod plants (Family Chenopodiaceae) are distributed

worldwide but are especially prominent in some wet and dry

saline or alkaline situations. Chenopods are both herbaceous and

woody. The relative proportions of life-forms in the family is

demonstrated by data from the important center of chenopod

diversity in south-central Asia, where n = 341 species: 76 percent

are herbaceous (mostly annual), 23 percent are shrubs or subshrubs, and 1 percent is arborescent (McArthur and Sanderson

1984; Shishkin 1936). Chenopod shrubs grow over wide expanses

of Intermountain rangelands as well as on large saline and

alkaline tracts on all continents, except Antarctica. In the Intermountain area, seven genera (table 1) with 28 species make

important contributions to the landscapes or to revegetation

needs.

As a group the chenopod shrubs exhibit the ability to grow in

low fertility, salt-bearing soils as well as on more favorable sites.

Their presence is essential for maintaining a stable soil in many

xeric environments where soil is too salty or dry for most other

classes of plants to live (Blauer and others 1976; Goodall 1982;

Sanderson and Stutz 1994b; Wilkins and Klopatek 1984). Chenopod shrubs grow well in high concentrations of calcium and

potassium salts and can endure considerable concentrations of

Chenopod Shrubs
468 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

sodium salts (Gates and others 1956; McNulty 1969;

Moore and others 1972). Certain ones grow through a

much wider range of alkalinity than others (table 2).

A few tolerate such high salt concentrations that they

are the only plant life. As a group they are especially

suited for planting on depleted salt desert shrub type

range areas as well as on construction scars, mine

spoils, and roadcuts (McArthur and others 1978b;

Plummer 1977).

Within species, large differences exist between

ecotypes in morphological and physiological traits.

The smaller ecotypes are usually from dry, severe

sites, whereas the larger ones are from more favorable

areas. Size of the shrubs is, to a large extent, genetically controlled as demonstrated by maintenance of

ecotypic form in accessions grown together in uniform

gardens (McArthur and others 1983a). Furthermore,

Stutz and Sanderson (1983) have shown in shadscale that the smallest ecotypes growing on the most

severe sites are higher polyploids, whereas lower

ploidy levels occur on more favorable mesic sites.

The gigas form of fourwing saltbush reported by

Stutz and others (1975) is a diploid. Dunford (1985)

Chapter Contents by Species

Atriplex canescens (fourwing saltbush)................................................................................................. 471

Atriplex confertifolia (shadscale) ........................................................................................................... 476

Atriplex corrugata (mat saltbush) .......................................................................................................... 479

Atriplex cuneata (Castle Valley clover or cuneate saltbush) ................................................................. 480

Atriplex gardneri (gardner saltbush) ...................................................................................................... 481

Gardner saltbush relatives .................................................................................................................... 481

Atriplex buxifolia ............................................................................................................................. 481

Atriplex falcata ................................................................................................................................ 481

Atriplex tridentata ............................................................................................................................ 481

Other Saltbushes .................................................................................................................................. 482

Atriplex lentiformis .......................................................................................................................... 482

Atriplex obovata .............................................................................................................................. 482

Atriplex polycarpa ........................................................................................................................... 482

Atriplex hymenelytra ....................................................................................................................... 482

Ceratoides lanata (winterfat) ................................................................................................................. 484

Grayia brandegei (spineless hopsage) ................................................................................................. 486

Grayia spinosa (spiny hopsage)............................................................................................................ 487

Kochia americana (gray molly).............................................................................................................. 488

Kochia prostrata (forage kochia) ........................................................................................................... 489

Sarcobatus vermiculatus (black greasewood) ...................................................................................... 490

Other chenopod shrubs......................................................................................................................... 491

Allenrolfea occidentalis (iodine bush) ............................................................................................. 491

Camphorosoma monspeliaca (Mediterranean camphorfume) ....................................................... 491

Suaeda torreyana (desert sumpbush, desert seepweed, or desert blite) ....................................... 491

reported consistent morphological differences among

the chromosome races of fourwing saltbush. Even

though genetic factors are important, moisture does,

of course, markedly affect luxuriance and growth.

Wallace and others (1974) reported that 40 percent of

the biomass of fourwing saltbush is in the root system.

Shrubby chenopods have a chromosome number

based on x = 9. Polyploids occur in most genera

(Blauer and others 1976; McArthur and Sanderson

1984). They are wind pollinated, occur in sympatric

distributions, are monoecious, dioecious, or even

trioecious—factors that lead to common interspecific

hybridization (McArthur 1989). Woody chenopods

show remarkable capability to hybridize both naturally (Sanderson and Stutz 1994b; Stutz 1978, 1984)

and under controlled conditions (Blauer and others

1976; Drobnick and Plummer 1966; Stutz 1982).

Most of the hybridization occurs within Atriplex,

although some possible intergeneric hybrids have

been reported (Blauer and others 1976).

Hybridization allows opportunity to create new

gene combinations to fill new niches—for example,

those in newly created mine spoils. Stutz (1982),
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Chapter 20 Chenopod Shrubs

Table 1—Shrubby chenopods in Western America useful or

potentially useful for restoration and revegetation

(adapted and expanded from McArthur and others

1978b).

Scientific name Common name

Allenrolfea occidentalis Iodine bush or inkbush

Atriplex aptera Wingless saltbush

Atriplex bonnevillensis Bonneville saltbush

Atriplex canescens Fourwing saltbush

Atriplex confertifolia Shadscale

Atriplex corrugata Mat saltbush

Atriplex cuneata Castle Valley clover saltbush

or cuneate saltbush

Atriplex falcata Falcate saltbush

Atriplex gardneri Gardner saltbush

Atriplex hymenelytra Desert holly saltbush

Atriplex lentiformis Big saltbush or quailbush

Atriplex navajoensis Navajo saltbush

Atriplex obovata Broadscale saltbush

Atriplex polycarpa Cattle saltbush or allscale

saltbush

Atriplex robusta Robust saltbush

Atriplex semibaccataa Australian saltbush

Atriplex tridentata Trident saltbush

Camphosoma monspeliacaa Mediterranean

camphorfume

Ceratoides lanata Winterfat

Ceratoides latensa Pamirian winterfat

Grayia brandegei Spineless hopsage

Grayia spinosa Spiny hopsage

Kochia americana Gray molly

Kochia prostrataa Forage kochia

Sarcobatus vermiculatus Black greasewood

Sarcobatus baileyi Bailey greasewood

Suaeda suffrutescens Desert sumpbush

Zuckia arizonica Arizona zuckia

a

Introduced species.

Table 2—Physical dimensions and tolerance to alkalinity of some chenopod shrub

taxa (from McArthur and others 1978b).

Adaptation to alkalinity

Species Height/crown Optimum Range

Inches (cm)

Atriplex canescens 12-78/12-178

(Fourwing saltbush) (30-200/30-450) Low Low-medium

Atriplex confertifolia 6-47/6-30

(Shadscale) (15-120/15-75) High Low-high

Atriplex corrugata 1-8/8-40

(Mat saltbush) (2-20/20-100) Medium Medium-high

Grayia spinosa 12-47/12-70

(Spiny hopsage) (30-120/30-180) Low Low

Ceratoides lanata 2-47/6-30

(Winterfat) (6-120/15-75) Low Low-medium

Allenrolfea occidentalis 6-47/12-70

(Iodine bush) (15-120/30-180) High High

Sarcobatus vermiculatus 40-118/40-78

(Black greasewood) (100-300/100-200) Medium Low-high

Ferguson and Frischknecht (1985), and Stutz and

Estrada (1995) found that several hybrid combinations within Atriplex performed well on mine spoils in

Utah and New Mexico. Blauer and others (1976),

Carlson (1984), Stutz (1983, 1984), and Stutz and

Carlson (1985) showed that there is ample opportunity to: (1) make selections from natural populations,

(2) locate natural hybrids, and (3) make artificial

hybrids. All three plant classes can be useful in providing plant materials to rehabilitate harsh disturbed

landscapes. Sanderson and colleagues (Sanderson

and Stutz 1984, 1994a; Sanderson and others 1988,

1990) have identified flavonoid compounds that are

useful genetic markers in population biology and

hybridization studies.

Woody chenopods show remarkable ability to grow

through a wide climatic variation. Some grow in the

coldest parts of the northern desert shrublands and

adjacent grasslands to warmer desert ranges in the

Southwestern United States and Mexico (fig. 1).

Three examples are Gardner saltbush, black greasewood, and winterfat. Several chenopods readily establish, grow, and produce in areas of severe aridity or

where the average annual precipitation may be less

than 5 inches (12.7 cm). Important among these are

winterfat, shadscale, and falcate saltbush. However,

they can also quickly take advantage of three to four

times this precipitation to make luxuriant growth.

While the shrubby chenopods are most characteristic of the salt desert shrub type in Western North

America, some grow well in limey soils on mountain

ranges in association with ponderosa pine, Utah

juniper, pinyon pine, Saskatoon serviceberry, and

Utah serviceberry. Winterfat, fourwing saltbush, and

spiny hopsage grow best in these areas. However, on
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Chapter 20 Chenopod Shrubs

highly alkaline extrusions in these higher elevations

are precipitation areas on mountain ranges where

shrubby chenopods may grow to the virtual exclusion

of other species (McArthur and others 1978b).

All shrubby chenopods bear a carpellate, loculed

one-celled fruit known as a utricle, within which is a

single seed. Actual seeds are small, usually no larger

than an alfalfa seed. The utricle wall may be quite

thin and easily removed from the seed, as in winterfat; or may be hard, as found in shadscale and some

accessions of fourwing saltbush. Production of fruits

varies widely among plants and from season to season

as does percentage fill (Crofts 1977; Gamrath 1972;

McArthur and others 1978b; Monsen and McArthur

1985; Springfield 1970b). To a considerable degree

this relates to how favorable the growing season may

be. Unless seed coats are restrictive, germination

takes place quickly in a favorable environment of

temperature and moisture. Utricles having indurated

walls, such as in some ecotypes of fourwing saltbush

and shadscale, may lay a year or more in the ground

before germination. This may provide some survival

value for the species because it ensures that some

seeds remain in the ground to germinate and establish

when temperature and moisture are most favorable.

Apparently, saltbush seeds are not highly palatable

to small rodents (Everett and others 1978) so they are

retained in the soil. Some buildup occurs in the

ground of fruits that have a tendency for delayed

germination. Consequently, in favorable years many

seedlings become established. Young seedlings, however, are particularly susceptible to rodent depredation (Nord and Green 1977; Young and others 1984e).

Shrubby chenopods express a number of traits

that aid in artificial revegetation. The appendages

attached to the utricles can be removed from most

species. Except for winterfat, all seeds can be cleaned

to a form that is easily handled and planted with

conventional equipment (chapter 19). Cleaned seeds

are of a size and shape that different seeding rates

can be employed. Within a specific collection, seed

germination features are uniform. However, seed

germination is not so specific to a particular set of

climatic conditions that all seeds germinate at any

one time. An extended period of germination often

assures the survival of a satisfactory number of

plants. However, seed germination of winterfat is

usually confined to a short duration. Seedlings and

young plants are quite vigorous and are able to compete well with herbs (Giunta and others 1975).

The growth rate of nearly all shrubs is excellent.

Even under adverse conditions plants grow remarkably well, often exceeding most herbs. Plants are well

adapted to infertile soils and can be used to restore

vegetation throughout variable, but harsh, situations. Young plantings withstand heavy browsing by

game, livestock, and rodents. Individual plants normally reach maturity within 3 to 5 years, and natural

spread can be expected thereafter. Once established,

young plants persist under adverse climatic conditions. In addition, established plants provide high

yields even when grown with a dense understory.

Chenopod shrubs frequently grow with a sparse

understory but persist and grow well when seeded

with numerous other plants. These shrubs normally

occur throughout some principal sites or vegetative

conditions important to both livestock and wildlife.

The plants can be increased in density or seeded with

a combination of herbaceous plants to improve forage

or habitat. Many other shrubs exist within specific

site conditions and do not grow well when moved to

adjacent locations. Some ecotypes of many chenopod

shrubs also exhibit specific site adaptation requirements, but others can be extended to additional sites

(see McArthur and others 1983a).

Most shrubby chenopods are highly nutritious. They

have higher protein content than most plants, particularly in the winter (Chatterton and others 1971;

Esplin and others 1937; Tiedemann and others 1984a;

Welch and Monsen 1981). On several species and

Figure 1—Distribution of shrubby chenopods

in general and winterfat, greasewood, and the

Gardner saltbush complex in particular. Distributions from Hall and Clements (1923), Branson

(1966), and McArthur and others (1978b).

British

Columbia Alberta

Saskatchewan

Montana

Wash.

Oreg.

Idaho

Nevada

Utah

Colo.

Wyoming

N. Dak.

S. Dak.

Neb.

Kan.

Okla.

Texas

New

Mexico Arizona

Calif.

Mexico Shrubby chenopods

Ceratoides lanata

Atriplex gardneri complex

Sarcobatus vermiculatus
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 471

Chapter 20 Chenopod Shrubs

accessions, leaves persist and remain green well

through winter. These are preferred by grazing animals during the winter and provide an important

source of protein, which is vital to the well being of

browsing animals on desert ranges of the West.

Without doubt, the shrubby chenopods have an

important future for development of cover and forage

on the generally basic soils of arid ranges worldwide. They are widely used in land rehabilitation

programs (Aldon and Oakes 1982; Plummer 1984;

Tiedemann and others 1984b).

Recently, Nechaeva and others (1977), Rumbaugh

and others (1981, 1982), and Ostyina and others (1984)

showed that most shrubs, especially some chenopods,

provide complementary forage to grasses in mixed

stands on semiarid ranges. Forage quality is enhanced

in the mixed stands and the effective season of grazing

is expanded. In addition, the forage resource is made

less susceptible to attack from insects and diseases

than are monocultures (Moore and others 1982).

Nord and Green (1977) identified several chenopod

shrubs with potential to reduce the hazards of fire in

the California chaparral. Among these were North

American fourwing saltbush, Castle Valley clover saltbush, Gardner saltbush, allscale saltbush, and three

Australian saltbush species.

From 1985 through 1988 a significant die-off of

chenopod (and some other families) shrubs occurred

in the Intermountain area (McArthur and others

1990b; Nelson and others 1989; Wallace and Nelson

1990). This die-off was particularly serious for

shadscale. The die-off was not unique in time, but it

was large in scale; and recovery will be complicated by

the recent large-scale invasion of exotic annuals into

chenopod shrublands.

Important characteristics of a number of chenopod

shrubs follow. Seeding recommendation for principal

vegetative types and conditions are discussed in chapter 17. Chenopod shrubs adapted to these vegetative

types and conditions are included in the seeding recommendations. Seed characteristics are found in chapters 24 and 26.

Included in the remainder of this chapter are a brief

species description, ecological relationship, distribution, culture requirements, use, improved varieties,

and management of each of the species appearing in

the chapter contents list.

Atriplex canescens ______________

Fourwing saltbush

Fourwing saltbush is an upright shrub (fig. 2).

Its height may vary from 1 ft (0.3 m) to 6.7 ft (2.04 m),

depending on site conditions and genotype. It branches

freely from the base. Its mature branches are quite

brittle. Both young leaves and twigs are gray-green

because of the white, scurfy vestiture. The linear to

oblanceolate or spatulate evergreen leaves are 1 to 4

cm long and 2 to 6 mm wide. Fourwing saltbush is

usually dioecious with a low frequency of monoecy.

Pistillate (female) flowers are small and inconspicuous with no flower parts except pistils. Each pistil is

enclosed by a pair of small bracts that are united along

their edges to form winglike expansions. In addition,

each bract of the pair has a wing down the middle; so

the utricle (fruit) has, at maturity, a varying prominence of four wings (fig. 3, 4). The yellow to red to

brown staminate flowers are more or less naked anthers (only a few bracts are present) borne in glomerules 2 to 3 mm wide (Blauer and others 1976; McArthur

and others 1978c). Sex expression in fourwing saltbush is apparently a dioecious system for the only

diploid population known in Utah (Freeman and

McArthur 1989; McArthur and Freeman 1982; Stutz

and others 1975). Barrow (1987) has shown that

diploid New Mexico and Texas populations may contain rare monoecious plants. At the higher and more

common (tetraploid, hexaploid) ploidy levels, fourwing

Figure 2—Fourwing saltbush is an upright

shrub. (A) A large plant along the Snake River,

Payette County, ID, opposite Nyssa, OR. (B)

A ‘Rincon’ fourwing saltbush orchard at the

Upper Colorado Environmental Plant Center,

Meeker, CO.

A

B
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Chapter 20 Chenopod Shrubs

Figure 4—Fruits

(utricles) of various

chenopod species.

(A) Fourwing saltbush. (B) Shadscale.

(C) Mat saltbush.

(D) Castle Valley clover

saltbush. (E) Trident

saltbush. (F) Broadscale

saltbush. (G) Hybrid of

fourwing saltbush x

shadscale. (H) Fourwing saltbush x trident

saltbush. (I) Winterfat.

(J) Spineless hopsage.

(K) Spiny hopsage.

(L) Gray molly. (M) Forage kochia. (N) Black

greasewood. (After

Blauer and others

1976).

AB C

Figure 3—Flowers and fruits of fourwing saltbush (A) A pistillate (female) plant in fruit (McArthur 722, seeded

near Mountain Home, Elmore County, ID, from seed collected near Panaca, Lincoln County, NV; 0.65x). (B)

A staminate (male) plant in flower (McArthur 378, St. George, Washington County, UT; 0.87x). (C) A

monoecious plant (McArthur 723, Payette County, ID; 0.56x).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 473

Chapter 20 Chenopod Shrubs

saltbush is trioecious (three sexual states). In the

trioecious condition, some plants are lifetime constant

females, others lifetime constant males, and some

plants are apparently able to avoid stressful situations by switching away from the physiologically taxing female state (Barrow 1987; Freeman and McArthur

1989; Freeman and others 1984; McArthur 1977;

McArthur and Freeman 1982; McArthur and others

1992; Noller and others 1984). Gender differences

appear to be adaptive and allow fuller exploitation of

habitats (Freeman and others 1993, 1997; Pendleton

and others 1992). Dr. Daniel J. Fairbanks and his

students at Brigham Young University have made

progress toward understanding the molecular genetics basis of sex expression in woody Atriplex (Freeman

and others 1993, 1997; Randell 1997; Ruas 1997).

Ecological Relationships and Distribution

The species grows in a variety of soil types from the

Great Plains to Pacific Coast ranges and from

Canada to Mexico (fig. 1); it occurs from elevations

below sea level to about 8,000 ft (2,400 m). This

shrub is well suited to deep, well-drained sandy soil,

sand dunes, gravelly washes, mesas, ridges, and

slopes; vigorous plants have been found in heavy

clays. Ecotypes of the species may grow in widely

different soil types and plant communities (Dunford

1985; McArthur and others 1978b; Richardson and

McKell 1980). Fourwing saltbush is frequently found

intermixed with black greasewood, shadscale, Castle

Valley clover saltbush, basin big sagebrush, and occasionally black sagebrush and pinyon-juniper. In many

Western regions it is the main shrub over extensive

grassland areas. Important grasses on these tracts

are galleta grass and blue grama. It is not unusual to

find it growing in association with spring-growing

grasses such as bluebunch wheatgrass and Sandberg

bluegrass.

Plant Culture

This species may be propagated by direct seeding,

transplanting nursery or container stock, or by rooted

stem cuttings (Hennessey 1985; Kay and others 1977b;

McArthur and others 1978c; Richardson and others

1979; nine papers in the revegetation section of

Tiedemann and others 1984b). Shaw and Monsen

(1984) give information on growing fourwing saltbush

and other chenopod shrubs in a nursery setting to

provide suitable bare root transplant stock. Ferguson

(1980) recommended a growth medium of 50 percent

sphagnum peat moss, 30 percent arcillite aggregate,

and 20 percent vermiculite for growing Bonneville

saltbush and other plant species native to semiarid

alkaline soils. Use has been made of rooted stem

cuttings (McArthur and others 1984a) to establish

seed orchards that combine desired vegetative and

floral traits together for seed production (Briggs 1984;

McArthur and others 1978c; Noller and others 1984;

Van Epps and Benson 1979). Such orchards may not

only provide superior seed but may also provide income from marginal agricultural land.

Most wildland stands of fourwing saltbush produce

a harvestable seed crop at least 3 out of every 5 years.

Production and seed quality vary among sites, but

fourwing saltbush is an abundant seed producer. Seed

is collected by hand stripping or beating, although

some vacuum and reel-type harvestors have been

modified for this purpose (Monsen 1978; Monsen and

others 1985b; Van Epps 1978). Seed is reasonably easy

to harvest and process. Consequently, seed costs are

not excessive. Seed is harvested in late fall and early

winter, and seed can be cleaned immediately after

collection. Various commercial growers have recently

established seed orchards of fourwing saltbush.

Seed yields from these centers appear to be more

reliable than wildland stands. Seed quality differences may be better as well (Briggs 1984; McArthur

and others 1978c; Noller and others 1984; Van Epps

and Benson 1979).

The wings are removed from the utricles using

hammermills, flail-type beaters, Dybvigs or cylinder

grinders (chapter 24). If correctly operated, these

cleaners can be used to fracture the utricle to aid seed

germination. Removal of the wings greatly reduces

the bulk and allows the utricles to be planted with

most seeding equipment.

Only about 50 percent of the utricles contain a full

viable seed. Seed “fill” can be increased by cultivation and culture of nursery plantings. Seed fill from

wildland collections may vary from a low of 10 to

20 percent to as high as 80 percent (Crofts 1977;

Gamrath 1972; Gerard 1978; Springfield 1970b).

Seeds should be carefully inspected prior to harvest

to ensure the acquisition of high quality seed. Acceptable fill is about 40 percent. Seeds can be stored for a

number of years (up to 7) in an open warehouse

without loss of viability. Even after 15 years storage,

viability is acceptable (Stevens and others 1981a).

Seeds of fourwing saltbush normally require 20 to

30 days of stratification to initiate and assure uniform

germination. Thick impermeable utricles often require much longer periods before water and oxygen

can gain entry. Springfield (1966) and Potter and

others (1986) found temperatures required for germination varied among ecotypes, but the optimum

range was 15 to 18 ∞C (59.0 to 64.4 ∞F). Potter and

others (1986) reported seeds are able to germinate at

low media osmotic potential if optimum temperatures are maintained. However, ecotypes differed in

their tolerance to moisture stress. Seeds germinate

early in the spring, and if fall seeded, high mortality
474 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

to spring frost can occur. Consequently, spring

plantings are sometimes recommended. However,

seeds must be planted early enough to facilitate

stratification. If this is not possible or if seeds with

impermeable seedcoats or long-term stratification

period requirements are to be planted, fall seeding

should be conducted. Where the climate permits, fall

or winter planting is preferred.

Seed should not be covered more than 0.5 inch

(1.2 cm) deep. Broadcast seeding followed by chaining

has produced excellent stands. Satisfactory stands

have also developed from drill and interseeding. Plants

can also establish when seeded in separate rows with

herbs using a conventional rangeland drill. However,

fourwing saltbush seedlings are not able to compete

with herbs if drilled together in a single row. Separating seeds of different species and reducing herb seedling density by broadcast seeding is beneficial to shrub

establishment. Most satisfactory stands have been

achieved when fourwing saltbush was seeded at 1 to

2 lb PLS/acre (1.2 to 2.2 kg/ha) or between 2 to 4 lb

(2.2 to 4.5 kg) bulk or cleaned utricles. Between 0.3 and

4.67 percent of all viable seed planted ultimately

emerge and develop into mature plants (Richardson

and others 1986).

Fourwing saltbush is well adapted to interseeding.

Plants can be established in strips or furrows separate

from herbs (Giunta and others 1975; Ostyina and

others 1984; Rumbaugh and others 1982; Stevens

and others 1981b). Interseeding is a useful means of

seeding the shrub into established stands of perennial

herbs. Narrow furrows or strips approximately 28 to

30 inches (71.1 to 76.2 cm) in width can be created

using various interseeders (Giunta and others 1975;

Stevens and others 1981b). Herbicides have also been

used for this purpose (Petersen and others 1990).

Shrub seeds are placed in the clearing. Furrows can be

interspaced at desired intervals, usually 10 to 12 ft

(3.0 to 3.7 m) apart. Fourwing saltbush grows profusely and quickly produces a dense overstory.

Fourwing saltbush grows well with understory herbs

and other shrubs. When grown in association with

other species, herbage production of the fourwing

saltbush plants and the understory herbs remains

quite high (Rumbaugh and others 1981, 1982).

Fourwing saltbush is naturally susceptible to

basal decay disease in both plantation and natural

populations (Nelson and Welch 1984). This is not

usually a serious problem. Insects can be troublesome to fourwing saltbush plants and populations,

especially when plants occur at heavy densities

(Haws and others 1984; Moore and Stevens 1984). In

seed orchards the case-bearing moth can be devastating if not controlled. Malathion will effectively control

this pest (Moore and Stevens 1984).

Uses and Management

Fourwing saltbush is one of the most valuable forage

shrubs in arid rangelands because of its abundance,

accessibility, palatability, size, evergreen habit, nutritive value, rate of growth, and large volume of foliage

(McArthur and others 1978; Petersen and others 1987).

Its leaves, stems, and utricles provide browse in all

seasons. Sanderson and others (1987) have shown

that saponin content may be important in the palatability of fourwing saltbush. In addition to providing

forage and cover, this species is one of the most

important shrubs for use in rehabilitation of depleted

rangelands and in soil stabilization projects in Western desert areas (Blaisdell and Holmgren 1984; Blauer

and others 1976; Plummer 1984; Plummer and others

1966b). Stutz (1982) and Stutz and Carlson (1985)

have shown that fourwing saltbush hybridized with

other saltbush species can be effective in an in situ

selection process in revegetation efforts. Many lines of

potentially adaptive plants are established under this

technique. Additional onsite hybridization and rapid

natural selection allow sustained establishment of

adapted plant materials.

Fourwing saltbush is widely planted in revegetation

projects. When the seedings are carefully planted they

are usually successful. Evidence shows that fourwing

saltbush increases the value of otherwise herbaceous

pastures (Ostyina and others 1984; Rumbaugh and

others 1982; Ueckert and others 1990). Giunta and

others (1975) demonstrated the value of interplanting

fourwing saltbush in cheatgrass-dominated ranges.

Sometimes, however, stands do not persist, perhaps

because of planting in sites that will not support the

species or because the source planted is not adapted to

the site. The species is a secondary or facultative

selenium absorber and thus may be mildly poisonous

where selenium occurs in the soil (Davis 1972;

Kingsbury 1964). Minor losses have occurred when

livestock had little or nothing else to eat (Hitchcock

and others 1964).

In propagating fourwing saltbush, it is especially

important to be sure that the strain being planted is

adapted to the site. If possible, seed should be from a

source growing in a climate similar to that of the

site being seeded unless other tested strains are

known to have similar or better adaptation. Strains

adapted to the mountain brush types, where soils

tend to be neutral, mature rapidly; but in deeper soils

of this type, the life span has often been not more than

10 years. On drier sites having higher alkaline content, life span of the same types has been much longer.

However, persistence of different accessions has been

quite erratic. Various biological and climatic influences apparently affect persistence. Plants established

from seed collected in the blackbrush-mesquite belt in
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 475

Chapter 20 Chenopod Shrubs

southern Utah have not persisted in colder parts of

the State. Seed collected from colder areas have persisted well on warmer sites (Plummer and others

1966, 1968).

Varieties and Ecotypes

Fourwing saltbush probably has the greatest potential among the woody chenopods for reclaiming

arid ranges. It is one of the most widespread and

adaptable western shrubs. Fourwing saltbush is the

center of a large species complex. It hybridizes widely

with other woody saltbushes (Blauer and others 1976;

Sanderson and Stutz 1994b; Stutz 1978, 1984). Several subsidiary species have been recognized. These

species are all fourwinged and likely have A. canescens

in their ancestry or are directly derived from that

species. Five of the allies of fourwing saltbush (A.

aptera, A. bonnevillensis, A. garrettii, A. navajohensis,

and A. “robusta”) are listed in table 3. Each species has

a habitat requirement different from A. canescens, and

consequently would be useful for restoration and revegetation of particular kinds of sites (table 3).

Fourwing saltbush proper is composed of widely

occurring polyploid races (x = 9). Diploids are mostly

restricted to the southern portion of the species range—

New Mexico and Texas and south (Dunford 1984;

Stutz and Sanderson 1979). Hexaploids, too, are restricted mostly to the west Texas and New Mexico

area. Other high polyploids (up to 20x), usually restricted endemics, occur in the southern portions of

the species range (Sanderson and Stutz 1994a; Stutz

and Sanderson 1979). Tetraploids (4x = 2n = 36) are

the common plants of the Intermountain areas.

Fourwing saltbush, with its outcrossing habit and

common polyploidy, exhibits both allo- and autopolyploidy (Stutz 1978, 1984). Many of the common

tetraploids of the Intermountain area appear to be

autotetraploids (McArthur and others 1986).

Differences have been demonstrated among several

accessions in winter protein content (Welch and Monsen

1981, 1984) and in both growth performance and

survival (Geist and Edgerton 1984; McArthur and

others 1983a; Shaw and others 1984; Young and others 1984e). In planting fourwing saltbush, either a

proven accession or accessions obtained from nearby

or from higher latitudes or higher elevations than

the planting site should be used (McArthur and

others 1983a; Van Epps 1975).

Three releases of fourwing saltbush of proven quality and adaptability have been made (Carlson 1984).

Additional work is under way to provide more plant

material with known characteristics by identifying

and testing promising accessions and through hybridization and selection procedures (Carlson 1984; Stutz

and Carlson 1985).

The three releases are ‘Rincon’, ‘Marana’, and

‘Wytana’. ‘Rincon’ is well adapted to a wide range of

Table 3—Other fourwing saltbushes in addition to Atriplex canescens.

Postulated Distinguishing

Species origin Habitat Distribution characteristics References

Atriplex aptera A. canescens Clay hill Upper Missouri Diminutive (or lacking) Brown 1956;

x badlands River Drainage, utricle wings. Small Stutz 1978;

A. gardneri Montana, Wyoming, 24 inches (60 cm) Stutz and

the Dakotas plants others 1979.

Atriplex bonnevillensis A. canescens Alkaline Eastern and Central Small 30 inches Hanson 1962;

playas Great Basin, Utah (75 cm) highly Frischknecht and

and Nevada variable plants Ferguson 1979.

Atriplex garretti — Rocky, sandy Colorado River and Early flowering, Brown 1956;

soils; in tributary canyons, succulent, yellow- Hanson 1962;

canyons southern Utah, green leaves. Small Blauer and others

northern Arizona 16 inches (40 cm) 1976.

Atriplex navajoensis — Rocky, sandy Vicinity of Navajo Woody, yellow- Hanson 1962.

soils Bridge, Coconino green leaves.

County, AZ Up to 40 inches

(100 cm) tall

Atriplex “robusta”

a A. canescens Highway East of Wendover, Rapid growth rate, Stutz and others

x disturbances Tooele County, UT utricles rarely 1979.

A. tridentata in salt flats winged but with

variously shaped

protuberances

a

This taxa has not been formally named.
476 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

Figure 5—Shadscale growing near Elberta, Utah

County, UT.

soil textures: sandy areas, gravely washes, loamy

soils, heavy clay soils, and moderately saline soils.

This strain is best adapted to big sagebrush and

pinyon-juniper zones, but it also does well in the more

mesic portions of the salt desert shrub area. It appears

to have tolerance to salt similar to that of other

fourwing saltbush ecotypes. Because of its high elevation origin, ‘Rincon’ has shown an adaptability to more

northern climates than might be expected, considering its origin in northern New Mexico. It has performed well at 3,000 to 8,000 ft (910 to 2,400 m)

elevation where the average annual precipitation

ranges from 9 to 23 inches (22.9 to 58.4 cm) (McArthur

and others 1984b). ‘Rincon’ is being grown in seed

orchards for commercial seed production (fig. 2b). The

recommended design and early orchard performance

characteristics of ‘Rincon’ are given by Noller and

others (1984).

‘Marana’ is recommended for Mediterranean type

climates below 4,000 ft (1,200 m) in elevations with

minimum temperatures of 10 ∞F (–10 ∞C) generally in

California and adjacent areas (Carlson 1984; Hassell

1982; McArthur and others 1984b). ‘Wytana’ is closely

related to A. aptera (table 3), and is adapted to eastern

Montana and Wyoming (Carlson 1984; McArthur and

others 1984b).

Several other accessions may have utility for widespread planting although they haven’t been formally

released (McArthur and others 1983a; Petersen and

others 1987; Shaw and others 1984; Young and others

1984e). One particular accession with a proven performance record is the gigas diploid (up to 10 ft [3 m]

tall) that grows naturally only in Utah’s Little Sahara

Sand Dunes (Stutz and others 1975). This giant form

layers readily and appears to have an important place

in the stabilization of dunes or other sandy areas. In

trials since the 1980’s, the giant form has grown well

on clay, as well as sandy soils.

Several other Utah accessions perform well in

pinyon-juniper and big sagebrush communities and

on mine disturbances throughout the Intermountain

region (McArthur and others 1983a). Some selections or ecotypes are nearly evergreen; that is, they

retain a high percentage of their leaves on a yearround basis. Ecotypes that have this feature occur in

both the warmer and cooler portions of the plant’s

natural range. Consequently, the feature is not restricted to limited areas. Among ecotypes considerable differences exist in seed size, utricle thickness,

and germination patterns (Monsen and McArthur

1985; Springfield 1970b). Similar variation also

occurs among seeds collected from a single bush.

However, germination traits are distinct with different ecotypes, and the features can be used to increase

stand establishment. Small utricles normally germinate more rapidly and uniformly than large utricles.

Large, thick utricles must be mechanically fractured to attain uniform germination (Monsen and

McArthur 1985).

Atriplex confertifolia _____________

Shadscale

Shadscale is a compact spinescent shrub, growing typically in dense clumps from 0.5 to 4 ft high

(15.2 to 121.9 cm) and 1 to 5.5 ft (30.5 to 167.6 cm)

wide (fig. 5). The rigid, brittle branches are scruffy

when young but become smooth and spiny with age.

The leaves are nearly circular to elliptic, oval, or

oblong, 9 to 25 mm long, 4 to 20 mm wide. They are

scruffy and gray especially at maturity (Blauer and

others 1976).

Flowers of shadscale are similar to those of fourwing saltbush except in the nature of the bracts

enclosing the seed. The bracts of shadscale are foliose,

5 to 12 mm long, broadly oval to almost round, united

at the base, and have entire, free, somewhat spreading

margins (fig. 5). Shadscale blooms from late March in

the southern portion of its ranges to mid-June in the

northern portion (Hanson 1962). Utricles mature about

15 weeks after blooming. They are fairly persistent

through the winter months (Blauer and others 1976).

Although sex expression in shadscale has not been

studied in the detail accorded fourwing saltbush, it

shares some of the characteristics of that species. For

example, shadscale populations may exhibit biased

sex ratios and include monoecious as well as pistillate

and staminate individuals, and individual plants of

different genders behave different physiologically

(Freeman and Harper 1980; Freeman and McArthur

1982, 1984).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 477

Chapter 20 Chenopod Shrubs

Ecological Relationships and Distribution

Shadscale is widely distributed in the Western

United States from Chihuahua to northern Montana

and from North Dakota and Texas to California and

Oregon (Branson and others 1967; Hall and Clements

1923; Sanderson and others 1990; Stutz and Sanderson

1983). In broadness of distribution among the woody

chenopods it ranks next to fourwing saltbush and

winterfat and frequently grows in association with

both. It grows at elevations from 1,500 to 7,000 ft

(460 to 2,100 m) usually on fine textured alkaline soils

but also on coarser, sandier soils (Blaisdell and

Holmgren 1984; Branson 1966; McArthur and others

1978b).

It occurs in nearly pure stands and in mixtures with

winterfat, budsage, big sagebrush, black sagebrush,

rabbitbrush, spiny hopsage, black greasewood, gray

molly, horsebrush, juniper, other saltbushes, and several species of grass. It endures alkaline soils better

than most of these associates and can be found growing with such halophytes as glasswort (Hall and

Clements 1923). West, Caldwell, and their colleagues,

(Caldwell and others 1977; Moore and others 1972;

West 1985; West and Gasto 1978) showed how the

synecological and physiological relationships of

shadscale and winterfat growing in a mixed stand give

insight on why subtle environmental differences are

important in the distribution patterns of these and

other desert shrubs. A number of ecotypes grow on a

wide range of sites including highly alkaline soils.

Shadscale has a relatively short life span and can be

killed by drought and high water (Blaisdell and

Holmgren 1984; McArthur and others 1978b).

Plant Culture

Although shadscale readily spreads naturally from

seed, it is difficult to establish from seed. Only under

the most favorable climatic conditions do stands

become established from artificial seeding (McArthur

and others 1978b; Monsen and Richardson 1984). It

can be successfully transplanted when moisture

conditions are right in the spring or fall (Luke and

Monsen 1984). Fruit remains on the ground 1 year or

more before germinating and establishing. Understanding the riddles in shadscale germination and

stand establishment requires more research. Warren

and Kay (1984) demonstrated that dormancy in

shadscale is difficult to overcome by traditional mechanical and chemical methods. Young and others

(1984e), following a Stutz (1978) and Stutz and

Sanderson (1983), scenario of shadscale’s evolution in

a pluvial lake regime, suggested that a great deal of

ecotypic variation in shadscale germination should

occur. Oaks, Stutz, and Sanderson (unpublished) have

found ecotypic germination variation to be common.

Garvin and others (1996) using eight shadscale accessions concluded that the species germinates in a longterm temporally staggered fashion, suggesting the

operation of multiple dormancy mechanisms. A few

populations lack the species’ characteristic seed dormancy. Insects can cause severe damage to shadscale

(Haws and others 1984; Hutchings 1952).

Low seed fill often compounds low seed germination

problems. Seed is usually acquired from wildland

stands growing under arid conditions. Seed quality

varies from year to year and even among bushes. Most

commercial lots of seed are collected from upland

foothill ranges that receive higher amounts of precipitation. Harvested seed crops are produced about

every 3 to 5 years, yet some sites are more reliable

than others.

Seed or utricles are normally hand collected but

can be efficiently harvested with vacuum, rill, or striptype machinery (see discussion under fourwing saltbush). Seed usually ripens in late fall or early winter.

Utricles must be dry before they detach from the bush.

Early winter storms can interfere with seed harvesting. Once harvested, the wings are usually removed from the utricle with a hammermill or flail-type

beater. Seeds can be stored for a number of years

without loss of viability. However, unless stored in

cold chambers and treated with an insecticide, seed

borne insects can destroy stored lots. Good quality

seed has between 40 to 50 percent “filled” utricles.

However, a considerable amount of lower quality seed

is sold. Seed viability should be retested if stored over

1 year.

Because seed germination is often erratic and

varies among ecotypes, fall seeding is recommended

to aid in overcoming natural dormancy and breakdown of the utricle. Seed must be placed 0.25 to 0.5

inch (6.4 to 12.7 mm) deep to ensure germination and

seedling establishment. Shadscale is often planted on

heavy-textured salty soils. Soil particles may be flocculated, which creates a rough surface that aids in

seed burial. However, soils may crust easily, and

shadscale seedlings may not be able to emerge

through the surface crust. Shadscale seedlings grow

slowly and are not able to compete with herbs, particularly cheatgrass. Consequently, planting sites

must be cleared of weeds, and the shrub should be

seeded alone in separate rows from other species.

Once seedlings reach 2 years old, losses to competition are low. The plant has done well on barren mine

wastes, particularly infertile soils (Ferguson and

Frischknecht 1985).

Broadcast seeding with no followup measure to

cover the seed has not been very successful. Either

direct drilling or interseeding is recommended. Seed

should be planted at approximately 2 to 4 lb PLS/acre

(2.2 to 4.5 kg PLS/ha). Most conventional seeders can
478 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

be used to seed this shrub. Dewinged seeds are relatively small and flow freely through most seeding

devices. Seeds are relatively heavy and can be used in

seed mixtures to facilitate the planting of more fluffy

or trashy seeds through the seeder (Monsen, unpublished). Seeding rates can be easily regulated. Seed

germination may begin quite early in the spring and

continue until early summer.

Natural spread of shadscale occurs erratically, although large stands frequently develop. New seedlings normally require 3 to 8 years to attain a mature

status. Some ecotypes, however, from north-central

Nevada and northeastern Wyoming grow fast. Undoubtedly other collections have similar traits.

Transplants of shadscale can be successfully reared

as container or bare root stock. Even under favorable

conditions, shadscale seedings grow rather slowly.

The 1 year old stock usually are 3 to 7 inches (7.6 to

17.8 cm) in height but have a moderately diverse

root system.

Seedlings do not survive field planting well unless

extra care is given in handling and planting. Seedlings

must be dormant when planted. Because some leaves

tend to remain on the plant, it is difficult to determine if plants are clearly dormant. Seedlings also

require a growth medium that is similar to the original soil. Bare root seedlings are particularly sensitive

to overwatering. Maintaining plants as 2-0 stock result in much larger materials, but has not improved

field survival. Transplants do not survive if planted

amid herbaceous competition. Clearings of 30 inches

(76.2 cm) in diameter are required to reduce competition sufficiently. Field survival of 60 to 80 percent is

considered normal for large-scale plantings. Much

lower survival results if plants are planted offsite.

Shadscale does not establish or persist long when

planted on unadapted sites. Neither transplants nor

direct-seeded seedlings persist if planted offsite. Care

should be taken to assure seeding of adapted ecotypes.

Unlike fourwing saltbush, shadscale seedlings and

young plants are not selectively sought and grazed by

rodents, rabbits, or livestock.

Uses and Management

Despite its spiny character, shadscale is grazed by

livestock and game. Persistence of leaves varies

greatly over shadscale habitat. Forage value improves

with persistence of leaves. Spines save this species

from heavier grazing than it would otherwise receive,

and the spines are likely a factor in its common

widespread distribution. Seeds are a preferred and

nutritious part of the plant (USDA Forest Service

1937). According to Sampson and Jespersen (1963), as

the leaves and fruits drop in the autumn, they often

accumulate around the parent shrubs. Livestock

readily seek this accumulation. In some areas shadscale

is an increaser and becomes dominant as more palatable species such as budsage and black sagebrush are

killed by grazing (Blaisdell and Holmgren 1984). Because shadscale is generally more salt-tolerant than

fourwing saltbush and several other shrubby chenopods, it may be used to better advantage on the more

salty areas for range rehabilitation and soil stabilization (McArthur and others 1978b). Existing stands of

shadscale should be managed rather than mechanically manipulated because the species usually occurs

on areas that are difficult to manipulate (Bleak and

others 1965; Plummer 1966). This is because shadscale normally grows in semiarid conditions, often occupying sites that receive less than 10 inches (25.4 cm)

annual moisture. When disturbed these areas are

usually invaded by cheatgrass or other weeds. The

competition must be controlled to allow survival of

naturally or artificially seeded shadscale. Large extensive areas can be direct seeded if no competition

exists. Otherwise, some means of site preparation is

required to reduce competition.

Shadscale often occurs as near pure stands void of

most other shrubs. Plants from these pure stands do

not grow in mixtures with many other shrubs. Consequently, they are usually planted alone. Ecotypes that

extend into pinyon-juniper woodlands and big sagebrush communities have been more successfully seeded

in diverse habitats. These ecotypes, the diploid races

(see Varieties and Ecotypes section), when seeded

compete favorably with other species but are best

adapted to sites similar to those from where they have

been collected. Ecotypes collected from more arid salt

desert conditions, the higher polyploids, are less

adapted to foothill upland communities but appear

more drought tolerant.

Varieties and Ecotypes

Shadscale comprise chromosome races ranging

from diploid (2n = 18) to decaploid (2n = 90). The

chromosome races are distributed, in general, with the

high ploidy levels occurring in uniform stands on

valley floors and lower ploidy forms progressively

larger and more morphologically variable and occurring in more diverse communities (Sanderson and

others 1990; Stutz and Sanderson 1983). Little attention has been paid to differential adaptation of these

forms for revegetation purposes. However, the most

robust erect forms produce more seed and have generally been used more for seeding. More attention should

be paid to this feature because some of the higher

polyploids are narrow genetically and adaptively. Consequently, these are poorly adapted to many sites for

which the more diverse diploids and tetraploids are

adapted (Stutz 1982; Stutz and Sanderson 1983).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 479

Chapter 20 Chenopod Shrubs

Atriplex corrugata _______________

Mat Saltbush

Mat saltbush is a low shrub that forms dense,

extremely prostrate (6 inches [15.2 cm] tall) nearly

white mats five to 20 times wider than they are tall

(fig. 6). The prostrate branches often produce adventitious roots where they contact the soil. The bark is

soft, spongy, and white. The evergreen sessile leaves

are opposite on the lower parts of the stems, and

alternate above. The blades are densely scurfy and

measure 7 to 18 mm wide (Blauer and others 1976;

McArthur and others 1978b). Bushes are commonly

dioecious, but some are monoecious. The sexual system appears to operate much like that of fourwing

and shadscale (Freeman and McArthur 1984; Pope

1976). The yellow to light-brown staminate flowers

are borne of glomerules 3 to 6 mm wide on nearly

naked spikes. The pairs of fruiting bracts that enclose the pistils of the female flowers are sessile or

subsessile, 3 to 5 mm long, 4 to 6 mm wide, united

along seven-tenths of their length and usually densely

tuberculate on the lower one-third. Mat saltbush

flowers from April to June, and the fruit ripens 6 to

10 weeks later.

Ecological Relationships and Distribution

Mat saltbush is distributed mainly on soils derived from the Mancos shale formation in eastern

Utah, southwestern Wyoming, western Colorado,

and northwestern New Mexico at elevations from

4,000 to 7,000 ft (1,200 to 2,100 m). It tolerates up to

13,000 ppm soluble salts and is often the only perennial plant present where such high concentrations

of salt occur (Hanson 1962). Mat saltbush is probably

the most halophytic shrub in the genus, but it is

frequently found in areas where the salt concentration

is not so high. In these areas, it may be associated with

shrubs such as winterfat, cuneate saltbush, shadscale,

fourwing saltbush, black greasewood, budsage, and

gray molly. Young plants have been successfully transplanted into a former greasewood type in the Great

Basin (Blauer and others 1976).

Plant Culture

The seed or utricle does not require hammermilling.

The seed is usually screened with a fanning mill to

remove debris, and then seeded with most conventional equipment. Cleaned seed can be stored for

several years without serious loss in viability. Most

native stands produce between 40 to 60 percent filled

utricles.

Seed production is often quite low, yet even during

serious drought and prolonged use, some shrubs produce seed. Seed fill or quality is usually low. Some

stands, particularly those in south-central Wyoming,

produce unusually abundant crops yearly. Large

fields are machine harvested and sold each year.

Hand collection is quite slow, yet field combines are

able to harvest seed easily.

Uses and Management

The plant is important as a winter forage. It often

grows interspaced with other plant communities at

mid to high elevations. Under these circumstances it

is encountered on exposed shaley outcrops with few

other plants. These sites are often windblown slopes

that are accessible to winter grazing. The shrub withstands heavy use, even under unstable soil conditions.

Plants may be heavily browsed and seriously trampled.

Big game animals often concentrate on these exposed

sites during periods of deep snow accumulation. The

shrubs are able to recover from serious abuse without

noticeable loss of vigor.

Fall seeding is normally recommended. Where possible, midwinter planting is successful. Seedlings are

susceptible to early spring frosts. If serious frosts

are expected, spring seeding can be used. Seeds

should be planted near the soil surface or not more

than 0.5 inch (1.3 cm) deep. Seeding is usually done on

sites that once supported the shrub. These areas

usually are not infested with annual weeds, consequently, direct seeding without other preparatory

measures is all that is necessary.

Seedlings usually emerge rapidly and grow rather

vigorously the first year. If moisture remains available, seedlings normally continue growth all summer.

Mat saltbush has been seeded with selected grasses

including Russian wildrye, tall wheatgrass, and crested

wheatgrass. Although these herbs may improve the

Figure 6—Male mat saltbush (left) and

female Castle Valley clover saltbush (right)

growing along highway near Fremont

Junction, Emery County, UT.
480 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

forage production, they usually do not persist in dense

stands throughout the site. Young grass seedlings

and mature plants do suppress shrub establishment.

However, grasses interseeded into established shrub

stands do not decrease shrub density or vigor.

Mat saltbush can be broadcast seeded and covered

with a drag or harrow if planted alone. When seeded

with herbs, the shrub seeds should be planted in

separate rows or spots. Between 2 to 4 lb PLS/acre

(2.2 to 4.5 kg PLS/ha) is recommended. However,

satisfactory stands have developed from lower rates.

When seeding on the heavy clay soils that normally

support this shrub, care must be taken to avoid surface

compaction and crusting.

Transplanting either nursery-grown bare root stock,

wildings, or container materials is successful. Wildings can be dug from native stands and transplanted

directly on disturbances. Wildings and bare root stock

survive abusive treatment, but should not be unnecessarily mistreated. Transplants should not be allowed

to “heat up,” and roots must be placed in a moist soil.

When transplanted on unstable and seemly infertile

soils, the transplants survive remarkably well. Natural enlargement of the crown occurs soon after planting, providing excellent soil protection even under

adverse conditions.

Mat saltbush has performed well from either direct

seeding or transplanting disturbed sites. It is particularly useful as a shrub for seeding mine disturbances, road construction sites, and similar areas

where heavy textured soils and substrata are exposed.

It has established, persisted, and grown well when

planted on pinyon-juniper and big sagebrush sites if

soils are quite basic and heavy.

Small seedlings normally are not destroyed by grazing or insect attack. The shrub should be seeded in

areas where remnant plants remain. Attempting to

convert mat saltbush communities to fourwing saltbush, winterfat, or blackbrush is not advised. Although these and other shrubs may be more palatable, converted communities usually do not remain.

Varieties and Ecotypes

Little study has been accomplished in this area.

Hanson (1962) and Pope (1976) have demonstrated

some populational differences. Mat saltbush will hybridize with other saltbush species (Blauer and others

1976; Stutz 1978, 1984).

Atriplex cuneata ________________

Castle Valley Clover or Cuneate Saltbush

Castle Valley clover saltbush is a low shrub 0.3 to

1.5 ft tall (9.1 to 45.7 cm) with a more or less prostrate,

woody, much-branched base and erect branches

(fig. 6). The light grey-green, spatulate to broadly

elliptic evergreen leaves are 2 to 6 cm long and 0.5 to

2.5 cm wide (Hall and Clements 1923). In common

with some other woody saltbush species, this species is

apparently trioecious (Freeman and McArthur 1984).

Its yellow to brown staminate flowers are borne in

glomerules arranged in panicles. The pistillate flowers are borne in axillary clusters and consist of pistils

enclosed by wingless bracts. At maturity, the bracts

are 5 to 9 mm wide, irregularly toothed along their

margins, and have numerous, flattened, crestlike tubercles on their sides (fig. 4d). Blooming occurs from

mid-April to July, depending on elevation and climatic

conditions. Ripening of the fruit follows about 7 weeks

later (Hanson 1962).

Ecological Relationships and Distribution

The species occurs in high to moderately alkaline

soils in eastern Utah, southwestern Colorado, and

northern New Mexico. In these areas, Castle Valley

clover saltbush is often dominant or is codominant

with shadscale and mat saltbush.

Plant Culture

Seed production of Castle Valley clover saltbush is

similar to that of other woody saltbushes. Annual

yields are erratic, and only about 50 percent of all

utricles produce viable seeds. Seed germination is

also erratic and varies among ecotypes. Seeds or

utricles are relatively small and can be planted with

most equipment. Seedlings are quite vigorous and

grow moderately rapidly. Young plants are competitive but can be suppressed by perennial grasses and

annual weeds. Plants invade open disturbances, yet

success from artificial seeding is not consistent. The

plant should not be seeded directly with herbaceous

plants but is able to compete favorably when seeded

with fourwing saltbush, big sagebrush, green ephedra, and spiny hopsage.

The shrub should not be seeded on areas other

than sites naturally occupied by this shrub. The species is not well adapted to the big sagebrush and

upland communities, but can be used throughout

most salt desert shrublands. Like other woody saltbushes, the shrub does well on fresh disturbances and

can be used to treat exposed substrata created from

mining, road construction, and so forth.

Uses and Management

Castle Valley clover saltbush, so named by stockmen because of the high livestock preference in all

seasons, remains green and succulent throughout

the winter. The shrub shows particular promise in

artificial restoration of certain winter game ranges

and on disturbed sites having saline-alkaline soils.
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Chapter 20 Chenopod Shrubs

It grows through a wide range of salinity and may be

associated with any of the other shrubby chenopods

and many of the sagebrush species. It grows well with

a variety of desert grasses and forbs. Its protein

content helps to provide a balanced diet for livestock

and game, particularly in the winter when much of

the associated vegetation is dry (McArthur and others

1978b).

Varieties and Ecotypes

Castle Valley clover saltbush hybridizes readily

with a number of the saltbushes, but particularly so

with fourwing saltbush, both naturally and artificially

(Blauer and others 1976; Stutz 1984). A stabilized

hybrid of these could be a boon to our winter ranges.

Such a plant would be taller than Castle Valley clover

saltbush and retain more green foliage than fourwing. A stabilized, widely adapted hybrid might be

found in nature.

Atriplex gardneri ________________

Gardner Saltbush

Gardner saltbush is a low subshrub, usually less

than 12 inches (30.5 cm) (fig. 7). The lower one-fourth

to three-fourths of the plant is slightly woody and the

rest is herbaceous. Its habit varies from decumbent

creeping forms to rounded forms three to five times

broader than they are high. The spineless decumbent

branches usually produce adventitious roots where

they contact the soil. The lightly scurfy leaves are

evergreen spatulate to oblaceolate to obovate 15 to

55 mm long, and 5 to 12 mm wide. Plants of this

species are dioecious or sometimes monoecious

(Hanson 1962; Pope 1976). The brown staminate

flowers are borne in glomerules 3 to 5 mm wide on

nearly naked one-branched terminal panicles. The

pistillate flowers are borne on leafy spikes. Fruiting

bracts that enclose the female flowers are 3 to 6 mm

long, 2.5 to 5.0 mm wide, and are sessile. Their

surfaces range from smooth and free of tubercles to

densely tuberculate. The apex of the bract ends in an

oval terminal tooth subtended by two slightly smaller

lateral teeth. Gardner saltbush flowers from midMay to the first of July and intermittently following

heavy rains. The fruit ripens about 7 weeks after

flowering.

Ecological Relationships and Distribution

Gardner saltbush is abundant throughout much of

Wyoming and Montana on clay and occasionally sandy

soils. It is tetraploid, 2n = 36 (Stutz and others 1979).

It is useful year-around forage for big game and livestock (McArthur and others 1978b).

Plant Culture

Field establishment may be difficult because of low

seedling vigor (Ansley and Abernethy 1984a,b). Ansley

and Abernethy (1984b) demonstrated that plants are

best established in a field situation when seeds are

scarified and stratified.

For production fields and commercial harvest,

Carlson and others (1984) recommend direct seeding

on a firm weed-free seedbed with 36 inch (91.4 cm)

rows in the fall at a rate of 20 pure live seed per foot

(66 per m). Once the plants are established and producing seed, the plants can be cut by hayswather,

windrowed, and the utricles collected by conventional

grain combine.

Uses and Management

Gardner saltbush can be successfully seeded on

mine disturbances and rangelands. Although initial

establishment is often erratic, a slow but general

improvement normally results. When seeded in areas

free of weedy competition, seedling survival is usually

adequate. Shrub seedlings grow quite favorably and

are able to survive normal grazing pressure.

Gardner Saltbush Relatives _______

Atriplex buxifolia, Atriplex falcata, Atriplex

tridentata

Much of the literature dealing with Gardner saltbush is confusing in that there are several related

species considered as a species complex. The taxonomic complexity has not been completely unraveled.

The species complex has been alternatively treated

Figure 7—Gardner saltbush (right) growing with

budsage (left) in the Red Desert, Sweetwater

County, WY.
482 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

as the A. gardneri or the A. nuttallii complex. These

suffrutescent shrubs exhibit population variability,

hybridization between populations of the same and

different taxa, and polyploid chromosome races (Stutz

1978; Sanderson and Stutz 1994b). Atriplex nuttalliiis

a name that has recently been lectotypified from the

northern Great Plains (Saskatchewan) by Bassett and

others (1983); that name (A. nuttallii) had been without standing prior to then (Fosberg 1941; Stutz and

others 1979). We believe the plants of the Intermountain area including A. buxifolia, A. falcata (falcate

saltbush), and A. tridentata (tridentate saltbush) are

best referred to as part of the A. gardneri complex

reserving A. nuttallii for plants of the northern Great

Plains. Drs. Howard Stutz and Stewart Sanderson

(n.d.) are actively working on this group of plants from

a cytogenetic and morphological standpoint. They

propose the name of A. canadensis as an alternative

name for A. nuttallii.

Atriplex buxifolia is similar to A. gardneri in habitat

preference (heavy clay soils) and in morphology

(Hanson 1962; McArthur and others 1978b). Stutz

(personal communication) believes that there is no

good reason to maintain the two as separate species

but to refer both to A. gardneri. Plants that have been

referred to as A. buxifolia were larger than A. gardneri

(up to 50 cm; 19.7 inches) and distributed more to the

north and east (northern Nebraska to southern

Saskatchewan) (Hanson 1962; McArthur and others

1978b).

Both A. tridentata and A. falcata are generally

larger statured than A. gardneri. Atriplex tridentata

plants may be from 4 inches to 3 ft tall (10.2 to 91.4 cm).

They root sprout profusely (fig. 8). This is the plant

referred to as A. gardneri by Nord and others (1969)

(see Stutz and others 1979). Atriplex tridentata can

tolerate salt concentrations of over 30,000 ppm and

grows with iodine bush and inland saltgrass

(Hanson 1962). It is distributed mostly in the Lake

Bonneville Basin of western Utah, eastern Nevada,

and southern Idaho, but also in eastern Utah and

southern Wyoming (Hanson 1962; Stutz and others

1979). Atriplex falcata grows up to 20 inches (50.8 cm)

tall. It is distributed through the northern Great

Basin extending northward to eastern Idaho and

eastern Oregon. It grows in soils with 9 to 50 percent

sand often between or in association with shadscale,

Wyoming big sagebrush, and winterfat communities

(Hanson 1962; Yensen and Smith 1984). Atriplex tridentata is mostly hexaploid, but tetraploids are also

known (Stutz 1978; Stutz and others 1979). All the

saltbushes treated in this section are useful yeararound forages as is A. gardneri itself. They may

prove useful in rehabilitation of disturbed saline sites.

The edaphic adaption of each taxa should be borne in

mind in rehabilitation efforts.

Research with A. tridentata and A. falcata have

been hampered by poor seed production. Although

both species are capable of good seed yields, concentrated efforts have not been developed to acquire

adequate samples. Both species have been successfully established by fall seeding. The shrubs appear

capable of growing over a wide range of sites, including areas naturally occupied by fourwing saltbush.

The erect growth habit of A. tridentata and A.

falcata can be grown as bare root or container transplant stock. They survive field planting quite well.

Individual plants grow well under nursery conditions

and may require pruning to reduce plant size.

Atriplex tridentata and A. falcata often lack a herbaceous understory. This is often due to heavy grazing.

Both shrubs persist under intense use, although

seed production is reduced. Both shrubs are capable

of growing with a diverse understory. These shrubs

have the unique ability to persist with various forage

species, and yet remain quite productive themselves.

Other Saltbushes _______________

Atriplex lentiformis (quailbush),

Atriplex obovata (broadscale saltbush),

Atriplex polycarpa (allscale saltbush),

Atriplex hymenelytra (desert holly saltbush)

Some other saltbush species occur in parts of the

Intermountain area—especially in the southern

parts. Quailbush (Atriplex lentiformis) is a large

shrub up to 11.5 ft (3.5 m) tall and 24.5 ft wide (7.5 m)

(fig. 9). Its leaves are evergreen, alternate, gray-green,

scurfy. The distinctive utricles are small flat discs

about 3 to 4 mm in diameter. The similar species,

Figure 8—Example of root sprouting in trident

saltbush from west Sanpete Valley near Ephraim,

Sanpete County, UT.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 483

Chapter 20 Chenopod Shrubs

Torrey saltbush differs by being smaller, up to 10 ft

(3.0 m) tall and 16 ft (4.9 m) wide, and in having anled

stems (versus terete for quailbush). Both species

occur in the St. George area of Utah and in other

southern and western margins of the Intermountain

area (Brown 1956; Hanson 1962). One selection, ‘Casa’,

of quailbush has been released for upland game use

and erosion control (Carlson 1984). In contrast to some

other Atriplex species, quailbush utricles require burial

for effective plant establishment (Young and others

1980).

Three other southern saltbush species are broadscale saltbush, allscale saltbush, and desert holly

saltbush. Broadscale saltbush is a subshrub 0.6 to 1.5

ft (0.15 to 0.45 m) in height with a woody, spreading

base that produces numerous ascending to erect

branches (fig. 10). The silvery, scurfy, deciduous leaves

Figure 9—A large quailbush saltbush near

St. George, Washington County, UT.

are elliptical to obovate, 1 to 3.5 cm long, 1 to 2 cm wide,

with a short petiole. This species has yellow staminate

flowers borne in small glomerules and arranged 500 to

5,000 to a panicle. The fruiting bracts are broadly

cuneate or obovate, 4 to 5 mm long, 5 to 9 mm broad,

with smooth or sometimes slightly tuberculed surfaces (Blauer and others 1976). Broadscale saltbush

tolerates salinity from 165 to 4,900 ppm soluble salts

(Hanson 1962). It is commonly associated with such

halophytes as black greasewood, seepweed, and alkali

sacaton, and other saltbushes (McArthur and others

1978b; Wagner and Aldon 1978).

Broadscale saltbush is an important browse plant

in alkaline areas (Edgar and Springield 1977; Hall

and Clements 1923). It grows rapidly and is most

succulent in the spring. Within areas of adaptation,

this species should be useful for stabilizing disturbed

sites. Frischknecht and Ferguson (1984) reported

that broadscale saltbush was the most successful of

seven chenopod shrubs transplanted into a processed

oil shale matrix in eastern Utah. After 6 years, over

80 percent of the shrubs survived in both irrigated

and nonirrigated plots. Broadscale saltbush seed

germinates best following an after-ripening period

(Edgar and Springfield 1977). It has been observed to

invade the disturbed margins of newly constructed

highways.

Allscale saltbush is an intricately branched, rounded

shrub to 5 ft (1.5 m) tall. Its twigs are delicate, even

approaching spininess. Leaves are obovate to linearly

obovate. It occurs in the Sonoran and Mohave Deserts

and in drier portions of the San Juaquin and Owens

Valleys of California. This species has value as browse

and as upland game habitat (Brown 1956; Chatterton

and others 1971; Hall and Clements 1923; Sankary

and Barbour 1972). Chatterton and others (1971)

reported that allscale saltbush is an excellent nutrient

source for browsing animals especially late in the year.

Graves and others (1975) reported that allscale saltbush was useful for revegetating Mohave Desert land.

For successful reseedings, they suggested that allscale

saltbush be drill seeded in the early winter. Sankary

and Barbour (1972) reported that November is the

time of natural seed dispersal and seedling establishment for allscale saltbush. Mixing activated carbon

directly with utricles in the seedbox enhances germination by inactivating inhibitory substances found in

the utricles (Graves and others 1975).

Desert holly saltbush is a compactly branching,

rounded shrub to 3 ft (91.4 cm) tall. The twigs are

brittle but not spiny. Its leaves are obicular in outline,

but leaf margins are irregularly undulate or coarsely

toothed and as long as 1.6 inches (4 cm). Desert holly

is distributed on dry alkaline alluvial fans and hills

from Owens Valley, California, to southwestern Utah

and south to Mexico (Brown 1956).

Figure 10—Broadscale saltbush growing in

Hamblin Wash, Coconino County, AZ. Ruler is

12 inches (30 cm) long.
484 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

Ceratoides lanata _______________

Winterfat

Winterfat is an erect or spreading subshrub that

shows wide variation in stature from dwarf forms

less than 8 inches (20.3 cm) in height to larger forms

to 4 ft (1.2 m) (fig. 11). The dwarf forms are herbaceous

above a woody base. Taller forms tend to be woody

throughout. Branches and leaves are covered with a

dense coating of stellate and simple hairs that are

white when young but become rust colored with age.

The leaves are alternate, linear, 5 to 50 mm long,

with entire, strongly revolute margins. Winterfat is

predominately monoecious, but dioecious plants also

may occur. The flowers are borne in dense paniculate

clusters along the upper portion of the branches. The

pistillate flowers are below the staminate flowers on

monoecious plants. Staminate flowers lack bracts

and petals. They have four sepals and four stamens

borne opposite the sepals. Pistillate flowers lack both

sepals and petals. Pistils are enclosed by a pair of

bracts that are united more than half their length.

The bracts are covered and often obscured by long,

silky hairs (fig. 4i). These long hairs distinguish winterfat from species of Atriplex. Site and climate permitting, winterfat blooms between May and August.

The fruit ripens from September to November and is

dispersed by wind in late fall and winter. Seed production is extremely variable. A scant crop is produced in most years under arid circumstances.

Sources from foothill ranges usually produce an adequate crop each year. A heavy seed crop can be

produced in years of good summer storms coupled

with the absence of summer grazing. Such productive

crops on dry desert ranges may occur but once in a

decade (Blauer and others 1976).

Ecological Relationships and Distribution

Winterfat is remarkably resistant to drought, and

even on dry sites will produce seed in the third and

fourth years after establishment. On favorable sites,

plants have produced some seed in their first year of

growth. The species has a deep taproot and numerous

extensive lateral roots. The plant may appear dead

after unusually dry years, but it normally recovers

after rain. It exhibits strong reproductive qualities;

however, establishment from artificial seeding is hindered because seedlings are highly sensitive to frost

damage (Stevens and others 1977a).

Winterfat stands often attain old ages. For example,

in southern Idaho a stand of winterfat had plants

thought to be an average of 72 years old, whereas the

corresponding figure for falcate saltbush was 24 years

(Yensen and Smith 1984). Determining the ages of

chenopod shrubs is difficult and imprecise (Stewart

and others 1940; Yensen and Smith 1984).

Winterfat is most abundant on lower foothills,

plains, and valleys with dry subalkaline soils in Utah,

Nevada, Wyoming, Arizona, and New Mexico (Riedl

and others 1964; Stevens and others 1977). In the

Great Basin, it often occurs in pure stands over

thousands of acres (Benson and Darrow 1945) and is

an important component over millions of acres of the

salt desert shrub type. It also grows intermixed with

basin big sagebrush, pinyon-juniper, and ponderosa

pine. The ecotypes for these upland communities

have been used to seed areas often lacking this

shrub. Winterfat ranges from Canada through the

Great Basin and Rocky Mountain States to Mexico,

from California and eastward to Texas and North

Dakota (Branson 1966; fig. 1). It also grows over a wide

range of altitudes. In Utah, it occurs from the lower

Sonoran zone to the alpine ridges. Dwarf forms usually occur on desert floors, on areas of high salt concentration, and on high mountain tops. The larger

forms occur on alluvial fans, foothills, and mesas on

ponderosa pine and juniper-pinyon sites.

Plant Culture

Winterfat can be established in seed production

fields. To do so requires a firm, weed-free seedbed.

Carlson and others (1984) recommended direct seeding at a depth of 0.12 inch (3.2 mm) in 36 inch (91.4 cm)

rows. Fields require cultivation or hoeing and may

require irrigation. Seeds can be combine harvested,

usually from late October to mid-November. See chapter 24 for additional information on seed harvest.

Seed is often hammermilled to remove the fluffy

utricles (Carlson and others 1984). Booth (1982), however, reported better establishment by seeding whole

fruits on the soil surface. Hammermilling also damages many fruits. However, a problem with not

Figure 11—‘Hatch’ winterfat growing at Bell Rapids,

Gooding County, ID.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 485

Chapter 20 Chenopod Shrubs

hammermilling is that fruits may be difficult to

separate from stems, leaves, and litter collected in

the field. Seed should be cleaned using a barley

debearder to detach separate utricles from the flower

head, and then processed with a fanning mill to separate utricles from debris.

Winterfat can be successfully seeded and transplanted in rehabilitation plantings (Clary and

Tiedemann 1984; Frischknecht and Ferguson 1984;

Luke and Monsen 1984; Plummer 1977; Shaw and

Monsen 1984; Stevens and others 1981b). When seeded

alone, winterfat seed does not flow through a drill or

other seeding equipment. If mixed with heavier seed

or a carrier, the winterfat seed is forced through the

machinery. Even when mixed with heavier seeds,

winterfat debris can cause bridging and clogging.

Seed must be carefully cleaned to facilitate seeding.

Sticks and other debris should be removed by screening. Chopping or hammermilling leaves a considerable amount of material that restricts seeding operation. Pelleting the seed has improved seeding without

reducing seed germination. The heavy coated seed can

also be broadcast seeded with less chance of drift.

However, coated or pelleted seed must be properly

cleaned prior to treatment, or sticks and leaves can

still impede seeding.

Winterfat seed maintains viability only for relatively short periods (6 months to 4 years) without

special treatment (Kay and others 1977d; Stevens and

others 1977, 1981a). Special treatments—such as use

of desiccants, sealed containers, and cold storage—

can prolong viability up to 8 years (Kay and others

1977d; Springfield 1974). Winterfat seed requires an

after-ripening period for maximum germination and

germinates best at warm temperatures (77 to 80 ∞F

[25.0 to 26.7 ∞C]) (Springfield 1968, 1972b; Young and

others 1984a). Current research seeks ways to better

establish winterfat by seeding because seedlings are

generally vulnerable to spring frosts (Stevens and

others 1977a).

Winterfat seed is usually cleaned to a purity of

between 20 to 50 percent. Seed viability normally

ranges between 20 to 50 percent. Approximately 1 lb

(PLS) per acre (1.1 k/ha) is recommended for drill

seeding. Seeds can be successfully established by broadcast seeding, if lightly covered with a drag, harrow, or

pipe harrow. Seeds should be covered with not more

than 0.25 inch (6.4 mm) of soil. Some seedlings establish from surface seeding, but failure to cover the seed

is a major reason for poor stands on semiarid sites.

Because spring frosts seriously weaken young seedlings, spring plantings are recommended in areas

where spring frosts are expected.

Winterfat germinates quickly and seedlings grow

rapidly. These traits, coupled with early germination,

cause the plant to be highly competitive with herbs.

Seedlings of few other shrubs survive and grow as well

as winterfat amid competition. However, seedlings

can be weakened by competition, and weedy annuals

and established perennials can and do reduce seeding

survival. Winterfat can be established by broadcast

seeding directly with herbs and other shrubs. When

drill seeded it is best to plant winterfat in separate

furrows from the herbs to reduce competition and to

set the furrower depth for proper seed placement.

Winterfat establishes well when seeded with a cultipacker seeder. Although seeds need only shallow

coverage, planting in a firm seedbed is recommended.

Young plantings are extremely persistent and are able

to withstand heavy grazing, trampling, and drought.

Winterfat is unusually well-adapted to mixed

seedings, particularly when planted in pinyon-juniper

communities. Although desert types normally grow

alone, the tall robust ecotypes do well when seeded in

mixtures. However, to encourage spread, openings

must be created to allow seedling survival.

Winterfat is easily grown as container or bare root

nursery stock. Seedlings are susceptible to damping

off organisms, and entire plantings can be eliminated

in a few days. Maintaining clean, disease-free conditions is the best control measure, although soil drenches

and fumigation can be helpful. Young seedlings transplant well. Plants are difficult to “harden-off” prior to

field planting. Transplants that are not hardened

are susceptible to stress by drought and freezing.

Spring planting is usually recommended, yet transplants are extremely hardy and survive even midsummer planting. The stems of young seedlings become

quite brittle and can be easily broken off during lifting

and transplanting.

Uses and Management

Winterfat is a superior nutritious winter browse for

livestock and big game. Sheep, cattle, antelope, elk,

deer, and rabbits consume it. Except for the woody base

and larger stems, the plant is edible (McArthur and

others 1978b; Riedl and others 1964). Overgrazing has

greatly reduced and even eliminated winterfat in some

areas, even though it is relatively tolerant to grazing

(Stevens and others 1977a). Rasmussen and

Brotherson (1986b) demonstrated that late winter

grazing of winterfat by cattle reduces the vigor of

winterfat stands and opens those stands to less

desirable forage species. Winterfat is potentially one

of the most useful shrubs for planting to increase cover

and forage on alkaline soils of desert ranges in Utah and

adjacent States where the average annual precipitation is less than 10 inches (25.4 cm). It is one of a few

shrubs that can be reliably seeded under arid circumstances. A great deal of winterfat is planted on Western ranges. Plummer (1984) estimated that 20,000 lb
486 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

(9,100 kg) of seed were sold annually. Winterfat ecotypes

from upland communities have proven useful for improving deteriorated pinyon-juniper game winter

ranges.

Sampson and Jespersen (1963) give winterfat a

browse rating of excellent to good for cattle and

sheep, excellent to fair for goats, good to fair for

deer, and fair for horses. Shaw and others (1984)

showed that some winterfat ecotypes have potential as

upland game cover. Ostyina and others (1984) reported that winterfat is useful in increasing range

productivity for fall and winter use of grass pastures.

The species is a good natural increaser and should be

highly useful for stabilization in areas where it naturally occurs. Some good stands have resulted from

broadcasting utricles on such areas (McArthur and

others 1978b).

Varieties and Ecotypes

The different ecotypes of winterfat not only show

wide variation in stature but also in seed production,

seed size, seedling germination and vigor, pubescence

on fruit and seed, and tolerance to varying pH in

soils. These ecotypes maintain character differences

when planted in uniform gardens (Riedl and others

1964; Stevens and others 1977a). A more woody, somewhat spinescent form of winterfat (C. lanata ssp.

subspinosa) referred to as foothills winterfat occurs on

rocky hills in southern Utah, Arizona, California, and

Mexico. It apparently is the only form present in

southern Arizona. This subspecies shows intergradation in all characteristics with typical winterfat

(Kearney and Peebles 1960). Six other species of

Ceratoides are confined to the old world (McArthur

and Sanderson 1984).

Some controversy still exists as to whether the long

standing name of Eurotia retains its validity in Eurasia and in North America (Committee for

Spermatophyta 1978; Howell 1971; Reveal and

Holmgren 1972; Robertson 1982). There is also some

discussion that the proper generic name is

Krascheninnikova. Robertson (1982) suggested that

south-central Asian winterfat, also known as Pamirian

winterfat (Ceratoides latens or Eurotia ceratoides) has

value as a forage plant on North American ranges.

‘Hatch’ winterfat is a recently released selection

that is suitable for its ability to establish, persist, and

provide forage in sagebrush and pinyon-juniper communities (Monsen and others 1985a; Stevens and

Monsen 1988). ‘Hatch’ is a large ecotype capable of

providing winter forage available above the snow

(fig. 11). This selection also has potential for upland

game habitat (Shaw and others 1984).

Grayia brandegei ________________

Spineless Hopsage

Grayia, like Atriplex, has staminate flowers of four

or five sepals, four or five stamens, and pistillate

flowers that lack both sepals and petals but have

pistils enclosed by pairs of bracts. The margins of

each pair of bracts are united from base to apex except

for a minute apical opening. Pendleton and others

(1988) have shown that spineless hopsage has the

unusual monoecious floral system of heterodochogamy. In that system protandrous (“male” first) and

protogynous (“female” first) plants are synchronized

and reciprocal ensuring cross fertilization. On the

basis of a morphological, embryological, and chromatographic study, Collotzi (1966) recommended that

Grayia be reclassified as Atriplex. Welsh (1984) made

an alternate suggestion that G. brandegei be transferred to Zuckia as Z. brandegei var. brandegei. We

favor retaining Grayia as a genus including both the

species G. brandegei and G. spinosa. McArthur (unpublished) analyzed Collotzi’s (1966) chromatographic

data, which shows G. brandegei and G. spinosa to be

most similar among the 10 chenopod taxa. Zuckia was

not included in Collotzi’s (1966) study. More work is

required to resolve the Grayia-Atriplex-Zuckia taxonomic question.

Spineless hopsage is a subshrub to 3.5 ft (1.1 m) tall

(fig. 12) with linear-oblanceolate to obovate leaf blades

1.5 to 4.5 cm long (Collotzi 1966; fig. 13). The flowering

period varies between mid-June and mid-August.

Seed matures in late September through early October.

Utricles persist on the plants until January and some

may last through the winter. These are often removed

by small mammals and birds.

Ecological Relationships and Distribution

Spineless hopsage is generally restricted to shale

formations of the Upper Colorado River Drainage

(Stutz and others 1987) in southwestern Wyoming,

western Colorado, northwestern New Mexico, northeastern Arizona, and eastern Utah. Other populations occur in central Utah. It favors silty clay loam

soil derived from shales. It grows on tough sites that

few other plants occupy (Pendleton 1986; Pendleton

and others 1988). Collotzi (1966) noted it occurred

under mildly alkaline (pH 7.4 to 7.7) conditions.

Plant Culture

Seed viability and production is similar to saltbush

species. Low viability is often encountered due to the

arid conditions at the native sites. It germinates and

produces seedlings readily, but unless protected,

these are quickly taken by rodents and rabbits.
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Chapter 20 Chenopod Shrubs

Uses and Management

This species may be used as browse by livestock

and big game. Small mammals have shown a high

preference for seedlings established at several locations. The shrub should be particularly useful for

stabilizing disturbed shaley soils. It provides forage

for livestock and game, especially in the spring

when it is full leaf (McArthur and others 1978c).

Kingsbury (1964) listed spineless hopsage as possible

secondary or facultative selenium absorber. As such,

it could be mildly poisonous in areas where the soil

contains selenium. Some ecotypes demonstrate the

ability to grow on heavy soils within pinyon-juniper

and basin big sagebrush communities. Stands establish best from fall plantings. To date the shrub has

not grown well with other herbs or shrubs. However,

it is well suited to harsh situations. Seedlings grow

quickly and can be reared as transplant stock quite

easily.

Varieties and Ecotypes

Stutz and others (1987) showed that diploid (2x =

2n = 18) and tetraploid (2n = 36) populations differ

in phenotype and distribution. In general, the diploids are restricted to the more central portion of

the species range, are smaller (x = 13 inches tall and

29 inches wide; 33.0 by 73.7 cm) than tetraploids (x =

16 inches tall and 37 inches wide; 40.6 to 94.0 cm)

and have narrower leaves (x = 3.8 mm versus x =

11.2 mm) than tetraploids (Stutz and others 1987).

Grayia spinosa _________________

Spiny Hopsage

Spiny hopsage differs markedly from spineless

hopsage in many respects, but the fruits and flowers

Figure 13—Spineless hopsage growing in

Antelope Valley, Sanpete County, UT.

are similar. It has small, inconspicuous, unisexual

flowers. The sexual system appears to be temporally

monoecious to allow the species to more efficiently

occupy stressful, heterogeneous, patchy environments

(McArthur and Sanderson 1984). Spiny hopsage is an

erect, diffusely branched, spinescent shrub from 1 ft

(30.5 cm) to 4 ft (121.9 cm) high. Its leaves are fleshy

in the spring but usually dry before midsummer.

Fruits (fig. 4k) generally mature just before leaf fall

(Ackerman and others 1980). Unusual ecotypes in

northeastern Utah and southwestern Idaho retain a

considerable percentage of their leaves throughout

the growing season (Blauer and others 1976).

Ecological Relationships and Distribution

Spiny hopsage occupies plains, desert valleys, and

foothills over much of the Western United States

(eastern Washington south to southern California

and east to western Wyoming and northwestern

New Mexico) at elevations ranging from 2,500 to

6,900 ft (760 to 2,100 m) (Blauer and others 1976;

Collotzi 1966; Shaw 1992). It generally grows on welldrained soils of loamy to rocky or gravelly textures,

but it may occasionally be found on clay soils. Soils

usually range from neutral to alkaline; its adaptation

to salinity may vary ecotypically (McArthur and others 1978b; Shaw1992; Stark 1966). It often grows in

soils high in calcium (McArthur and others 1978b).

Spiny hopsage may occur in pure stands but more

often is a component of communities dominated by

other shrub species (Shaw 1992). It occurs in both

sagebrush and salt desert shrub communities in association with Wyoming and basin big sagebrush,

blackbrush, black greasewood, shadscale, winterfat,

and pinyon and juniper species (Sampson and

Jespersen 1963; Welsh 1984).

Figure 12—Spineless hopsage growing in

Antelope Valley, Sanpete County, UT.
488 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

Figure 14—Gray molly growing near Faust,

Tooele County, UT.

Plant Culture

Seed production is highly variable between years,

populations, and individual plants. During favorable

moisture years some shrubs produce large quantities of fruits, but during dry years few flowers or

fruits are produced (Shaw 1992). Under warehouse

conditions seed maintains viability up to 6 years

(Shaw 1992; Smith 1974a). Kay and others (1984)

demonstrated that seed life can be extended by drying

seed at 35 ∞C (95.0 ∞F) for 6 days and storing them in

glass jars with a desiccant at room temperature. This

procedure has maintained full seed viability for over 9

years.

Viable seeds germinate rapidly. For field establishment late fall seeding is best because moisture

levels are high (Shaw and others 1994). Bracts of

spiny hopsage are probably important for seed dispersal and seedling establishment (Shaw and others

1996). In some cases, however, removal of the bracts

may be preferred for seeding depending on the equipment used. Most drills are unable to seed the large

fruits unless seeded alone.

Young seedlings grow quite rapidly but become

dominant during the early summer. The plant often

grows without an understory of herbs, and the shrub

seedlings are not highly competitive with herbs. To be

successful, seedlings must be made in prepared seedbeds free of competition. Small seedlings are vulnerable to weedy annuals.

Broadcast seedings have erratic success. Natural

seeding is episodic; success depends on the availability

of soil water (Shaw and others 1994). When hand

seeded in pits or favorable sites and covered with soil,

moderate success has been achieved. Rodents appear

to forage upon young seedlings, but the extent of their

damage is not fully understood. When seeded with

herbs, small seedlings emerge and survive until about

midsummer. Seedlings then appear to become dormant, yet few recover the following season.

Field grown transplants also demonstrate early summer dormancy even when irrigated. However, greenhouse reared stock do not senesce prematurely. Field

survival of bare root transplants have been satisfactory when planted on mine disturbances and native

sites. Neither container or bare root stock do well

when planted offsite.

Uses and Management

Spiny hopsage is a valuable forage for livestock and

game from early May to midsummer while foliage

persists. It receives little use during its dormant

period (McArthur and others 1978b; Sampson and

Jespersen 1963; Welsh 1984). Spiny hopsage furnishes good cover for upland game in all seasons. The

plant has potential for use in revegetation but has

received little use in that capacity. Glazebrook (1941),

Wood and others (1976), and Kay and others (1977c)

have all recommended its use for rehabilitation

plantings. However, additional work on site requirements is needed to better assure artificial seeding

success.

Varieties and Ecotypes

No improved varieties have been developed. Spiny

hopsage is, however, composed of numerous ecotypes.

Use of these for various management purposes and

genetic improvement should both be possible. The

species is known only at the tetraploid (4x) level

(Sanderson and Stutz 1994b).

Kochia americana _______________

Gray Molly

Gray molly is a small, woody-based subshrub to 20

inches tall (50.8 cm) (fig. 14). Its numerous, annual,

erect branches and leaves are covered with long, silky

hairs. The linear, fleshy, more or less terete leaves

are 5 to 30 mm long. Hairy perfect or pistillate flowers

are borne singly or in small clusters in the axils of

leaves along almost the full length of the branches.

At anthesis, the hooded calyx lobes are about 1.5 mm

long and closely cover the ovary. At maturity, the fruit

is largely concealed in the persistent calyx, which

develops conspicuous, horizontal, fanlike papery

wings to 3 mm long. This species blooms from June to

August (Blauer and others 1976).

Gray molly usually occurs in saline or alkaline clay

on plains and foothills between 4,500 to 6,000 ft

(1,400 to 1,800 m). Its range extends from southern

Montana and eastern Oregon to New Mexico and

California. Gray molly summer cypress is quite widely
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Chapter 20 Chenopod Shrubs

used as winter forage by livestock when it is sufficiently abundant (Blauer and others 1976). It appears

to have usefulness in revegetation of areas with a high

salt concentration.

Varieties and Ecotypes

Both diploid (2x) and tetraploid (4x) populations

are known to exist but their characteristic differences, if any, are poorly understood (Sanderson and

McArthur, n.d.; Sanderson and Stutz 1994b).

Kochia prostrata ________________

Forage Kochia

Forage kochia is generally a long-lived, highly variable, woody-based subshrub (fig. 15). However, some

ecotypes may be definite upright shrubs. It ranges

from less than 12 inches to over 40 inches (30.5 to

101.6 cm) in height. Ascending branches are covered

with short to long woolly hairs. Leaves are flat, linear

to filiform, and hairy. Flowers are borne in small

clusters (glomerules) in the axils of slightly reduced

leaves on the upper part of the stem. As the fruit

develops, the calyx forms dorsal appendages around

it that are rounded, flat, and tuberclelike, or oblong

and winglike (Shishkin 1936). Blooming occurs from

July to September (Blauer and others 1976).

Ecological Relationships and Distribution

Forage kochia is native to arid and semiarid regions

of central Eurasia extending to the Mediterranean

basin and northeastern China. It grows on alkaline,

stony, and sandy steppes and plains (Francois 1976;

Shishkin 1936). Forage kochia tends to be evergreen.

Seeds of accessions from several types including both

subspecies of forage kochia were introduced into the

United States during the 1960’s (Keller and Bleak

1974). The species is well adapted to the climate and

soils of the Intermountain area. This is especially so

for the pinyon-juniper, sagebrush, and salt desert

shrub communities (Blauer and others 1976; Keller

and Bleak 1974; McArthur and others 1974; Stevens

and others 1985a). It grows at elevations in its native

range from 0 to 8,000 ft (0 to 2,400 m) (Balyan 1972;

McArthur and others 1974).

Plant Culture

Seed, under uncontrolled storage conditions, loses

viability rapidly in about 6 months (Balyan 1972;

Jorgensen and Davis 1984). Seed viability can be maintained up to 4 years by drying seed to 7 to 9 percent

moisture levels and storing in airtight containers

(Balyan 1972; Jorgensen and Davis 1984; Young and

others 1981c).

In seeding forage kochia, best results occur when

the seed is surface sown and not covered (Stevens

and Van Epps 1984). Aerial seedings have been successful. Seeding at a rate of 0.25 lb (PLS) per acre

(0.28 k/ha) is usually sufficient. If drilled, seeds should

be no deeper than 1⁄16 inch (1.6 mm) (Stevens and

others 1985a). The species can be transplanted with

good results in spring or fall when the ground is moist.

Once established forage kochia is a good natural

spreader. In concentrated stands, forage kochia attracts large populations of lygus bugs, but such concentrations do not occur in mixed plant communities

(Moore and others 1982).

Uses and Management

In its native habitat, forage kochia is a droughtresistant, salt-tolerant species and is highly valued as

a forage plant (Nechaeva and others 1977). The plant

has high nutritive value with a high protein and

carotene content (Balyan 1972; Davis 1979; Welch and

Davis 1984). Forage kochia has its highest nutritive

content in midsummer through fall when its crude

protein and carotene levels are high (Davis 1979;

Davis and Welch 1984). Frischknecht and Ferguson

(1979), McKell and others (1979), and Ferguson and

Frischknecht (1985) have shown forage kochia to be

an effective mine spoil stabilizing plant. Sheep, goats,

camels, and horses all consume it (Shishkin 1936). In

Utah this species is sought out by mule deer. Davis

and Welch (1985) showed some accessions are more

palatable to mule deer than others. Forage kochia

readily establishes from seed and grows rapidly, and

under favorable conditions, reaches sexual maturity

in 1 year (McArthur and others 1974). It spreads

well, naturally, from seed. It is a useful component in

shrub-grass pastures (Otsynia and others 1984). Once

Figure 15—Forage kochia growing on a

roadcut near the mouth of Salina Canyon,

Sevier County, UT.
490 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 20 Chenopod Shrubs

Figure 16—Black greasewood growing near Tuba

City, Coconino County, AZ.

established, forage kochia will compete with annuals

such as halogeton and cheatgrass (McArthur and

others 1990a; Monsen and Turnipseed 1990; Stevens

and McArthur 1990; Stevens and others 1985a). One

desirable characteristic for competing with cheatgrass is its fire tolerance (McArthur and others 1990a).

Forage kochia shows considerable potential for

becoming a valuable forage and cover plant on arid

Western ranges. In arid shrublands, it is among the

most desired species to seed. Concern over possible

poisonous properties is unnecessary (Davis 1979;

Williams 1980). Russian workers have been seeding

forage kochia on ranges since 1921 (Balyan 1972).

Ecotypes and Varieties

Although forage kochia is not native to North

America, it is well adapted to many areas in Western

North America (McArthur and Stevens 1983; Stevens

and others 1985a). A near relative, Kochia americana,

is widespread but less robust and aggressive than

K. prostrata (Blackwell and others 1978; Blauer and

others 1976).

Forage kochia comprises two subspecies and two

varieties with separate habitat preferences (Balyan

1972):

Clay habitats; Kochia prostrata ssp. virescens

(green)

Sandy habitats; Kochia prostrata ssp. grisea (grey) var.

canescens (densely

pubescent)

Solonetzes (salty clay soils) var. villosocansa

(white-villous)

Each of the above taxa has a relatively wide tolerance

to soil types; however, in general, the green subspecies (ssp. virescens) grows best in clay soils. It has

both mountain and lowland ecotypes (Balyan 1972).

Variety canescens of the grey subspecies (ssp. grisea)

is adapted best to sandy habitats, whereas variety

villosocansa is best adapted to salty clay soils (Balyan

1972).

The species is a polyploid complex based on x = 9

with diploid, tetraploid, and hexaploid populations

(Davis and Welch 1985; Herbel and others 1981; Pope

and McArthur 1977). One selection, ‘Immigrant’, has

been released (Stevens and others 1985a). This

selection (ssp. virescens), originally from Stavrapol

Botanical Gardens in Russia, has shown superior

performance in longevity, forage production and quality, palatability, and competitiveness with annuals.

It is adapted to sandy loam to heavy clay soils with best

adaptation to heavier soils. ‘Immigrant’ has demonstrated its adaptability to the pinyon-juniper, basin

and Wyoming big sagebrush, and greasewoodshadscale shrub vegetative types. McArthur and

others (1996) recently confirmed the outstanding

performance of ‘Immigrant’ forage kochia in an adaptation and performance test across an environmental

gradient. Another accession (ssp. grisea) also performed well.

Sarcobatus vermiculatus _________

Black Greasewood

Black greasewood is an erect, spiny-branched shrub

up to 10 ft (3.0 m) tall (fig. 16). The deciduous, brightgreen leaves 1 to 4 cm long are usually narrowly linear

and often semi-terete. The shrub is usually monoecious.

Black greasewood had traditionally been considered as a member of the chenopod family. However,

recent studies on chloroplast DNA and sieve element

plastids provide evidence that it should be separated

from that family (Behnke 1997; Downie and others

1997). Behnke (1997) reviews the taxonomic history

of Sarcobatus and concludes that this genus is distinct enough to merit its own family, Sarobataceae.

We have included it here because of its traditional

place in the Chenopodiaceae.

Ecological Relationship and Distribution

Black greasewood grows from Alberta south into

Mexico with stands occurring in all the Western States.

A smaller species, Bailey greasewood (Sarcobatus

baileyi), occurs in western Nevada in association

with shadscale and other desert shrubs. Shadscale

often grows with black greasewood as well. In the

more saline areas of its range, black greasewood

may be found in nearly pure stands. On such sites,

minor components of shadscale, Castle Valley clover

saltbush, and Gardner saltbush may occur. In less

saline areas, it may grow with threadleaf and whitestem
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Chapter 20 Chenopod Shrubs

rubber rabbitbrushes, spiny hopsage, basin big sagebrush, fourwing saltbush, winterfat, black sagebrush,

and broom snakeweed. Greasewood often occurs in

valley bottoms where subsoil moisture is high. Often

inland saltgrass, alkali sacaton, Nuttall alkaligrass,

alkali muhly, and basin wildrye grow with black

greasewood, they may form grassy understories. Russian wildrye, tall wheatgrass, and quackgrass have

been successfully planted into black greasewood stands

to make these stands more versatile grazing areas

(McArthur and others 1978b).

Plant Culture

Like other chenopod shrubs, black greasewood produces erratic seed crops that usually have low viability. Seed is more difficult to collect because of the

spiny, brittle nature of the stems. Poor seed quality

has decreased planting success. Seeds ripen in the

fall and respond best to fall or winter seeding. Establishment by natural or artificial seeding is unpredictable. Seeding with grasses is not recommended

unless the shrub can be seeded in separate rows.

Black greasewood often exists on sites with claytextured surface soils. Soils compact and crust easily,

often reducing seedling survival. Black greasewood

has done well when seeded or transplanted on mine

disturbances. Under these conditions it establishes

well on fresh exposures and infertile soils. It competes

well with herbs under these circumstances unless the

sites are fertilized heavily.

Uses and Management

Black greasewood contains soluble oxalates that

will sicken and kill hungry livestock that consume

large amounts of the foliage (Kingsbury 1964). However, where there is a good assortment of other species,

this danger is negligible. It is often eaten by livestock

(sheep and cattle) and wildlife in the spring. It is

usually not regarded as an important forage plant

although it furnishes cover and habitat. It is especially important on sites that flood, and it forms an

important zone within various riparian habitats. It is

highly important as cover to upland game birds and

big game animals. Sites that are burned, disked, or

plowed and seeded to herbs receive heavy use. When

seeded on mine and related disturbances, the shrub

is often heavily grazed.

Varieties and Ecotypes

Black greasewood exhibits considerable variation

both in natural stands and plantations. Roos (1984),

in a study involving 18 populations, concluded that

Sarcobatus vermiculatus was rich in genetic variation. Sanderson and others (1999) have documented

an interesting distribution of chromosome races in

Sarcobatus. Sarcobatus vermiculatus, black greasewood, is widely distributed in Western North America

(fig. 1). Tetraploid (x = 9, 4x populations) occur only on

the periphery of the distributional range to the north

(southern Alberta, southern Manitoba, northern Montana, eastern North Dakota), to the west (northeastern and east-central California and south-central Oregon), and to the south (southern Arizona and

northwestern Sonora). Throughout the rest of the

species vast range including the central portion of that

range, S. vermiculatus is octoploid (8x). It is apparently an old plant; no diploids are known (Sanderson

and Stutz 1994b; Sanderson and others 1999). Sarcobatus baileyi is dodecapoid (12x) throughout its more

limited range (western and south-central Nevada)

(Sanderson and Stutz 1994b; Sanderson and others

1999).

Other Chenopod Shrubs _________

Allenrolfea occidentalis (iodine bush),

Camphorosoma monspeliaca

(Mediterranean camphorfume), Suaeda

torreyana (desert sumpbush or desert

blite)

Iodine bush and desert sumpbush are two species

that grow in areas where salt concentrations are so

high that they are the only forms of higher plant life

that survive (McArthur and others 1978b). In so

doing, they provide a valuable soil-binding service.

Perhaps these species could be used more for rehabilitation work. Another species, an introduction

from Eurasia, Mediterranean camphorfume, has

been shown to be useful in rehabilitating mine spoil

materials (Ferguson and Frischknecht 1985). This

species is similar in growth habit to forage kochia,

and like forage kochia, it is a useful forage plant in

south-central Asia (Nechaeva and others 1977).
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Chapter 20 Chenopod Shrubs
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 493

E. Durant McArthur

Richard Stevens

21

Chapter

The sunflower family (Compositae or Asteraceae) is the largest

family of flowering plants. Its many species occur around the

world as annual and perennial herbs and as shrubs and trees

(Benson 1957; Cronquist 1968; Wagenitz 1977). Three shrubby

genera of the family—sagebrush (Artemisia), rabbitbrush

(Chrysothamnus), and matchbrush (Gutierrezia)—make plants

of this family among the most common and important plants of

the Intermountain area (McArthur and others 1979a; table 1).

Shrubs of these genera provide critically needed ground cover on

arid Western ranges, are important sources of browse for domestic livestock and big game, and serve as cover and forage for many

wildlife species. A number of sagebrush and rabbitbrush species

are important as cover for small birds, game birds, and mammals, and as browse plants for big game animals, especially on

winter and early spring ranges. Some species also provide forage

for livestock (sheep and cattle). Horsebrush and matchbrush also

contribute more forage than is generally believed; however, both

plants may, under certain conditions, be harmful to domestic

livestock (Benson and Darrow 1945; Johnson 1974a; McArthur

and others 1979a) and cause allergies in humans (Lewis and

Elvin-Lewis 1977; Rodriguez and others 1976).

Composite Shrubs
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Chapter 21 Composite Shrubs

Various forms of sagebrush and rabbitbrush may

also be directly seeded or transplanted for landscaping,

stabilizing, and beautifying disturbed landscapes and

may have potential for supplying industrial chemicals. Young wildings of both sagebrush and rabbitbrush transplant easily. Usually within 3 to 7 years,

direct seeded or transplanted plants are established

sufficiently to reproduce naturally from seed. Both

establish well when properly aerially or drill seeded

(McArthur and others 1974; Plummer 1977; Plummer

and others 1968).

In this chapter, we prefer to limit the term sagebrush to members of the subgenus Tridentatae

(McArthur 1979; McArthur and others 1981). Other

woody members of the genus Artemisia then become

“sage” or “wormwood.” However, in the broad sense in

this chapter, use of sagebrush alone may refer to any

woody, native North American Artemisia species.

Sagebrush and rabbitbrush species that are especially important on low elevation Intermountain area

Chapter Contents by Species

General sagebrush culture ................................................................................................................. 496

Artemisia abrotanum (oldman wormwood) ....................................................................................... 497

Artemisia arbuscula (low sagebrush ................................................................................................. 499

Artemisia bigelovii (Bigelow sagebrush) ........................................................................................... 500

Artemisia cana (silver sagebrush) ..................................................................................................... 502

Artemisia filifolia (sandsage or oldman sage) .................................................................................. 504

Artemisia frigida (fringed sage) ......................................................................................................... 505

Artemisia longiloba (alkali sagebrush).............................................................................................. 506

Artemisia nova (black sagebrush)...................................................................................................... 508

Artemisia pygmaea (pygmy sagebrush)............................................................................................. 509

Artemisia rigida (stiff or scabland sagebrush).................................................................................. 510

Artemisia spinescens (budsage).......................................................................................................... 511

Artemisia tridentata (big sagebrush) ................................................................................................. 513

Artemisia tripartita (threetip sagebrush) ......................................................................................... 521

Other sagebrushes .............................................................................................................................. 522

General rabbitbrush culture .............................................................................................................. 522

Chrysothamnus depressus (dwarf rabbitbrush) ................................................................................ 524

Chrysothamnus linifolius (spreading rabbitbrush) ............................................................................ 524

Chrysothamnus nauseosus (rubber rabbitbrush).............................................................................. 525

Chrysothamnus parryi (Parry rabbitbrush) ...................................................................................... 529

Chrysothamnus vaseyi (Vasey rabbitbrush) ..................................................................................... 531

Chrysothamnus viscidiflorus (low rabbitbrush) ............................................................................... 531

Other rabbitbrushes ........................................................................................................................... 533

Other composite shrubs...................................................................................................................... 534

Haplopappus species (goldenweed) ............................................................................................. 534

Tetradymia species (horsebrush) ................................................................................................ 534

Gutierrezia species (matchbrush or snakeweed) ........................................................................ 534

Lepidospartum latisquamatum (scalebroom) ............................................................................. 534

ranges include big sagebrush (A. tridentata), black

sagebrush (A. nova), threetip sagebrush (A. tripartita),

low sagebrush (A. arbuscula), budsage (A. spinescens),

fringed sage (A. frigida), sand sage (A. filifolia),

early sagebrush (A. longiloba), rubber rabbitbrush

(C. nauseosus), low rabbitbrush (C. viscidiflorus),

Greene’s rabbitbrush (C. greenei), spreading rabbitbrush (C. linifolius), and Parry’s rabbitbrush (C. parryi).

Some of these species, for example, threetip sagebrush

and Parry’s rabbitbrush, are usually found on ranges

at higher elevations than the others.

Virtually all of these species include an array of

ecotypes. Some are large taxonomic units that include

several subspecies (big sagebrush, rubber rabbitbrush,

low rabbitbrush, and Parry’s rabbitbrush) (McArthur

and others 1979a). These shrubs, especially under

conditions of heavy grazing, may form closed stands.

However, in many locations they mix with grasses and

forbs. Big sagebrush often occurs with the grasses,

bluebunch wheatgrass, western wheatgrass, the
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Chapter 21 Composite Shrubs

Table 1—Important Composite shrubs of the Intermountain

area.

Common name Scientific name

Oldman wormwooda Artemisia abrotanum

Low sagebrush Artemisia arbuscula

Bigelow sagebrush Artemisia bigelovii

Silver sagebrush Artemisia cana

Sand or oldman sage Artemisia filifolia

Fringed sage Artemisia frigida

Longleaf sage Artemisia longifolia

Alkali sagebrush Artemisia longiloba

Black sagebrush Artemisia nova

Birdsfoot sage Artemisia pedifida

Pygmy sagebrush Artemisia pygmaea

Stiff or scabland sagebrush Artemisia rigida

Budsage Artemisia spinescens

Big sagebrush Artemisia tridentata

Threetip sagebrush Artemisia tripartita

Alkali rabbitbrush Chrysothamnus albidus

Dwarf rabbitbrush Chrysothamnus depressus

Spreading rabbitbrush Chrysothamnus linifolius

Greene’s rabbitbrush Chrysothamnus greenei

Rubber rabbitbrush Chrysothamnus nauseosus

Parry rabbitbrush Chrysothamnus parryi

Vasey rabbitbrush Chrysothamnus vaseyi

Low rabbitbrush Chrysothamnus viscidiflorus

Small headed matchbrush Gutierrizia microcephala

Broom snakeweed Gutierrizia sarothrae

Gray horsebrush Tetradymia canescens

Littleleaf horsebrush Tetradymia glabrata

Nuttall horsebrush Tetradymia nuttallii

Spiny horsebrush Tetradymia spinosa

a

Introduced species.

introduced crested wheatgrasses and desert wheatgrasses, and intermediate wheatgrasses, Idaho fescue, sheep fescue, Indian ricegrass, Sandberg bluegrass, Thurber needlegrass, bottlebrush squirreltail,

and others.

A rich array of forbs also grow in the sagebrushgrasslands, including species of aster, lupine, locoweed

or milkvetch , balsamroot, Agroseris, wild buckwheat,

and goldenrod. Sagebrush species are the dominant

and codominant species of over 40 described habitat

types of the American West (Blaisdell and others

1982). Passey and others (1982) published a classic

study on sagebrush habitats of the Intermountain

West. Their statement of the importance of sagebrush:

Opinions differ among plant scientists as to the place

of sagebrush in plant communities on semiarid rangelands (Ellison 1960). Historical accounts of the amount

and distribution of sagebrush are also often contradictory. As noted, the relative amount and distribution of sagebrush species or subspecies have been

observed and evaluated on the several hundred tracts

examined in this study. One or more species of sagebrush were present on every area examined except

where there was unmistakable evidence of recent

burning, tillage, or mechanical or chemical treatment.

Many of these observations were made on rangelands

where the plant community was judged to be in nearclimax condition. Some areas, in fact, had no evidence

of use or disturbance of any kind except by native

fauna. On all study sites supporting subspecies of big

sagebrush, annual growth rings of the largest stems

were counted to determine the age of the plants and

to establish the length of time during which there had

been no fires. The oldest plants found were about 120

years old, and several locations supported plants at

least 100 years old. Many of the older plants had split

and nearly decayed stems on which the growth rings

could not be accurately counted; so some plants may

have been even older. With few exceptions, the oldest

(not necessarily the largest) plants grew on stony or

gravelly soils, and were of the Wyoming big sagebrush

subspecies. On disturbed relic areas, subspecies of big

sagebrush typically grew in open stands with individual plants or small clumps of plants uniformly

distributed. Where it grew, big sagebrush made up five

to 20 percent of the total annual production on the

study site.

Woody Artemisia, called wormwood in North Africa

and Eurasia, are principal components of the steppes

of those regions. There (Larin 1956), as in North

America (Blaisdell and others 1982; Laycock 1979;

McArthur and Plummer 1978), many range managers

regard them as a mixed blessing. They are aggressive

and increase on overgrazed land; on the other hand,

they provide much forage for wildlife and domestic

animals. This group of plants (Artemisia) may number 400 species and is divided into four subgenera

based on floral, anatomical, chemical, karyotypical,

and distributional characteristics (McArthur 1979;

McArthur 1983b; McArthur and others 1981; table 2).

Rabbitbrushes are equally as widespread as sagebrush (= N. American Tridentatae) but are generally

less dominant (McArthur and Meyer 1987). There are

17 rabbitbrush species with 40 subspecies divided

into five sections (table 4).

Seed of most composite shrubs can be collected by

hand stripping, beating into a container, and, in some

cases, with vacuum harvesters (chapter 24). For seed

to pass through most seeders, plumes and other flower

parts need to be partially removed. This can be done by

passing the seed through a barley debearder followed

by screening and fanning. Seeds are rather brittle;

consequently, care must be taken to ensure that they

are not cracked or broken during cleaning and seeding. Seed of most composites can be stored no more

than 1 to 3 years before seeding—otherwise, considerable viability will be lost (Stevens and others 1981a).

A principal trait of sagebrush and rabbitbrush is the

ability to persist under heavy abuse and poor management, and yet, be able to recover with proper management. Few other shrubs can recover in a reasonable

period without special management, including seeding. Most plants of this group can compete with weedy

herbs, and many species occupy arid sites where few

other shrubs persist.
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Chapter 21 Composite Shrubs

Important characteristics of a number of composite

shrubs are listed in the Introduction to section VII.

Seeding recommendations for principal vegetative

types and conditions are discussed in chapter 17.

Composite shrubs adapted to these vegetative types

and conditions are included in the seeding recommendations. Seed characteristics are found in chapters 24 and 26.

We include in this chapter a brief species description, ecological relationships, distribution, culture

requirements, use, improved varieties, and management of each of the species appearing in the chapter

contents list.

General Sagebrush Culture _______

Intermountain sagebrush species all have small

seed (see chapter 24). The smallest seeds are those

of fringed sage, with over 4.5 million seeds per lb

(9.9 million per kg) (100 percent purity). Pygmy sagebrush has relatively large seeds with about 500,000

Table 2—Taxonomic subgenera of Artemisia (McArthur and others 1979a).

Classical Modern Distinguishing Species mentioned

sectiona subgenerab characteristics Distribution in this chapter

Absinthium Artemisia Pistillate ray flowers Eurasia A. abrotanum

perfect disc flower, North Africa A. absinthium

predominantly North America A. frigida

herbaceous, but

Abrotanum a few are woody A. longifolia

A. ludoviciana

Drancunculus Drancunculus Pistillate ray flowers Eurasia A. filifolia

staminate disc flowers North America A. pedatifida

herbacious and woody. A. spinescens

Seriphidium Ray flowers lacking, Eurasia,

Seriphidium perfect disc flowers; North Africad

herbaceous and woody.

Tridentatae Ray flowering lacking, North America A. arbuscula

perfect disc flowers, A. argillosa

woody.c A. bigelovii

A. cana

A. longiloba

A. nova

A. pygmaea

A. rigida

A. rothrockii

A. tridentata

A. tripartita

a

De Candolle 1837; Hooker 1840; McArthur and Plummer 1978; McArthur 1979.

b

Rydberg 1916; Beetle 1960; Polyakova 1961; McArthur 1979; McArthur and others 1981.

c

The single exception, A. bigelovii, has zero to two pistillate ray flowers on otherwise discoid heads. d

Two anomalous American species have been referred to Seriphidium: A. palmeri of southern and Baja California and

A. mendozana Argentina.

per lb (1.1 million per kg). In big sagebrush, seed size

varies among subspecies. Basin big sagebrush has

just over 2.5 million seeds per lb (5.5 million per kg),

Wyoming big sagebrush just under 2.5 million

(5.5 million per kg), and mountain big sagebrush has

about 2 million (2.2 million/kg). Seed size, no doubt,

influences the depth at which seeds should be planted.

Seed of most sagebrush species should not be planted

deeper than 1⁄16 inch (1.6 mm). Good results can be

obtained when seeds are broadcast on top of disturbed

soils and not covered.

Adequate covering usually occurs from natural soil

sluff. Sagebrush seed can be placed on top of the

seedbed using drills that have had their seed drops or

tubes pulled out between the disk-furrow opener and

placed behind. Seeds then drop on the soil disturbed

by the disk furrow openers. Several effective methods

of planting sagebrush seed include broadcasting (aerial

or by ground rigs), and placing through a seed dribbler, thimble seeder, or browse seeder onto disturbed

soil. Drill seeding is a low priority option.
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Chapter 21 Composite Shrubs

Figure 1—Oldman wormwood plants growing

from stem cuttings along Ephraim Canyon Road,

Sanpete County, UT.

Artemisia abrotanum ____________

Oldman Wormwood

Oldman wormwood, also called southernwood, is a

shrub 1.5 to 6 ft (0.5 to 1.8 m) tall (Grieve 1931; Hall

and Clements 1923; Plummer 1974b). Stems are much

branched, erect or somewhat spreading, and form a

rounded bush (fig. 1). The species has a pleasing

aromatic fragrance and bright green, finely divided,

pinnately dissected leaves giving the shrubs a feathery appearance. It has a deep and extensive root

system allowing the plant to obtain nutrients and

water from considerable distances (Plummer 1974b).

This characteristic also allows the plant to establish

in raw subsoils. Principal leaves are 3.0 to 6.1 cm long,

two or three times pinnately dissected with revolute

margins, glabrous or sparsely puberulent on upper

surfaces, but lightly tomentulose on the lower surfaces (Hall and Clements 1923). The inflorescence is

an elongated terminal panicle, leafy at the base, 6 to

15 inches (15.2 to 38.1 cm) long and 1.5 to 6 inches

(3.8 to 15.2 cm) broad. Heads are heterogamous, short

Seeding sagebrush with most ground equipment

requires that the seed be cleaned to at least 60 percent

purity. Most sagebrush seed is collected between 10 to

15 percent purity. To increase purity, the collected

material is run through a hammermill or barley

debearder followed by fanning and screening. Barley

debearders are preferred (Booth and others 1997;

Welch 1995). Hammermills tend to ball up the collected materials, rendering it almost impossible to

seed mechanically.

As seed purity increases, volume of material decreases. Seed cleaned to more than 30 percent purity

requires being mixed with a carrier such as rice hulls

or screened sawdust. Range seeders and drills are not

equipped to handle seed the size of sagebrush seed.

Small seed size requires that volume of material be

increased to help ensure proper seed metering. Plant

materials and floral parts that are collected with

sagebrush seed will not flow through and will plug

most ground rig seeding devises.

Most sagebrush seed matures and is dispersed

October through December. This is also the most ideal

time to seed. We do not recommend spring seedings.

Sagebrush seedlings are susceptible to frost damage

and are intolerant of short drought. Many sagebrush

seedlings that establish are located in, under, and

next to downed and dead plant materials and next to

rocks—all areas that provide some protection from

adverse climatic conditions and grazing animals. When

seeding sagebrush in a mixture with other species

through ground equipment, we recommend that sagebrush seed be separated in the seeding device from

seed of more competitive species (those with strong

seedlings and fast rate of growth; most grasses fall in

this group). Individual compartments in drill seed

boxes can be constructed with cardboard partitions

and duct tape. This will allow sagebrush and compatible species to be seeded separately but at the same

time as seeding the noncompatible species another

compartment. Seeding sagebrush aerially generally

results in the seed of the various species in the mixture

being separated so they are not in competition with

each other.

When dried and stored in an open, untreated,

uncooled warehouse, seed of most sagebrush species

will retain good viability for up to 3 years following

collection (Stevens and others 1981a).

Most sagebrushes transplant well. When standard

bareroot and container stock handling, planting, site

condition, requirements, principles, procedures, and

methods are followed, excellent establishment success

can be expected (Richardson and others 1986; Stevens

and others 1981b).

A majority of sagebrush seed is hand collected by

stripping or beating the seed into some type of container. Seed of some species can be collected with a

mechanical seed stripper.
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Chapter 21 Composite Shrubs

peduncled, and nodding, and are about 2.0 to 2.5 mm

high, 2.5 to 3 mm broad. There are five to 15 ray

flowers with 1.5 mm long corollas and 10 to 20 fivetoothed disk flowers with campanulate corollas, 1.5 to

2 mm long. There are eight to 18 bracts. The receptacle

is naked.

Ecological Relationships and Distribution

Oldman wormwood is native to the Mediterranean

areas of Europe, Asia, and Africa (Hall and Clements

1923). Its alternate common name, southernwood,

distinguishes it from common wormwood (A. absinthium). The latter is common and adapted in northern

Europe including Great Britain, whereas oldman

wormwood was recognized as an introduction in these

areas (Grieve 1931). In Britain, oldman wormwood

rarely flowers, perhaps because of the high latitude. In

North America, the plant is occasionally adventive in

the Eastern United States as a garden escape. In the

Intermountain area, however, it occurs only where it

has been planted. Few viable seeds are produced and

only rarely are seedlings found (Plummer 1977). A

dwarf form, var. nana 1 ft (30.5 cm) tall, and another

regular sized ecotype have long been maintained by

the USDA-ARS High Plains Research Station at Cheyenne, WY. Although seed production remains rare, a

higher frequency of viable seed production has occurred with these additional plant genotypes growing

in proximity to the formerly near sterile ecotype. This

suggests that outcrossing facilitates seed production

in oldman wormwood. It can be established by cuttings

in many soil types including raw subsoils in various

vegetation types big sagebrush to the subalpine, 5,000

to 10,500 ft (1,500 to 3,200 m) elevation, 12 to 40 inches

(30.5 to 101.6 cm) precipitation (Plummer 1974b,

1976, 1977). It can tolerate moderately alkaline to

moderately acidic soils (Plummer 1974b). Although

this species is adapted to subsoils such as roadcuts it

is not universally adapted to mine disturbances.

Plants are apparently sensitive to heavy metals or

other mine waste materials. Plants have not persisted

more than 5 years on acidic soils of the Idaho Batholith

(Monsen, unpublished).

Plant Culture

This species is, in essence, maintained vegetatively

in the Intermountain area. It is easily propagated by

stem cuttings. The process for establishing the plant

in a nursery or in outplantings is the same. Place

cuttings in the ground in spring when and where soil

is moist. Plummer (1974b) gave instructions for cuttings. Stems 10 to 18 inches (25.4 to 45.7 cm) and up to

0.5 inch (12.7 mm) in diameter give good results.

Smaller cuttings root well but are difficult to stick in

the ground. For best results, leaves and small twigs

should be removed from the cuttings. Sharpening the

large end of the cutting will facilitate penetration of a

hard soil surface and minimize damage to the cuttings. The cuttings perform best if planted within 48

hours after being taken. However, cuttings have rooted

after 6 weeks in cold storage (32 to 40 ∞F; 0 to 4.4 ∞C).

Cuttings should be kept moist and cool until planted.

In general, roots are forming and new buds leafing out

about 2 weeks after planting. Plants should be placed

about every 2 to 4 ft (0.6 to 1.2 m) for good stands.

Nursery plants remain vigorous and disease-free for

years (Nelson and Krebill 1981).

Uses and Management

Oldman wormwood is useful for stabilizing subsoil

sites (Plummer 1974b, 1977). Its deep extensive root

system makes the plant especially useful for erosion

control. Plants placed in staggered rows about 2 to 4 ft

(0.6 to 1.2 m) apart make good ground cover. It is

especially useful on steep sites such as roadcuts and

fills. On such sites, new roots are produced on freshly

covered stems with each earth movement, thus providing additional stabilization. This species is useful

as a nurse crop in that it helps stabilize and modify raw

soils so that natural invasion of plant materials from

the surrounding areas can occur. Individual plants

may persist for 15 to 25 years or more, but eventually

they will go out without reproducing themselves. By

that time, however, they have usually served their

purpose. Plants are consumed by small animals (at

times girdling the bark under snow) and insects, but

are not often eaten by large animals. Plants do provide

excellent cover for upland game species. Formerly the

plant was much used in herbal medicine as a tonic,

anthelmintic, emmenagogue, and antiseptic (Grieve

1931). It is also used in ornamental plantings (Hall

and Clements 1923; Plummer 1974b).

This plant requires little management. Nurseries

can be maintained as a source of cutting stock with

only routine maintenance. Occasional weed control

and supplemental water on dry sites is all that is

required. At outplanting sites where the plant has

been put in to help with soil stabilization no care is

necessary. These sites, however, often provide additional cutting stock.

Varieties and Ecotypes

Two varieties exist, the common one (var. abrotanum)

and a dwarf one (var. nana). We know of no plant

improvement work or ecotype comparisons. The common variety is best, in most cases, for raw soil stabilization planting because of its more vigorous growth.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 499

Chapter 21 Composite Shrubs

A B

Figure 3—Two subspecies of low sagebrush: (A) A. arbuscula ssp. arbuscula;

(B) A. arbuscula ssp. thermopola (Shultz

1986).

Artemisia arbuscula _____________

Low Sagebrush

Low sagebrush (fig. 2) is a low spreading, irregularly

branched shrub up to 20 inches (50.8 cm) high. The

slender, erect twigs are densely canescent but may

become nearly glabrous and thus darker green in

late summer. The plant layers infrequently. Leaves

are broadly cuneate, or fan shaped, 0.5 to 1.5 cm long,

0.3 to 1 cm wide, and are usually three-toothed (occasionally four to five in ephemeral leaves) or cleft at the

apex (fig. 3). Leaves on the upper part of the flowering

shoots may become entire. Flowerheads are grouped

into elongated, narrow racemose panicles. The heads

usually contain five to 11 disc flowers with corollas

3 to 4 mm long. The 10 to 15 involucral bracts are

canescent. Flowering occurs August to September,

depending upon strain and elevation. Seed ripens in

October and November (McArthur and others 1979a).

Ecological Relationships and Distribution

Low sagebrush grows on dry, sterile, often rocky and

alkaline, usually clay, soils between 2,300 and

11,500 ft (700 and 3,500 m) on approximately

39,112 mi2 (101,300 km2) in 11 Western States (Beetle

1960; McArthur and Plummer 1978; Ward 1953). In

the warmer, drier parts of its range, particularly in

southwestern Utah and Nevada, it may grow well into

the mountains above 9,800 ft (3,000 m). In some areas,

for example, east-central Idaho, low sagebrush occurs

on disjunct low and high elevation bands (McArthur

and others 1979a). Low sagebrush ranges from

southern Colorado to western Montana and west

throughout Utah and Idaho to northern California,

Oregon, and Washington. Normally its sites are drier

and more rocky than those on which big sagebrush

occurs. It may grow in mosaics with big sagebrush

where each is confined to a particular soil type. Low

sagebrush and black sagebrush only rarely occur in

intermixed stands, for example, Lost River-Lemhi

Range area of Idaho (McArthur and others 1979a). In

areas where the distribution of these two species

overlap, low sagebrush is usually found in the more

moist habitats or at slightly higher elevations than

black sagebrush (Tisdale and Hironaka 1981; Ward

1953; Zamora and Tueller 1973).

Plant Culture

Plants can be established on adapted sites by direct

seeding, broadcasting, and drilling. Seeding should

be conducted in the fall on disturbed soil. Care must

be taken to ensure that seeds are not covered more

than 1⁄16 inch (1.6 mm). Wildings of this species can

be transplanted in the spring or fall. Experimental

nursery plantings have successfully reproduced.

Plants of low sagebrush established at the Snow Field

Station were more vigorous than most other sagebrush plantings there (Nelson and Krebill 1981). Plants

establish well by direct seeding. Seedlings emerge

rapidly and grow quickly, although few attain large

stature at first. Roots develop rapidly, and plants

are able to survive dry surface conditions. Low

Figure 2—Low sagebrush growing on

shallow, stony soil in Logan Canyon,

Cache County, UT.
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Chapter 21 Composite Shrubs

sagebrush recovers quickly after burning and other

disturbances. It persists and invades sites seeded to

perennial grasses. It expresses good seedling vigor

and can be seeded with herbs. Seeds should be planted

at a shallow depth with a firm seedbed. Seed lots of low

sagebrush usually contain less inert or floral materials than processed seed lots of big sagebrush. Thus,

seeds can be more easily planted. Seed of low sagebrush is more difficult to collect than the more robust

big sagebrush. Seed prices are, thus, somewhat

higher, and lower quality seed is often sold. Low

sagebrush may often occur with little understory.

When seeded with herbs, a mixed community normally develops, but the understory often weakens

and recedes. The relationship with introduced and

native herbs is not well understood.

Uses and Management

On winter ranges, and to a limited extent on summer

ranges, low sagebrush is browsed by big game and

livestock (Kufeld and others 1973; McArthur and

others 1979a). Considerable variation exists in how

animals browse it in different locations. In Nevada a

gray-green form may be heavily browsed, while the

green form is only lightly browsed (Brunner 1972). In

addition to the direct browsing use of the plant, its

communities are important as habitat for a wide range

of domestic and wild animals (Dealy and others 1981).

Low sagebrush grows on difficult sites and is, therefore, not often a candidate for manipulation. Because

low sagebrush grows on exposed sites it is important

to wintering animals. Few other shrubs are adapted to

similar conditions. Consequently, maintenance of the

species is often critical. Attempts to maintain low

sagebrush on sites not similar to its origin have proven

unsuccessful. Seedlings usually establish when planted

“offsite,” and plants may attain maturity. However,

plants usually succumb after a few years with little or

no regeneration.

Varieties and Ecotypes

We are not aware of any substantial plant improvement work on this species. There are numerous ecotypes (Brunner 1972; McArthur and others 1979a)

and three subspecies (fig. 3a,b shows two of these

subspecies). Low sagebrush can apparently hybridize

with its Tridentatae relatives (McArthur and others

1979a). Beetle (1959, 1960) named a dwarf form

hotsprings sagebrush. This form occurs in the

Stanley Basin area of Idaho, Jackson Hole area of

Wyoming, east-central Oregon, and perhaps other

locations. Beetle speculated that this form arose as a

result of hybridization between typical low sagebrush

and threetip sagebrush. Another important low

sagebrush is Lahonton low sagebrush (Winward and

others 1986; Winward and McArthur 1995). This

plant occurs in western Nevada. We believe that it is

a stabilized hybrid between low and big sagebrush.

The big sagebrush parent is probably Wyoming big

sagebrush (Winward and McArthur 1995).

Artemisia bigelovii ______________

Bigelow Sagebrush

Bigelow sagebrush is a low shrub 8 to 16 inches

(20 to 40 cm) high with numerous spreading branches

(fig. 4). It is also known as flat sagebrush (Hall and

Clements 1923). The flowering stems are slender and

erect and bear inflorescences that are long, narrow

panicles with short, recurved branches (fig. 5). New

growth is covered with a silvery-canescent pubescence. The leaves of vegetative branches are similar

to those of big sagebrush. They are narrowly cuneate,

1 to 2 cm long, 2 to 5 mm wide, and normal tridentate,

but may show various abnormal tips. The odor of

crushed leaves is mild and pleasant. The heads are

arranged into elongated, narrow panicles and normally contain one, but occasionally zero to two, ray

flowers and one to three, usually two, disc flowers.

The turbinate involucre consists of eight to 12 short,

densely tomentose, bracts 2 to 4 mm long and 1.5 to

2.5 mm broad. Flowering occurs August to October.

Bigelow sagebrush closely resembles, and is often

mistaken for, low forms of big sagebrush produced by

overgrazing. In contrast to big sagebrush, it has ray

flowers. Furthermore, lobes of Bigelow sagebrush’s

vegetative leaves are always more shallow and more

sharply dentate than those of big sagebrush (McArthur

and others 1979a).

Figure 4—Bigelow sagebrush growing at the Snow

Field Station. The ruler is 12 inches (30 cm) long.
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Chapter 21 Composite Shrubs

Figure 6—Bigelow sagebrush, foreground,

growing with fourwing saltbush in a gravelly

draw near Bicknell, Wayne County, UT.

Ecological Relationships and Distribution

Bigelow sagebrush has a more southerly distribution than other sagebrushes. It is one of the most

drought-resistant of the sagebrushes. It occurs over

approximately 34,010 mi2 (88,086 km2) through western Texas, southern Colorado, New Mexico, Arizona,

Utah, Nevada, and California in canyons, gravelly

draws, and dry flats (fig. 6) from 3,000 to 7,900 ft

(900 to 2,400 m) (Beetle 1960; Kearney and Peebles

1960; Ward 1953). This species is often found mixed

with big sagebrush, black sagebrush, leafless green

rabbitbrush, shadscale, and especially broom

snakeweed. It is also characteristic in rocky soils of

the southern portion of the pinyon-juniper woodlands

(Hall and Clements 1923). Bigelow sagebrush is normally free of the common rust diseases and insect

galls common on other Tridentatae taxa (Beetle 1960).

However, in common with other Tridentatae, it is susceptible to a wilt disease (Nelson and Krebill 1981).

Seedlings of Bigelow sagebrush spread well naturally.

Seedlings grow rapidly and are robust and vigorous.

Direct seeding of this shrub has been limited to

experimental plantings. Where seeded, the shrub responds favorably and natural spread normally occurs

around the parental plant. Plants appear to have good

drought tolerance, even as young seedlings. Direct

seeding should not be a handicap to the use of this

species.

Figure 5—Bigelow sagebrush (McArthur and

others 1979a).

Plant Culture

Wildings can be successfully transplanted in the

spring or fall. Nursery plantings successfully reproduce. Seedlings grow in the wild, suggesting that

Bigelow sagebrush can be successfully direct seeded.

Uses and Management

Bigelow sagebrush is palatable to livestock and

game in all areas where it occurs. Its twigs are less

woody, the odor milder, and taste less bitter than most

of the big sagebrush complex (Hall and Clements

1923). This species can withstand considerable

browsing; generally more than other sagebrushes.

However, the plants are usually scattered, and consequently, not an abundant forage source except for

some large stands in the Four Corners area.

Varieties and Ecotypes

Ecotype variation has not been documented although it exists in chromosome races from 2x to 8x

(McArthur and others 1981; McArthur and Sanderson

1999). Bigelow sagebrush occupies a taxonomic position between the true sagebrushes (subgenus

Tridentatae) and the Artemisia (subgenus Artemisia)

species. We have chosen to treat it as a member of the

Tridentatae because of its characteristic growth habit,

woody anatomy, leaf form, chromosomal karyotype,

and RAPD molecular genetic markers (McArthur and

others 1981; McArthur and others 1998a). The confusion arises because its flowers have zero to two ray

flowers as well as the characteristic disc flowers of the

Tridentatae.
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Chapter 21 Composite Shrubs

Artemisia cana__________________

Silver Sagebrush

Silver sagebrush is an erect, freely branched, rounded

shrub up to 5 ft (1.5 m) tall. Older branches have dark

brown, fibrous bark while younger branches are covered with a dense white to yellowish-green tomentum.

Leaves on the vegetative branches are 1 to 10 mm wide

and 2 to 8 cm long, linear to linear-oblanceolate, entire

or occasionally with one or two irregular teeth or lobes,

silver-canescent becoming slightly viscid with age

(fig. 7). Leaves on the flowering stems are similar, but

they may be slightly smaller, especially on the upper

parts of the stems. The foliage emits a mild to pungent

aromatic odor when crushed. Numerous heads are

arranged into dense, narrow, leafy panicles, sometimes reduced to a raceme or spikelike inflorescence.

Stems layer when in contact with the soil, thus producing additional root systems. Each head contains

four to 20 disc flowers. Ray flowers are lacking.

Achenes are granuliferous. Blooming occurs during

August and September. Seed ripens in October and

November.

Ecological Relationships and Distribution

Aside from big sagebrush, silver sagebrush is the

most widely distributed sagebrush. It occurs over

approximately 53,221 mi2 (137,842 km2) in British

Columbia and Saskatchewan in the north, south to

Arizona and New Mexico, and west to Oregon and

California (Beetle 1960; McArthur and Plummer 1978).

It grows on soils that are less mature, with less

phosphorus, potassium, nitrogen, organic matter, and

lower cation exchange capabilities than do soils that

support big sagebrush (Hazlett and Hoffman 1975).

However, it may grow on a variety of soils (Morris and

others 1976).

Plant Culture

Wildings of silver sagebrush can be transplanted in

the spring or fall. Establishment by direct seeding

has also been successful (Kelsey 1986), especially

when done in the fall on the soil surface or at shallow

depths (Wasser 1982). In most natural situations,

however, few seedlings survive (Walton and others

1986). The species can also be propagated from hardwood cuttings. Harvey (1981) successfully rooted 87

percent of the cuttings of plains silver sagebrush.

He used 6 inch (15.2 cm) cuttings with the basal,

1.5 inch (3.8 cm) stripped of leaves, dipped in “rootone”

and placed in a misting propagation bench with a

growth medium of 50 percent sand, 25 percent vermiculite, 15 percent peat, and 10 percent perlite.

Monsen (personal communication) has had excellent

success seeding silver sagebrush on mine disturbances

on semiarid ranges in southern Idaho and Wyoming

and under nursery conditions. Seedlings develop rapidly often attaining a larger stature than plants of

basin big sagebrush. Seedlings respond dramatically

to moist conditions and may reach 15 to 20 inches (38.1

to 50.8 cm) in 1 year. Plants are usually quite uniform

in size and form. Young plants grow well with understory herbs. In undisturbed sites, conditions often

support a diverse understory. Plants appear well

adapted to harsh disturbances when artificially seeded,

but initially are slow to invade raw sites.

Use and Management

Silver sagebrush is important through its range as

a browse shrub and is used quite extensively by livestock and big game, particularly when other food is

scare (Kufeld 1973; Kufeld and others 1973; Wasser

1982). Bolander silver sagebrush was among the most

preferred of the sagebrush taxa offered to mule deer

and sheep in winter and fall feeding trials (Sheehy

and Winward 1981). Mountain silver sagebrush is

grazed by sheep in the fall after grasses and forbs

dry. In the Western Great Plains, silver sagebrush is

an important antelope survival food. Like big sagebrush, this species has been used by white settlers

and Native Americans for fuel (McArthur and others

1979a). Silver sagebrush is not as susceptible to fire

as many other sagebrushes (Wright and others 1979;

White and Currie 1983). In efforts to manage stand

densities by prescribed burning, White and Currie

(1983) reported that on both fall and spring burns fire

intensity was directly related to plant mortality and

Figure 7—Mountain silver sagebrush on Targhee

National Forest, Sublette County, WY, showing

leaves and flower heads.
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Chapter 21 Composite Shrubs

inversely related to subsequent growth. In general,

fall burns are more effective, as a silver sagebrush

control measure, than are spring burns. Internal water stress in silver sagebrush is greater as the season

progresses and soils dry out, and that stress is compounded in thick stands where there is interplant

competition (White and Currie 1984). An ecotype from

the Sheridan, WY, area shows promise as a fire resistant competition to cheatgrass (Monsen, unpublished).

Varieties and Ecotypes

No plant improvement work has been pursued on

silver sagebrush. The species, however, hybridizes

with other sagebrushes (Beetle 1960; McArthur and

others 1979a; Ward 1953). It is, in all likelihood, a

parent along with mountain big sagebrush for spicate

big sagebrush, which we believe to be a widely adapted

stabilized hybrid in Intermountain areas (Goodrich

and others 1985; McArthur and Goodrich 1986). It

has also been proposed that silver sagebrush is one of

the parents, along with early sagebrush, of the narrowly endemic coaltown sagebrush (Beetle 1960). Silver sagebrush is differentiated into three subspecies (fig. 8) separated from one another geographically,

ecologically, and morphologically (Beetle 1960, 1977;

Harvey 1981; McArthur and others 1979a; Shultz

1983; Walton and others 1986). RAPD genetic markers confirm both the relationship between and the

integrity of subspecies (McArthur and others 1998a,c).

Plains silver sagebrush is an erect, round, canescent, freely branched shrub up to 5 ft (1.5 m) tall. It

layers whenever conditions are suitable. This subspecies may spread rapidly, particularly after burning, by rootsprouting and by rhizomes (Beetle 1960).

Leaves of the vegetative branches are linearoblanceolate, entire or rarely with one or two irregular teeth or lobes, 1 to 10 mm wide, 2 to 8 cm long, and

are densely silky-canescent. Crushed foliage emits a

pungent turpentine odor (Beetle 1960; Ward 1953).

Flowerheads are usually arranged into dense, leafy

panicles and may contain one to 20 disc flowers.

Blooming occurs during September, and the seeds

ripen during October and November. It is octoploid, 8x

(McArthur and Sanderson 1999). Putative natural

hybrids between plains silver sagebrush and big sagebrush subspecies have been found (Beetle 1960; Ward

1953). This subspecies has a more eastern distribution

than the other two silver sagebrush subspecies. It

occurs southern Canada southward, but mostly east of

the Continental Divide, through Montana, the Dakotas, Wyoming, western Nebraska, and northern Colorado. It grows particularly well on the well watered,

deep soils of the northern Great Plains, especially

along stream bottoms and drainageways (Johnson

1978; McArthur and others 1979a; Walton and others

1986).

Mountain silver sagebrush (A. cana ssp. viscidula)

is an erect shrub that readily layers. It usually is not

more than 3.3 ft (1.0 m) tall. Leaves on the vegetative

branches are 1 to 5 mm wide, up to 7 cm long, and are

often crowded in dark green clusters. The leaves

typically are simple and entire, but occasionally are

variously toothed or lobed. This subspecies varies in

appearance but is always darker green than mountain

big sagebrush with which it is often growing (Beetle

1960). Mountain silver sagebrush is distinguished

from plains silver sagebrush by its smaller, darker

green leaves, its lower stature, and more western

distribution. Flowerheads are arranged into dense,

short racemes or spikelike inflorescences 1 to 3 cm

long. Each head contains four to 15 disc flowers.

Flowers bloom during August and September. Seed

matures during October and November. Mountain

silver sagebrush occurs in mountainous regions around

5,500 ft (1,800 m) and above. It is usually found in

areas of heavy, lingering snowpack from the southwestern corner of Montana, south along the Continental Divide to New Mexico, and west to Arizona,

Nevada, and Oregon (Beetle 1960, 1977; McArthur

and others 1979a; Tisdale and Hironaka 1981; Winward

1980). It occurs as both diploid, 2x, and tetraploid, 4x,

populations (McArthur and others 1981).

Bolander silver sagebrush (A. cana ssp. bolanderi)

has narrow leaves like mountain silver sagebrush, but

they are canescent like plains silver sagebrush. It is

distributed in poorly drained, alkaline soils from central

Oregon to eastern California (Beetle 1960; McArthur

and others 1979a; Tisdale and Hironaka 1981; Winward

1980). It is known only as diploid, 2x (Ward 1953).

Figure 8—Silver sagebrush subspecies:

(A) A. cana ssp. cana; (B) A. cana ssp.

bolanderi; (C) A. cana ssp. viscidula

(Shultz 1986).

A BC
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Artemisia filifolia ________________

Sandsage or Oldman Sage

Sandsage is a round, freely branching shrub up to

5 ft (1.5 m) tall (fig. 9). Young branches are covered

with a canescent pubescence while the older stems are

covered by a dark-gray or blackish bark. Leaves are

long, filiform, and silvery-white canescent; 3 to 8 cm

long, less than 0.5 mm wide, entire or alternately

divided into filiform divisions, and are often fascicled

(fig. 10). Numerous, nodding heads containing two or

three fertile, pistillate ray flowers and one to six

perfect, but sterile, disc flowers are arranged into

leafy, narrow panicles. Each head is subtended by

five to nine canescent involucral bracts. Both the

receptacle and achenes are glabrous. Flowers bloom

during August and September. Seed ripens from

October to December (McArthur and others 1979a).

Other seed characteristics are given in tables in chapters 24 to 26.

Ecological Relationships and Distribution

Sandsage is an excellent indicator of sand. It is probably the most widespread shrub on sand dunes and

sandhills the southern Black Hills of South Dakota

southward to Texas and Chihuahua and westward

to Arizona and Nevada (Hall and Clements 1923;

McArthur and others 1979a; Rasmsussen and

Brotherson 1986). In a study in southwestern Utah,

Rasmussen and Brotherson (1986a) found that

sandsage communities were floristically less diverse

than adjacent plant communities, but that there was

a higher density of plants in the sandsage communities. In the same study, they reported that despite the poor relative quality of the sandsage community soil

profile, sandsage nutrient quality was not affected.

Sandsage accumulates mineral nutrients well above

levels found in the soils in which it grows. At the study

locations, at least, sandsage is adapted to soils of low

fertility.

Plant Culture

Wildings can be transplanted in the spring or fall.

Plants can be easily reared from seed. Seeds are

smaller than those of most other sagebrushes but grow

well once established. Plants mature quickly even

when planted offsite. Survival is reduced, however,

when the species is planted out of its natural range

(Nelson and Krebill 1981). It grows well on infertile

soils. It has been seeded alone in most plantings but

appears to develop rapidly and can be seeded with

herbs adapted to arid sandy sites. It has been easy to

grow as bareroot stock and survives well when field

planted. It does not require special media for rearing

(Monsen n.d.).

Figure 9—Sandsage growing near Moccasin,

Mohave County, AZ.

Figure 10—Sandsage (McArthur and others

1979a).
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Uses and Management

The browse value of sandsage depends on where it

grows. It is seldom eaten in grasslands where other

food is adequate, but in more arid, desert regions it is

consumed by cattle, sheep, and big game. It is particularly important in dry years (Hall and Clements 1923;

McArthur and others 1979a). Sandsage is habitat for

some small nongame birds. This species helps prevent wind erosion by helping to stabilize light sandy

soils. It is an important candidate for harsh sandy

sites but often grows in such arid sites that seed

production is quite low. It is particularly useful in

blackbrush and pinyon-juniper sites where sandy outcrops occur.

Varieties and Ecotypes

No ecotypic differentiation has been documented,

but it may well occur. It does have some cytological

and chemical similarities with the true sagebrushes

(subgenus Tridentatae) although it is assigned to the

subgenus Dracunculus and differs from the Tridentatae in floral characteristics and wood anatomy

(Kelsey and Shafizadeh 1979; McArthur and Pope

1979; Moss 1940).

Artemisia frigida ________________

Fringed Sage

Fringed sage is a fragrant, aromatic, mat-forming

perennial subshrub 8 to 24 inches (20.3 to 61.0 cm)

tall (fig. 11). The lower woody stems are spreading and

often much branched. Adventitious rooting may occur

when stems contact the soil. The upper herbaceous

stems are erect and leafy. The whole plant is densely

silver-canescent. The numerous small, silky, canescent leaves are 6 to 12 mm long and are two or three

times pinnately divided (fig. 12). This species has a

deep perennial taproot with numerous extensive laterals that help it withstand drought. Numerous small

flowerheads are borne in nodding racemes or open

panicles. Small, densely hairy involucral bracts occur

in several series around each flowerhead. Each head

contains 10 to 17 outer, seed-producing, pistillate ray

flowers and numerous (25 to 50) tubular, funnelform,

perfect seed-producing disc flowers. Flower receptacles are densely villous. Fringed sage blooms from

June at high elevations and latitudes, to November

at lower elevations and latitudes. Seed matures between September and December (McArthur and others

1979a).

Ecological Relationships And Distribution

Fringed sage is probably, on a worldwide basis, the

most widely distributed and abundant species of

Artemisia. Its range extends from Mexico northward

through most of the western United States and Western Canada into Alaska, then on to Siberia, Mongolia,

and Kazakhstan (Harvey 1981; USDA Forest Service

1937). Fringed sage is a common plant of the high

plains along the eastern slope of the Rocky Mountains,

but also occurs in valleys and mountains. In the

United States, it is most abundant in the eastern and

northern parts of its range. This species ranges from

low semidesert valleys to more than 11,000 ft (3,400

m) elevation throughout the Rocky Mountain, Intermountain, and adjacent plains regions. Fringed sage

inhabits a wide variety of sites. Most typically, it

grows in full sunlight in dry, coarse, shallow soils. On

winter ranges in western Utah and eastern Nevada,

fringed sage may occur in dense stands along shallow

depressions that collect moisture from summer rains.

In such areas, it is frequently associated with winterfat,

shadscale, and rabbitbrushes. On plains, foothills, and

mountain slopes, this species may be associated with

a variety of grasses and forbs as well as with various

shrubs such as big sagebrush, Bigelow sagebrush,

sandsage, and especially in overgrazed areas, with

broom snakeweed. It is a common understory plant in

ponderosa pine in several Western States (McArthur

and others 1979a).

Figure 11—Fringed sage growing at Black

Mesa, Gunnison County, CO.
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Chapter 21 Composite Shrubs

Plant Culture

Fringed sage can be established by seed or by

transplanting young plants or segments of old plants.

Surface seeding on disturbed soils is recommended

(Wasser 1982). It produces an abundance of small

seeds, approximately 1,000 seeds per inch (2.5 cm) of

inflorescence (Harvey 1981). There are 4.5 million

seeds per lb (9.9 million per kg) (see chapter 24). In a

nursery, plants require little care and maintain reasonable vigor (Nelson and Krebill 1981). Individual

plants are susceptible to overgrazing, but on a populational basis, fringed sage seems to be stimulated and

increases plant numbers with heavy grazing pressure

(Cooperrider and Bailey 1986).

Uses and Management

The forage value of fringed sage varies considerably

with location and season (Dietz 1972). Its value as

browse is highest in late fall, winter, and early spring

on western ranges where it is eaten readily by big

game and livestock (Cooperrider and Bailey 1986;

Kufeld 1973; USDA Forest Service 1937; Wasser 1982).

It is also important food for sage-grouse (Wallestad

and others 1975). Nutritive quality is highest in the

spring, but remains adequate for animals for much of

the year (Cooperrider and Bailey 1986; Rauzi 1982). In

Figure 12—Fringed sage (McArthur and

others 1979a).

other areas, such as the grasslands of the Northwest

and Great Plains, fringed sage may be less palatable

and occasionally invades deteriorated grasslands. However, on the Great Plains, fringed sage is an important

winter antelope food and is used to a lesser extent the

year round (McArthur and others 1979a).

It is well suited for growth with grasses and broadleaf herbs. Its seedlings are competitive and can be

direct seeded into areas with herbaceous competition.

It is adapted to mine disturbances, perhaps better

than any other Artemisia. It can be used to control the

rapid influx of weeds on large disturbances.

This species has strong reproductive qualities and is

a good pioneer shrub for stabilizing disturbed areas. It

is often used in seeding western strip mines, especially

coal areas. It has excellent reproduction by seed, and

young plants or segments of old plants are readily

transplanted in early spring (McArthur and others

1979a). Its strong taproot and numerous lateral roots

make it effective in stabilizing gullies and reducing

soil erosion. These rooting characteristics make the

species capable of withstanding considerable grazing

and trampling use. Fringed sage has some value as a

medicinal plant (Hall and Clements 1923). It can be

controlled when it is too abundant (Alley 1972).

Varieties and Ecotypes

Undoubtedly, fringed sage, with its wide geographical and altitudinal distribution, has considerable genetic variation. However, no subspecific taxa

are recognized and no plant improvement work is

under way. Unlike several other widespread Artemisia species, fringed sage is known only at the diploid

(2n = 2x = 18) chromosome level (McArthur and Pope

1979).

Artemisia longiloba ______________

Alkali Sagebrush

Alkali sagebrush has also been called early sagebrush and low sagebrush. Alkali sagebrush is a misnomer because it grows mostly on neutral rather than

on alkaline soils (Passey and Hugie 1962b; Tisdale and

Hironaka 1981). It, however, is adapted to drought

conditions (Robertson and others 1966; Zamora and

Tueller 1973).

Alkali sagebrush is a low shrub up to 18 inches

(45.7 cm) tall (fig. 13). It has lax, spreading stems that

frequently layer. The bark is dark brown to black on

the older stems. The whole plant has a dark graygreen appearance (Beetle 1960). Leaves on the vegetative stems are broadly cuneate, up to 2 cm long, and

are deeply three-lobed (fig. 14). Leaves of the flowering stems are similar but smaller. This species is
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Chapter 21 Composite Shrubs

readily distinguished from other low statured sagebrushes by its large heads and early blooming (Beetle

1959). Its heads contains six to 11 disc flowers and

are 3 to 5 mm broad as opposed to 3 mm or less for

other short statured sagebrushes (low sagebrush and

black sagebrush). It flowers during mid-June to early

August and its seed ripens in August and September.

Beetle (1960) points out that this species has in the

past been confused with silver sagebrush because of

its large heads; with big sagebrush because of its

broadly cuneate, 3 lobed leaves; and with low sagebrush because of its dwarf size.

Ecological Relationships and

Distributions

Alkali sagebrush characteristically grows in heavy,

highly impermeable soils generally with dense “B”

horizons (Passey and Hugie 1962b; Tisdale and

Hironaka 1981). It is, however, also found on lighter

soils (McArthur and others 1979a). It occurs at elevations from 6,000 to 8,000 ft (1,800 to 2,400 m) along

foothills and in basins of the ranges forming the

Continental Divide in southwestern Montana, south

through Wyoming to northwestern Colorado, and scattered westward through northern Utah and Idaho to

Nevada and Oregon (Beetle 1960; Winward 1980).

Plant Culture

Little is known except the plant can be transplanted. Natural seeding occurs rapidly following

fires and other disturbances, yet mechanical tillage

can limit seedling establishment if seeds are buried

too deep or the seedbed is disrupted (Monsen and

Shaw 1986).

Uses and Management

The habitat of some stands of alkali sagebrush,

when in good condition, supports a mixed understory

of perennial grasses and annual and perennial forbs

providing forage and cover for sage-grouse, antelope,

and other wildlife as well as livestock (Dealy and

others 1981; Monsen and Shaw 1986). Other stands

are quite depauperate of associated vegetation and

provide little forage and habitat for animals. Alkali

sagebrush itself provides, depending on the site,

browse for sheep (often preferred), and habitat and

food for small animals (Dealy and others 1981;

McArthur and others 1979a). The plant should be

useful in rehabilitation plantings, particularly on

heavy, seasonally dry soils. It has been observed invading roadcuts in Echo Canyon, UT, and barrow pits

near Kemmerer, WY, (McArthur and others 1979a).

Monsen and Shaw (1986) reported that decadent

stands can be rejuvenated by mechanical means.

Sites that lack a satisfactory understory remain closed

stands unless these shrubs are reduced in number by

fire or mechanical means.

Varieties and Ecotypes

Alkali sagebrush has not been studied sufficiently

enough to make substantial comments here. However,

because of its relatively broad and scattered distribution and its differential use by animals, we suspect it

harbors considerable genetic variation. Beetle (1960)

considered it to be a parent to the restricted endemic

coaltown sagebrush.

Figure 13—Alkali sagebrush growing at

Wasatch Station, Summit County, UT. Note

the abundant, large flower heads.

Figure 14—Alkali sagebrush (Shultz 1986).
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Chapter 21 Composite Shrubs

Artemisia nova _________________

Black Sagebrush

Black sagebrush is a small spreading, aromatic

shrub 6 to 8 inches (15.2 to 20.3 cm) or occasionally to

30 inches (76.2 cm) tall with a dull grayish-tomentose

vestiture that causes most populations to appear

darker than big sagebrush and low sagebrush (fig. 15).

However, some forms may be as light in color as those

species (Beetle 1960; Brunner 1972). Numerous erect

branches arise from a spreading base (fig. 16). This

shrub had not been observed to layer or stump sprout

(Beetle 1960; Tisdale and Hironaka 1981). However,

some layering of black sagebrush on a roadcut near

Kolob Reservoir, Washington County, UT, has been

noted (McArthur and others 1979a). Typical leaves

are evergreen, cuneate, viscid from a glandular pubescence, 0.5 to 2 cm long, 2 to 8 mm wide, and threetoothed at the apex. The uppermost leaves, particularly on the flowering stems, may be entire.

Flowerheads are grouped into tall, narrow, spikelike

panicles that extend above the herbage. The inflorescence stalks are red-brown and persistent. The heads

usually contain three to five disc flowers with corollas

1.8 to 3 mm long. The eight to 12 involucral bracts are

greenish-yellow and nearly glabrous. The principal

difference between black sagebrush and low sagebrush is that low sagebrush has five to 11 flowers per

head, 10 to 15 canescent involucral bracts, and is light

in color. Also, the flower stalks of black sagebrush are

denser, much darker, and more persistent than those

of low sagebrush (McArthur and others 1979a; Ward

1953). Black sagebrush is characterized by the presence of leaf hairs readily visible at 10x magnification

when compared to its relatives low sagebrush, alkali

Figure 15—Black sagebrush, with dark flower

stalks, and Wyoming big sagebrush growing

at ecotone near Gabbs, Nye County, NV.

Figure 16—Black sagebrush (Shultz 1986).

sagebrush, threetip sagebrush, and the subspecies of

big sagebrush (Kelsey 1984). Three-fourths of a large

sample (n = 152) of black sagebrush plants across the

species range showed this characteristic conspicuously, whereas, no other taxa exceeded a 4 percent

display (n = 1,849).

Ecological Relationships and Distribution

Black sagebrush covers approximately 43,301 mi2

(112,150 km2) in the 11 Western States (Beetle 1960).

It is most abundant at elevations from 4,900 to 7,900 ft

(1,500 to 2,400 m) and normally grows on drier, more

shallow stony soil than basin, mountain, or Wyoming

big sagebrushes or low sagebrush (Beatley 1976;

Tisdale and Hironaka 1981). Zamora and Tueller

(1973) reported root restricting layers at depths of

11 to 27 inches (28.0 to 68.6 cm) in half of their black

sagebrush study sites. Other soils in which black sagebrush commonly occurs are usually underlain by gravelly and sandy loam strata. Most, but not all, soils

supporting black sagebrush are calcareous. Most

black sagebrush stands do not burn because of their

sparseness (Tisdale and Hironaka 1981). However, we

have observed large burns of black sagebrush stands

in central Utah. Insect galls are numerous on black

sagebrush, but rust diseases are less common (Beetle

1960). Nelson and Krebill (1981) found black sagebrush to be less susceptible to a wilt disease than

most other Artemisia taxa growing at the Snow Field
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Station in Ephraim, UT. Clary (1986) reported that

winter livestock grazing (principally sheep) has had a

significant and often severe effect on black sagebrush.

Some large areas in Nevada formerly occupied by

black sagebrush have been invaded recently by Utah

juniper and singleleaf pinyon pine.

Plant Culture

Black sagebrush has been successfully transplanted

(Luke and Monsen 1984). Seed production in nurseries is often good. It should be seeded in fall or early

winter. Seeds should be covered lightly up to 0.25 inch

(6.4 mm) deep. Once established, black sagebrush can

spread rapidly (Stevens 1986b).

Seeds of black sagebrush are larger than those of

most sagebrushes (see chapters 20 and 24). Seeds

tend to remain attached to the flower but can be processed easier than seeds of other sagebrush. Seedlings

are vigorous, and young plants often grow rapidly.

However, even under favorable conditions, young

plants do not grow as rapidly as big sagebrush. Once

established, small seedings persist well even under

adverse conditions. Plants are able to establish amid

rocky exposed surfaces. Under these conditions the

shrub seedlings persist and usually exclude weedy

annuals. Black sagebrush can be established when

seeded on sites other than its origin. It grows well on

areas normally occupied by big sagebrush and may

persist and reproduce. Young stands grow well with

seeded herbs, but as stands mature a reduction in

understory herbs occurs. Black sagebrush establishes

well from direct seeding. The larger seed is not as

likely to be seeded too deep as smaller seeded species

of sagebrush. Under favorable moisture conditions,

mature plants produce abundant seed crops. Natural

spread occurs quickly, consequently interseeding or

other techniques can be used to establish a seed source

and allow natural spread to populate large areas.

Use and Management

Black sagebrush is usually held in high regard as a

palatable forage for wildlife and livestock, especially

sheep, antelope, and deer (Clary 1986; McArthur and

Plummer 1978). However, it, like big sagebrush, has

populations that differ dramatically in mule deer

preference (Behan and Welch 1985; Welch and others

1981). Welch and his colleagues (Behan and Welch

1985; Welch and others 1981) have shown that some

accessions are not eaten whereas others are highly

preferred—one accession had 60 percent of its current

annual growth eaten in 1978 and 82 percent in 1982.

Scholl and others (1977) also reported accessional

differences of mule deer preference between black

sagebrush accessions. In comparison with other sagebrushes, black sagebrush was preferred by mule deer

(Nagy and Regelin 1977); however, Smith (1950) and

Sheehy and Winward (1981) found no such preference. Striby and others (1982) reported that mule deer

and elk preferred black sagebrush over basin big

sagebrush, but not over Wyoming and mountain big

sagebrushes. However, Wambolt (1996) found that

elk and mule deer preferred black sagebrush least in

comparison with mountain, basin, and Wyoming big

sagebrush. In the preference tests it should be remembered that there are considerable intraspecific

variations as well as interspecific ones being compared. Black sagebrush has good winter nutritive

quality (Welch 1983b); not as good as big sagebrush, but

the two are adapted to different habitats. Black sagebrush is a good conservation plant for dry, shallow,

stony soils. The plant is an aggressive natural spreader

from seed. Because it usually grows on dry rocky sites,

it is usually not a candidate for plant control.

Varieties and Ecotypes

Welsh and Goodrich (1995) recently described a new

variety, A. nova var. duchesnicola to go with the

longstanding typical taxon, A. nova ssp. or var. nova.

Typical black sagebrush occurs in two color morphs

(Beetle 1960). The gray-green form is generally browsed

more than the darker glossy-green form (Brunner

1972; Stevens and McArthur 1974; Winward 1980).

Welch and others (1981) and Behan and Welch (1985)

have shown that an accession Pine Valley Ridge in

Millard County, UT, is a fine candidate for wider

planting because of browsing animal preference. That

accession has been transplanted and survived in several locations. It has been released as a germplasm

(Welch and others 1994). Black sagebrush has many

ecotypes that deserve additional attention. Black sagebrush may have had a role in the parentage of Wyoming big sagebrush (McArthur 1983a; Winward 1975).

Beetle and Johnson (1982) reported that some forms of

black sagebrush approach Wyoming big sagebrush

in appearance. We have made similar observations on

the Kaibab foothills in northern Arizona.

Artemisia pygmaea ______________

Pygmy Sagebrush

Pygmy sagebrush (fig. 17) is a dwarf, depressed,

evergreen, cushionlike shrub less than 8 inches

(20.3 cm) tall. Bark on older stems becomes dark

brown and fibrous. On young branches the bark is

nearly white to straw-colored and somewhat puberulent. Leaves on the vegetative stems are green, nearly

glabrous, viscidulous, 2 to 4 mm wide, 2 to 8 mm long,

and are pinnatified with three to 11 lobes, or sometimes may be only toothed (fig. 18). Leaves on the

flowering branches are usually reduced and may be
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Chapter 21 Composite Shrubs

entire. Heads with three to five disc flowers are arranged into spikelike inflorescences. Ray flowers are

lacking. Twelve to 18 greenish-yellow bracts subtend

each head. Achenes are glabrous. Flowers bloom in

August and September, and seed matures in October.

Seeds are large for Artemisia (McArthur and others

1979a; see chapter 24).

Pygmy sagebrush is limited to calcareous soils in

desert areas over approximately 21 mi2 (54 km2) eastern

Figure 17—Pygmy sagebrush growing at the

Snow Field Station, Sanpete County, UT.

Figure 18—Pygmy sagebrush (Shultz 1986).

Utah to western Nevada and northern Arizona (Beetle

1960; McArthur and Plummer 1978; Ward 1953). In

Nevada, this species is often associated with halophytic threadleaf rubber rabbitbrush. Some fairly

large stands occur with black sagebrush in Utah.

Because of its scarcity and small size, this species

has little value as browse although it is eagerly taken

when available. It does, however, provide important

ground cover in the dry, alkaline areas where little

else will grow. It establishes readily by transplanting

divided plants. Although it spreads well from naturally dispersed seed, artificial planting of seed has not,

as yet, been successful. This is due probably to poor

quality seed and not the inability to establish by

artificial planting. It is a good candidate to plant in

heavy soils. It may be of considerable value for mine

and roadway disturbances. Natural spreading has

occurred when plants have been established in research plantings.

Artemisia rigida _________________

Stiff or Scabland Sagebrush

Stiff sagebrush is a low, pungently aromatic shrub

with thick, rigid, somewhat brittle branches up to

16 inches (40.6 cm) high (fig. 19). It is not known to

resprout or layer. The wider deciduous, silverycanescent, spatulate leaves are mostly 1 to 4 cm long

and deeply divided into three to five narrowly linear

lobes (fig. 20). Occasionally some leaves are linear

and entire. The inflorescence is a leafy spike with

heads sessile or in small clusters in the axils of their

subtending leaves, which generally are all longer than

the heads. The campanulate involucre is 4 to 5 mm

long with numerous, canescent bracts. Each head

consists of five to 16 perfect disc flowers. Flowering

occurs from late August to early October; seeds ripen

in October and November (see chapters 20 and 24).

This species resembles threetip sagebrush somewhat

in its small size, silvery pubescence, and the deeply,

narrowly lobed leaves, but may be distinguished by

the spikelike inflorescence, large leafy bracts that

subtend the heads, and the deciduous leaves.

Stiff sagebrush occurs in dry rocky scablands in the

Columbia and Snake River basins and spills over into

the northern end of the Great Basin. It occurs primarily, if not exclusively, over basaltic bedrock

(Daubenmire 1982; Tisdale and Hironaka 1981). It

grows at elevations from 3,000 to 5,000 ft (910 to

1,500 m) in Idaho, central and eastern Oregon, and

central and eastern Washington. Otherwise its ecological niche is similar to that of low sagebrush (Ward

1953). The report of this species growing in Montana

are apparently in error (Morris and others 1976;

McArthur and others 1979a).
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Chapter 21 Composite Shrubs

Figure 20—Stiff sagebrush (Shultz 1986).

Because of its scant foliage and stiff branches, stiff

sagebrush has been considered of little value for

browse, except for sheep (Hall and Clements 1923). It

provides forage on the dry sites on which it grows in

the midsummer when herbaceous plants are dry.

This species provides cover on the poor rocky soils

where it grows. It appears to have a wider range of

adaptation than is indicated by its present natural

range of occurrence, thus giving it potential use in

reclamation of some harsh disturbed sites.

Seed production is normally quite low and seed

quality is questionable (Monsen, unpublished). Attempts to seed and transplant this species in arid

sagebrush ranges near Boise, ID, have not been successful. In most cases plantings were made on sites

where cheatgrass and other annuals had invaded the

shrublands. Stiff sagebrush seedlings were not able

to compete with the weeds. Plants grow slowly and

appear to require 2 to 4 years to establish and

attain reasonable size. The invasion of cheatgrass

undoubtedly limits the natural distribution and

occurrence of this shrub. However, it is an important

plant for harsh rocky sites. Plantings have successfully established on prepared seedbeds. It is able to

seed naturally on harsh sites and has been able to

survive wildfires.

Artemisia spinescens ____________

Budsage

Budsage is a low, spinescent, pungently aromatic,

rounded shrub 4 to 20 inches (10.2 to 50.8 cm) high

(fig. 21). It is profusely branched from the base and

has white-tomentose pubescence on young twigs and

leaves. This pubescence is grayish and stiff on older

branches. Leaves are small, mostly 2 cm or less in

length, including the petiole. Leaves are three to five

palmately parted, with the divisions again divided

into three linear-spatulate lobes. Leaves are crowded

on the short stems, with those near the apex being

smaller and more entire (fig. 22). Unlike most species

of Artemisia, budsage is deciduous, with the leaves

falling by midsummer. Early in the spring, when

budsage first shows signs of breaking dormancy, but

before the buds elongate, the bark the last season’s

growth can easily be pulled off. At this developmental

stage, budsage is sought out by big game and livestock.

Sheepmen refer to this condition as “slipping.” As

early as February or March, new bright-green leaves

are produced.

Budsage is well adapted to xeric conditions. It has

an extensive root system that grows primarily in the

top 6 to 22 inches (15.2 to 55.9 cm) of soil. Interxylary

cork is formed annually over the last year’s wood in

both the roots and the stem. This layer of cork restricts

Figure 19—Stiff sagebrush growing at the Snow

Field Station, Sanpete County, UT. The ruler is

30 cm long.
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Chapter 21 Composite Shrubs

the upward movement of water to the narrow zone of

wood formed by the current year’s growth. The corky

tissue develops during early summer and thus helps

to prevent excessive water loss during the dormant

season (Wood 1966). Many other xerically adapted

woody Artemisia also have interxylary cork (Moss

1940). Budsage bears small flowerheads, 3 to 5 mm

long, in glomerate racemes of one to three heads in leaf

axils of the flower branches. Each head contains two

to six fertile, pistillate ray flowers and five to 13

perfect, but sterile, disc flowers with aboritive ovaries.

The loose flowerheads are held together by long, matted hairs that cover the corolla and especially the

achenes. The heads often fall off the plant intact,

without breaking apart to release the seed. However,

some seeds are usually dispersed independently. In

some instances, seeds germinate while still in the

head (Wood 1966). Seeds mature by early June. Good

seed production occurs infrequently. The flowers

bloom so early in the spring that developing embryos

frequently are frozen. Abundant reproduction occurs

in years of plentiful seed and favorable moisture.

Figure 21—Budsage growing near Sevier

Lake, Millard County, UT.

Figure 22—Budsage (McArthur and others 1979a).

Terminal and lateral buds of budsage generally expand and begin to elongate in late March and early

April during the latter part of the “slipping” period.

Blooming normally occurs the last week in April

through the last week in May, although it has been

found in bloom as early as late March and as late as

mid-June (Wood 1966). Although budsage ordinarily

begins growth early in the spring and then becomes

dormant by early or midsummer, it occasionally may

break dormancy in response to late summer storms.

The plants then remain green all winter to provide

succulent forage throughout the winter and spring.

Ecological Relationships and Distribution

Budsage is a drought-resistant shrub quite common

in semiarid valley bottoms, benches, and foothills over

much of the interior Western United States (Hall and

Clements 1923; Wood and Brotherson 1986). It occurs

from Wyoming through southwestern Montana,

southern Idaho, and eastern Oregon south to northwestern New Mexico, northern Arizona, and southeastern California. It is often associated with shadscale,

winterfat, and other salt-tolerant shrubs (McArthur

and others 1979a; Wood and Brotherson 1986).

Plant Culture

Summer dormancy can be broken by supplemental

water. Wood and Brotherson (1986) reported that

supplemental water advanced the beginning of

growth by 2 weeks, but did not increase total productivity by the time of the onset of cold weather. Seed

quality is usually low, but during infrequent years,

natural seeding occurs. Some sites have improved well

through protection from livestock grazing. Blaisdell

and Holmgren (1984) reported significant recovery of

budsage on areas protected from spring grazing. Artificial seeding has been hampered by the use of low

quality seed, not the inability of the plant to establish

from direct seeding. Plants do not reach maturity

quickly, but seedlings and young plants are competitive. They persist under adverse conditions where few

other species exist. Plants grow well with winterfat

and scattered amounts of grass. Disking is detrimental to the survival of budsage, but anchor chaining

usually has little effect upon the shrub.

Uses and Management

Budsage is a palatable, nutritious forage plant for

upland birds, small game, big game, and sheep in the

winter. It is especially high in calcium, magnesium,

phosphorus, and protein (Wood and Brotherson 1986).

Generally, it is more palatable in the late winter than

during the early winter (Holmgren and Hutchings

1972). During this time it is of tremendous value to

the welfare of grazing animals, especially where there
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Chapter 21 Composite Shrubs

is an abundance of dry grass. Care must be taken in

grazing budsage in late winter and early spring. Even

light grazing during this period is detrimental, and

continual heavy grazing may eliminate budsage from

areas (Holmgren and Hutchings 1972).

Varieties and Ecotypes

Despite its wide distribution and isolated population, little morphological variation has been described

in budsage. However, polyploidy is known (McArthur

and Pope 1979).

Artemisia tridentata _____________

Big Sagebrush

Big sagebrush (Artemisia tridentata) is a highly

polymorphic species with numerous ecotypes and biotypes (fig. 23). Three subspecies—basin, Wyoming,

and mountain (tridentata, wyomingensis, and

vaseyana) big sagebrush—are generally recognized

(Beetle 1960; Beetle and Young 1965) (see chapter 21).

Additional subspecies, spicate big sagebrush

(spiciformis) and xeric big sagebrush (xericensis), have

recently been proposed (Goodrich and others 1985;

Rosentreter and Kelsey 1991). Subspecies will be

discussed in the Varieties and Ecotypes section. Big

sagebrush is composed of aromatic, evergreen shrubs

ranging from dwarf to tall, arborescent forms up to 15

ft (4.6 m) tall (fig. 24). The lower forms generally have

several main stems arising from the base, whereas the

tall forms often have a single short trunk. Older

branches are covered with a gray to brown to black

shredded bark. Younger branches and leaves have a

Figure 23—Individual big sagebrush plants of

basin, mountain, and Wyoming subspecies.

Plants of each subspecies are randomly placed

at the Gordon Creek Wildlife Management Area,

Carbon County, UT. Plants are even aged.

Figure 24—Large basin big sagebrush

near Meeker, Rio Blanco County, CO.

white to gray tomentum that gives the plants a silvery

cast. Typical leaves are narrowly cuneate or oblanceolate and terminate with three blunt teeth at their

truncate apexes. However, considerable variation occurs, ranging from linear, entire leaves with rounded

to acute apexes (fig. 25) to broadly cuneate leaves

with varying numbers of teeth or shallow lobes. The

leaves also range in size from 2 mm to 2 cm broad and

1 cm to 6.5 cm long. Normally, leaves of vegetative

shoots are more characteristic and less variable than

those on flowering shoots. Also, persistent, that is

overwintering, leaves are less variable than leaves of

the spring growth flush (Miller and Shultz 1987;

Winward 1970, 1980), which are shed by midsummer.

Heads of this species contain three to eight disc flowers, (except spicate big sagebrush, which may have

up to 12 flowers per head) and are arranged into leafy

panicles with erect or sometimes drooping branches.

In some forms, the inflorescence becomes spikate.

Blooming occurs from July to October. Seeds mature

in October, November, and December. Big sagebrush

plants often live 70 to 100 years or more. Specimens

from several sites were found to be more than 200

years old (Ferguson 1964). Usually, however, a high

rate of individual plant turnover occurs within a 20

year period (Stevens 1986b). Characteristics of big

sagebrush are detailed in table 3.
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Chapter 21 Composite Shrubs

Ecological Relationships and Distribution

Big sagebrush is the most widespread and common

shrub of Western North America. It is especially

common in the Great Basin. Beetle (1960) reports the

species as covering approximately 226,374 mi2

(586,309 km2) in the 11 Western States. Beetle’s estimate of coverage for this and other species is probably

too high, because it includes areas where Artemisia

are partial dominants (McArthur 1981, 1983b;

A B

Figure 25—Four subspecies of big sagebrush: (A) A.

tridentata ssp. tridentata; (B) A. tridentata ssp. spiciformis;

(C) A. tridentata ssp. vaseyana; (D) A. tridentata ssp.

wyomingensis (Shultz 1986).

C D

McArthur and Ott 1996). Nevertheless, his figures

show how extensively sagebrush is distributed and

the relative abundance of each Artemisia species when

compared to related species. Big sagebrush grows in a

variety of soils on arid plains, valleys, and foothills to

mountain slopes from 1,600 to 11,200 ft (490 to 3,400

m) and is frequently associated with such shrubs as

shadscale, saltbush, rubber rabbitbrush, low rabbitbrush, fourwing saltbush, spiny hopsage, spiny

horsebrush, winterfat, broom snakeweed, antelope

bitterbrush, snowberry, and serviceberry. Although it

is tolerant of quite alkaline as well as quite acidic soils,

its optimum growth is in deep, fertile, alluvial loams

(Sampson and Jesperson 1963).

Although big sagebrush has spread with settlement

of the West (Cottam 1961; Christensen and Johnson

1964; Hull and Hull 1974), it was clearly an important

and widespread plant before this settlement. Vale

(1975), quoting early pioneer and explorer diaries, has

shown it was a common Western plant prior to 1850.

Johnson (1986) presented photographic evidence

showing sagebrush as a widespread dominant in the

1870’s. As early as the Pleistocene Epoch, sagebrush

was already an important part of the Intermountain

flora. Some 80,000 to 10,000 years ago, sagebrush

dominated large tracts of land in areas where it is still

found (Tidwell and others 1972; Van Devender 1977).

Big sagebrush is an indicator of site quality (Kearney

and others 1914; McArthur and Welch 1982; Passey

and others 1982; Stevens and others 1974). Some

populations, especially basin big sagebrush, have

been taken out and the land they occupied put into

agriculture.

Big sagebrush and its subgenus Tridentatae relatives are subject to insect and microbial pests and

benefactors—most notably the sagebrush defoliator

moth (Aroga websteri) (see chapter 21). Wide ranging

and periodic outbreaks of this insect have caused

extensive sagebrush mortality over much of the range

of A. tridentata and its relatives (Hall 1965; Henry

1961; Hsiao 1986; McArthur and others 1979a). However, except on some critical winter game ranges,

Aroga websteri is not believed to have serious, longterm effects. The moth is subject to insect parasites

and predators and does not completely kill entire

sagebrush stands (Hall 1965; Hsiao 1986). In time,

sagebrush naturally reinvades its old sites that in the

meantime have become more diverse plant communities. During drought, grasshoppers have been known

to defoliate big sagebrush (Schlatterer, personal communication). Galls of many kinds of flies (Diptera) are

found on sagebrush (Hall 1965; Jones 1971). The

effect of the galls is not known. Sagebrush hosts many

others insects of various orders (see Graham and

others 1995; Messina and others 1996). Some of

these may protect the plant disease vectors. Several

microbial-induced diseases are known (Krebill 1972).
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Chapter 21 Composite Shrubs

Table 3—Characteristics of subspecies of Artemisia tridentata (adapted from McArthur 1983b).

Subspecies

Characteristics tridentata vaseyana wyomingensis spiciformis References

Habitat and range Foothills and Foothills and Foothills and valley High mountain Beetle and Young 1965;

valley floors. mountains. floors. 2,500 to 7,000 ft areas. 7,000 to Morris and others 1976;

4,000 to 7,000 ft 3,000 to 19,000 ft (760 to 2,135 m). 12,000 ft (2,135 to Winward and Tisdale 1977;

(1,220 to 2,135 m). (915 to 5,790 m). Montana, Washington 3,660 m). McArthur and Plummer 1978;

British Columbia British Columbia to Arizona. Oregon and McArthur and others 1979a;

and Montana to and Alberta to Montana to Dealy and others 1981;

Baja California California and Nevada and Winward 1980; Harvey 1981.

and New Mexico. New Mexico. Colorado.

Smell Biter pungent Pleasant Bitter pungent Pungent, not bitter McArthur and others 1974.

Essential oil x = 1.4 percent x = 2.2 percent x = 1.1 percent x = ? Welch and McArthur 1981.

Leaf shape narrowly cuneate Cuneate to Cuneate Cuneate to Marchand and others 1966;

spatulate narrowly McArthur and others 1974;

cuneate McDonough and others

1975; Winward and Tisdale

1977.

Common height 3 to 13 ft 2 to 5 ft 1.5 to 3 ft 2 to 5 ft McArthur and others 1979a;

ranges (0.9 to 4 m) (0.6 to 1.5 m) (0.5 to 0.9 m) (0.6 to 1.5 m) Winward 1980.

Sesquiterpenes 4 to 7 3 to 6 2 5 Kelsey and others 1973.

compounds

Ultraviolet Trace Abundant Trace, but often Abundant Shafizdeh and Melinknoff

visible more than 1970; Stevens and McArthur

coumarins tridentata. 1974; Brown and others

1975; McArthur and others

1981.

Tendency to None Mild None Very strong Beetle and Young 1965;

layer Winward 1980; Goodrich

and others 1985.

Palatability to Low Usually highest Highest for sheep High for sheep in Hanks and others 1973;

deer and sheep sometimes high late summer and Sheehy and Winward 1981;

for deer. fall. Sometimes Welch and others 1981;

under snow in Striby and others 1982;

season of use. Welch and McArthur 1986;

unpublished; Welch and

others 1987.

Protein content High Low Low ? Welch and McArthur 1979b.

Seed germination High Low Intermediate ? Harniss and McDonough

prior to stratification 1976.

2n chromosome Commonly 18, Commonly 18, Consistently 36 18, 36 Ward 1953; Taylor and

some 36 some 36 others 1964; Winward and

Tisdale 1977; Kelsey and

others 1975; McArthur and

others 1981; McArthur and

Sanderson 1999

Flower and seed Late Early Late-intermediate Early Marchand and others 1966;

phenology Hanks and others 1973;

Winward and Tisdale 1977.

Flower stalk to x = 1.8 x = 3.4 x = 2.2 x = 3.1 Winward and Tisdale 1977.

vegetative shoot

length ratios

Flower per head 3 to 6 4 to 8 3 to 8 6 to 12 Beetle and Young 1965;

Winward and Tisdale 1977;

Goodrich and others 1985.

Effect of fire on Negative Positive Neutral Positive? Chaplin and Winward 1982.

seed germination

Leaf and stem water Lower Higher Lower Higher? Miller and others 1982;

potential differences Shumar 1984; McArthur

and others 1998b

Shrub shape Uneven topped Even topped Uneven topped Even topped Winward 1980; Goodrich

(usually) and others 1985.

Position of Throughout Above crown Throughout Above crown Winward 1980; Goodrich

flower stalks crown crown and others 1985.

Seedling growth Longest sustained Slowest root Both above ground and ? Welch and Jacobson 1988;

root growth rate growth rate root growth is initially Booth and others 1990.

more rapid than other ssp.
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Chapter 21 Composite Shrubs

Some of these are widespread and may be locally

destructive; however, sagebrush populations are resilient and generally are not significantly affected in

the long run. Nelson and Krebill (1981) isolated several fungal species of the genera Gliocladium,

Fusarium, and Rhizoctonia from dying sagebrush in

uniform gardens. Similar symptoms (dying desiccated

plants) have been observed in natural populations

(McArthur and others 1979a). Nelson and Sturges

(1986; Sturges and Nelson 1986) reported that

snowmold disease can be serious in mountain big

sagebrush populations. Diseases induced by these

fungi may be among the most serious sagebrush diseases. Some microbes are likely useful for the vigor

and growth of sagebrush. Wallace and Romney (1972)

found preliminary evidence that big sagebrush

formed symbiotic relationships with microbial endophytes to fix atmospheric nitrogen. Williams and

Aldon (1976) found endomycorrhizae within big sagebrush roots and abundant spores around the roots.

Endomycorrhizae, in general, have a beneficial influence on plant growth by promoting nutrient absorption through infected roots (Williams and Aldon 1976).

For sagebrush, the beneficial effect has recently been

demonstrated (Allen 1984).

Plant Culture

Big sagebrush is one of few shrubs with potential for

easy establishment by artificial seeding. Seedlings

grow rapidly and compete well with herbaceous

plants. The plant establishes well from broadcast

planting on irregular but firm seedbeds. Consequently,

plants can be established by broadcast seeding even

when seeded in mixtures with herbs. Natural seeding

often takes place amid established stands of seeded

grasses. Because natural spread generally occurs,

sagebrush can be planted at low rates and allowed to

spread. This has been successful.

Sagebrush seed is difficult to separate from the

chaff. Large stalks and debris are usually removed

and seed is sold and seeded at a purity of 10 to 20

percent. Seeds are small and planting rates of 0.10

PLS (1 lb bulk per acre or 1.1 kg per ha) can produce

adequate stands from broadcast or drill seeding. Seeds

require minimal coverage but must be planted on a

firm seedbed. Planting on a loose surface is not advised. Seeding should be delayed until early winter

when soil surfaces are moist and firm. Seeds require

afterripening and stratification to attain uniform

germination. Small seedlings are not frost tolerant,

and extensive losses occur from spring frosts.

Sagebrush often establishes better from broadcast

seeding than drill seeding, as seed are placed too deep

with conventional drills. Seedings using the cultipacker

seeder are successful as seeds are not placed more

than 0.25 inch (6.4 mm) deep and the soil surface is

compacted or “firmed up” with this machine.

Although established stands of big sagebrush can

reduce the productivity of associated herbs, young

sagebrush plants do not diminish the density or vigor

of developing herbs. Consequently, sagebrush can be

directly seeded with herbs to provide a mixed composition of plants. Big sagebrush is also compatible with

numerous shrubs, so mixed seedings can be done even

though young sagebrush plants develop more rapidly

than serviceberry, snowberry, Martin ceanothus,

green ephreda, and Wyeth eriogonum.

Big sagebrush wildings can easily be transplanted

into nurseries and field locations when they are relatively dormant and moisture conditions are right—

fall and early spring. Wildings about 6 to 8 inches

(15.2 to 20.3 cm) tall transplant best. McArthur and

Plummer (1978) reported that of the 829 big sagebrush plants transplanted to the Snow Field Station

in Ephraim, UT, in 1969 to 1970, 590 established

(71.2 percent). Five years later 421 plants were vigorous and growing. Luke and Monsen (1984) reported

high success in transplanting sagebrush on a Wyoming

mine spoil site. Individual big sagebrush plants can

be propagated by the rooting of stem cuttings

(Alvarez-Cordero and McKell 1979). Tissue culture

propagation is also possible (Neville and McArthur

1986).

Big sagebrush can be successfully drill (fig. 26, 27),

or broadcast seeded (fig. 28). Care should be taken that

seeds are not covered too deeply, 1/8 inch (3.2 mm) or

less is preferred (Monsen and Richardson 1984;

Richardson and others 1986; Young and Evans 1986b).

Seeding should be done in the fall. Planting in subsoil

substrates has had mixed success. Luke and Monsen

(1984) and Monsen and Richardson (1984) were not

successful, but Stevens and others (1981b) successfully seeded big sagebrush in subsoil scalps (fig. 29).

Figure 26—Drill seeded big sagebrush with

grass mixture in southern Idaho. Sagebrush

seed was separated from grass seed in seed

boxes and drill rows.
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Chapter 21 Composite Shrubs

Matching seed source with planting site habitat is

important for emergence and survival of big sagebrush (Meyer and Monsen 1991). In general, regardless

of subspecies, big sagebrush seed germination is

adapted to climate of collection sites (Meyer and others 1990b). A wilt disease can be troublesome in

nurseries (Nelson and Krebill 1981). Symptoms of

this disease are present also in natural populations.

Snowmold can also cause lack of vigor and death in

sagebrush (Nelson and Sturgess 1986; Sturgess and

Nelson 1986). During cold winters with little snow

cover, winter injury and death has been documented

for large stands of sagebrush (Nelson and Tiernan

1983).

Uses and Management

Big sagebrush is one of the more nutritious shrubs

on Western winter livestock and game ranges. Basin

big sagebrush has higher protein levels than the other

subspecies (Welch and McArthur 1979b). Palatability

of the different populations of this shrub to mule deer

and other animals varies widely. For deer, basin big

sagebrush is generally less palatable than Wyoming

big sagebrush; both are generally less palatable than

mountain big sagebrush (Hanks and others 1973;

Scholl and others 1977; Sheehy and Winward 1981;

Wambolt 1996; Welch and McArthur 1986; Welch

and others 1981). However, Striby and others (1982)

Figure 27—Seeded big sagebrush in subsoil

scalps in an intermediate wheatgrass seeding, Millard County, UT.

Figure 28—Plant establishment from aerial seeding of big sagebrush with crested wheatgrass,

alfalfa, and Utah sweetvetch in south-central Utah.

reported that Wyoming big sagebrush was most palatable to deer and elk at their Montana study site.

Wyoming big sagebrush is generally most palatable

to sheep (Welch and others 1987). Palatability of

spicate big sagebrush has not been studied to any

great extent. It is browsed by sheep in the late summer

and early fall and by big game when available in

winter. Palatability of xeric big sagebrush is good for

mule deer and sheep. Sagebrush is important food and

cover for upland birds. For example, sagebrush comprised 62 percent of the annual diet of sage-grouse in

Montana (Wallestad and others 1975). Big sagebrush

is an important component of antelope diet. Olsen and

Hansen (1977) found sagebrush comprised 78 percent

of the annual diet for antelope in Wyoming’s Red

Desert. Even though sagebrush is a valuable forage, it

is generally a less preferred browse for deer and other

Figure 29—An even-topped mountain big

sagebrush.
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Chapter 21 Composite Shrubs

browsers than many rosaceous shrubs such as the

mountain mahoganies, bitterbrush, and cliffrose

(McArthur and others 1978b; Smith and Hubbard

1954; Smith and others 1965). Because of its widespread abundance, its ability to grow with associated

grasses, forbs, and other shrubs and its nutritious

nature, big sagebrush is the most important winter

forage in foothill areas throughout much of the West

for livestock and big game. Sustained heavy use can,

however, result in loss of vigor and mortality of big

sagebrush and other sagebrush species (Cook and

Child 1971; McArthur and others 1988a; Smith 1949).

It is one of the best shrubs available for use in revegetation of depleted winter game ranges in the Intermountain area (Plummer 1974a; Welch 1983a).

Meyer and colleagues showed that sagebrush seeds

are site-adapted. Generally seeds from the area to be

planted or a similar area will establish and perform

better (Meyer 1994; Meyer and Monsen 1992; Meyer

and others 1990b).

All forms of big sagebrush usually recover by naturally seeding sites that have been sprayed, burned,

disked, or chained. Natural spread can be reduced if

soils are plowed or deeply disked causing seeds to be

deeply buried. Mountain big sagebrush recovers well

even amid the presence of established herbs. However,

mountain big sagebrush usually grows on sites that

receive high amounts of summer moisture to sustain

seedling growth. In addition, this sagebrush is able to

produce good seed crops each year. Under more arid

conditions, natural seed production is more erratic.

Poor seed crops often occur and seed viability is diminished after 2 to 3 years of storage (Stevens and others

1981a).

Big sagebrush is aggressive and persistent and

sometimes forms closed stands. These stands may

require thinning and rejuvenation. These techniques

are treated in chapter 17, and in Plummer and others

(1955), Pechanec and others (1965), Laycock (1979),

Utah State University (1979) (especially Keller’s contribution), Marion and others (1986), and Whisenant

(1986b). Big sagebrush stands are unexcelled in providing ground cover and forage when grazed to maintain a balance between the sagebrush and associated

herbs and shrubs. Because big sagebrush has the

potential to establish rapidly from both transplanting

and direct seeding, it is useful for stabilizing washes,

gullies, roadcuts, and other raw, exposed sites (Clary

and Tiedemann 1984; McArthur and others 1979a;

Monsen and Richardson 1984) and planting depleted

game ranges (Plummer and others 1968). Big sagebrush, especially basin big sagebrush, has shown

promise as a living snowfence (Laycock and Shoop

1986). Winward (1980, 1983) reported that associated

species richness with each subspecies (see next section) varies greatly. Of the three common subspecies,

the order and quantity of diversified associated species is vaseyana > tridentata > wyomingensis. This

information should be kept in mind for carrying capacity and manipulative considerations.

Varieties and Ecotypes

Because of its plasticity and the apparent ease

with which it hybridizes, there are good opportunities for developing improved forms of big sagebrush

for different purposes (McArthur 1981; McArthur and

others 1979a, 1985, 1988b; Noller and McArthur 1986;

Welch and McArthur 1979a). One accession has been

released as a named germplasm; ‘Hobble Creek’ is a

vigorous mountain big sagebrush from near the

mouth of Hobble Creek Canyon, UT. It is palatable

to mule deer and sheep; it persists in relatively dry

sites receiving from 13 to 14 inches (33.0 to 35.6 cm) of

rain; and has good growth characteristics (Welch and

others 1986). Other accessions also show promise and

may qualify for later release.

Basin big sagebrush (A. tridentata ssp. tridentata)

is an erect, heavily branched, unevenly topped shrub

(fig. 24). Similar contrasting characteristics of basin

and other big sagebrush subspecies are given in

table 3. This subspecies has undivided or at least

trunklike, main stems. Most shrubs range between

3 to 6.5 ft (0.9 to 2.0 m) in height. Some forms, however,

may reach 15 ft (4.6 m) in suitable habitats. Mature

shrubs of this subspecies are the largest members of

the big sagebrush complex. The evergreen, vegetative

leaves are narrowly lanceolate, up to 5 cm long by 5 mm

wide, and typically three-toothed at the apex. The

leaves of the flowering stems gradually become smaller

and may be linear or oblanceolate and entire. Winward

(1970) found the average length-to-width ratio of persistent leaves is 4.6 to 5.6. The gray-canescent foliage

passes a strongly pungent, aromatic odor. Flowering

stems arise through the uneven crown and bear numerous flowerheads in erect, leafy panicles. The heads

contain three to six small yellowish or brownish,

trumpet-shaped, perfect disc flowers. The narrowly

campanulate involucre consists of canescent bracts

3 to 4 mm long and about 2 mm wide that form four to

five overlapping series around each head. The outermost bracts are less than a fourth as long as the

innermost bracts. Flowering occurs from late August

to October. Seed matures, depending on site, from

October to early December. Seed can be collected fairly

easily by beating (see chapters 20 and 24). Basin big

sagebrush is probably the most abundant shrub in

Western North America on lowland ranges. It normally occurs on dry, deep, well-drained soils on plains,

valleys, and foothills below 7,000 ft (2,100 m) elevation.

Above this elevation subspecies vaseyana and spiciformis, and occasionally subspecies wyomingensis are
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Chapter 21 Composite Shrubs

Figure 30—Wyoming big sagebrush growing

near Daniel, Sublette County, WY.

more prevalent. Vigorously growing basin big sagebrush is considered indicative of productive ranges

because it often grows in deep, fertile soil. This subspecies has generally been regarded as intolerant of

alkali, but there are ecotypes that grow in relatively

high alkalinity in association with such alkali-tolerant

plants as black greasewood, shadscale, and saltgrass.

A similar taxa restricted to southern California is ssp.

parishii distinguished by its exceptionally hairy

achenes. Subspecies parishii may only be an ecotype of

basin big sagebrush.

Mountain big sagebrush (A. t. ssp. vaseyana) is normally a smaller shrub than basin big sagebrush. Its

main stems are usually divided at or near the ground,

and it tends to have a spreading, evenly topped

crown (fig. 29). Vegetative branches are usually less

than 3 ft (91.4 cm) high and occasionally layer at the

base. Some lower elevation ecotypes reach about 6 ft

(1.8 m) in height. The persistent vegetative leaves are

broadly cuneate to spatulate and are characteristically wider than those of both basin and Wyoming big

sagebrushes. When looking down at shrubs of mountain big sagebrush, terminal leaves on each twig

appear to be distinctly whorled. Basin big sagebrush

does not show this trait, but Wyoming big sagebrush

shows the trait to some extent. Mountain big sagebrush grows in slightly acidic to slightly alkaline soils

(Welch and McArthur n.d.). Unlike ssp. tridentata,

ssp. vaseyana is rarely associated with any of the

saltbushes. Mountain big sagebrush has larger seeds

than Wyoming and basin big sagebrushes. It usually

comes better from direct seeding than the other two

subspecies. There are two varieties of mountain big

sagebrush. Each variety occurs in large distinct

populations but also integrates into other populations.

The most common variety outside of the Pacific Northwest is small-headed mountain big sagebrush

(Goodrich and others 1985). This is the sagebrush

that Ward (1953) referred to as that of “timbered or

mountainous areas in which the plants are very uniform in size, usually about 2 ft (61.0 cm) in height, and

of a rather spreading, flat-topped habit of growth, with

the inflorescence extending upward like plumes

above the rest of the bush.” The small-headed form has

four to six flowers per head. The other variety, var.

vaseyana, has seven to 11 flowers per head and has a

more restricted geographical range in the upper

elevational sagebrush areas of Washington, Oregon,

and Idaho with only occasional small patches occurring outside of that area (Goodrich and others 1985).

Rosentreter and Kelsey (1991), on Winward’s (1970)

suggestion maintained that xeric big sagebrush, (A. t.

ssp. xericensis) be recognized as a distinct entity. This

subspecies is apparently derived from hybridization

between ssp. tridentata and vaseyana (McArthur and

others 1979a; Rosentreter and Kelsey 1991). It is

adapted to dry sites between 2,600 and 4,600 ft (790 to

1,400 m) in southern Idaho and surrounding areas. Its

introgression with ssp. tridentata is apparent in that

plants are large and ragged topped, but chromatographically, cytologically, and phenologically it most

closely resembles ssp. vaseyana.

Wyoming big sagebrush (A. t. ssp. wyomingensis) is

somewhat intermediate in distribution, ecology, and

morphology between basin big sagebrush and mountain big sagebrush, but more diminutive than either

(fig. 30). It is adapted to drier sites than other subspecies by virtue of its rapid and long root growth and

early, but relatively small, aboveground growth (Booth

and others 1990; Welch and Jacobson 1988). The

accession ‘Gordon Creek’ has recently been released as

a named germplasm (Welch and others 1992b). It

combines wide adaptability, palatability, favorable

growth characteristics, and drought tolerance. Occasionally, the subspecies tridentata, vaseyana, and

wyomingensis may be found growing together.

Whenever it is found associated with ssp. tridentata,

ssp. wyomingensis is growing in the poorer, more

shallow soils (Beetle and Young 1965). Subspecies

wyomingensis is a low shrub usually less than 3 ft

(91.4 cm) in height. It has an uneven to round top with

flower stalks arising throughout the crown like ssp.

tridentata. Its main stem branches at or near the

ground level like ssp. vaseyana, but it does not layer.

Leaves are 1 to 2 cm long, narrowly cuneate to cuneate,

and have an average length-to-width ratio of about 3:1

for the persistent leaves (Winward 1970). Flowerheads

contain three to eight disc flowers and are arranged

into panicles narrower than the paniculate inflorescence of tridentata, and is wider than the spicate inflorescence of vaseyana(Beetle and Young 1965; Winward

and Tisdale 1977). Flowering and seed ripening take
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place later than vaseyana and earlier than tridentata.

Wyoming big sagebrush plants have an effective near

surface root system (Sturges 1977). This may be one

reason why relatively few plant species grow with it.

This subspecies may have arisen from hybridization

between ssp. tridentata and vaseyana (Beetle and

Young 1965; Hanks and others 1973) or ssp. tridentata

and A. nova (Winward 1975), or a combination of all

three taxa (McArthur 1983b). Wyoming big sagebrush

grows throughout the Intermountain region. It is

particularly abundant east of the Continental Divide

in Montana, Wyoming, and parts of Colorado in dry,

shallow, gravelly soil; usually from 5,000 to 7,000 ft

(1,500 to 2,100 m) (Beetle and Young 1965). In Idaho,

this subspecies is found from 2,500 to 6,500 ft (760 to

2,000 m) in the hotter, drier portions of the state

(Winward 1970).

Timberline big sagebrush (A. t.ssp. spiciformis) has

been the subject of some taxonomic confusion (see

A. rothrockii discussion in Goodrich and others 1985

and McArthur and others 1979a, 1981). This taxon is

the widespread root sprouting big sagebrush of high

elevations in the Intermountain area (Goodrich and

others 1985; McArthur and Goodrich 1986). Its habitat is often in openings in aspen, spruce, or fir woods

or where drifting snow accumulates (fig. 31). It had

formerly been confounded with Rothrock sagebrush

(Artemisia rothrockii), a similar taxon of the Sierra

Nevada (Goodrich and others 1985; Shultz 1983; Ward

1953). Rothrock sagebrush differs from timberline

sagebrush by its dark resinous leaves, restricted

California distribution, and high polyploidy. Timberline big sagebrush is thought to be a stabilized hybrid

between mountain big sagebrush and mountain silver

sagebrush (Goodrich and others 1985). In growth

form, timberline big sagebrush resembles mountain

Figure 31—Timberline big sagebrush growing

near Minturn, Eagle County, CO. Note the

large flower heads and flowering stalks.

big sagebrush in stem and leaf characteristics, but

mountain silver sagebrush in floral characteristics

(fig. 32). It has 10 to 18 flowers per head. Timberline

big sagebrush flowers in August and September in

native habitats, but in early June when transplanted

to low elevations (4,500 to 5,600 ft; 1,400 to 1,700 m).

It is a palatable sagebrush often browsed heavily in

the fall by sheep and mule deer. In winter it is usually

unavailable under snow. It has shown potential as a

low landscape hedge (McArthur and others 1979a).

Although basin, Wyoming, and mountain big sagebrush occur in distinct sites, all three of these common subspecies express a wide range of adaptation.

When planted throughout the big sagebrush, pinyonjuniper, and mountain brush communities all three

subspecies demonstrate adaptiveness. Although

sources from high elevations are not suited to arid

circumstances, and sources collected from light textured soils are not adapted to heavy clay textured

soils, most selections have established and persisted

when planted over a wide range of sites.

The subspecies of big sagebrush as well as near relatives (=subgenus Tridentatae) hybridize at many of

their contact points. In fact, hybridization is thought

to have been a key mechanism in the evolution and

speciation of the group (Beetle 1960; Hanks and others

1973; McArthur and others 1981, 1988; Ward 1953).

Artificial hybridization may be useful in selecting and

combining traits for management purposes such as

fire tolerance and palatability (McArthur and others

1979a, 1988b,c, 1998a; Weber and others 1994). Some

Figure 32—A specimen of timberline big sagebrush collected from Wolf Creek Summit, Wasatch

County, UT.
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Chapter 21 Composite Shrubs

current studies of big sagebrush zones of hybridization are yielding results in an effort to better understand the role of hybridization in nature and to help

determine the importance of the permanence and

function of hybrid zones. These studies have concentrated on morphological, chemical, physiological, cytological, ecological, edaphic, and entomological characteristics (Byrd 1997; Byrd and others 1999; Freeman

and others 1991, 1995, 1999; Graham and others

1995; McArthur and others 1988b,c, 1998a, 1999b;

McArthur and Sanderson 1999; Messina and others

1996; Wang 1996; Wang and others 1997, 1998,

1999).

Artemisia tripartita ______________

Threetip Sagebrush

Threetip sagebrush is a round, evergreen shrub up

to 6 ft (1.8 m) high. It may have a simple, trunklike

main stem or several branches arising from the base.

The bark on young branches is canescent but becomes

shredded and grayish, light brown to dark brown or

black on older stems. This species can layer, sometimes sprout back after a burn, and may sprout from

the stump following herbicide treatment (Beetle 1960;

Pechanec and others 1965; Schlatterer 1973; Winward

1980). Leaves of the vegetative branches are canescent, 0.5 to 4 cm long, and typically deeply divided

into three linear or narrowly linear-lanceolate lobes,

which in turn may be three-cleft (fig. 33). Some of the

upper leaves are often entire. Crushed foliage emits a

pungent odor. Flowerheads contain three to 11 disc

flowers and are normally arranged into panicles. Ray

flowers are lacking. Each head is subtended by eight

to 12 canescent involucral bracts. Achenes are resinous-granuliferous. Blooming occurs from July to

September and seed is collected from mid-October to

mid-December.

Ecological Relationships and Distribution

Threetip sagebrush covers approximately 13,000 mi2

(33,670 km2) in the Northern Rocky Mountains and

Great Basin States from British Columbia south

through Montana and Wyoming to Colorado and west

to Washington, Oregon, northern Nevada, and northern Utah at elevations between 3,000 to 9,000 ft

(910 to 2,700 m) (Beetle 1960). In some places, particularly in Idaho, this species occurs between the lower,

hot, dry sites dominated by Wyoming big sagebrush

and the higher, cooler sites dominated by mountain

big sagebrush (Schlatterer 1973). It usually grows on

moderate to deep, well-drained, loamy and sandy loam

soils (Winward 1980).

A B

Figure 33—Threetip sagebrush (Shultz 1986):

(A) A. tripartita ssp. rupicola; (B) A. tripartita

ssp. tripartita.

Uses and Management

Threetip sagebrush is a vigorous seeder, but unfortunately some forms are not particularly palatable.

However, Kufeld (1973) reported use by elk, and Kufeld

and others (1973) reported use by deer. Beetle (1960)

reported it was of low palatability for both livestock

and game, and Brunner (1972) observed that it was

never grazed in Nevada. However, a form near Salmon,

ID, is palatable to deer. The form near Salmon may

have been introgressed by a nearby population of

Wyoming big sagebrush (McArthur and others 1979a).

Stands of threetip sagebrush sometimes require

thinning by mechanical and chemical means

(Pechanec and others 1965).

Threetip sagebrush has not been difficult to establish by direct seeding, rearing, or transplanting. Usually a large amount of high quality seed is produced

annually. Seeds can be harvested without difficulty

from the upright bushes. Within its area of occurrence,

threetip sagebrush establishes and performs well.

Seedlings are able to persist within both a dense

perennial and annual understory. But threetip sagebrush is not any more competitive with cheatgrass

and other annuals than other species of sagebrush.

Plants grow quickly and can attain a mature stature

in 3 to 5 years. The shrub frequently exists with a

number of other shrubs and herbs. Seeding a mixture

of species usually does not reduce the survival or

productivity of this shrub.
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Attempts to establish threetip sagebrush onto arid

sites dominated by Wyoming big sagebrush have not

been successful. Plants initially appear but fail to

persist if reseeded. However, some ecotypes can be

extended to drier sites.

Threetip sagebrush recovers well following extensive disturbances. However, hot summer fires can kill

large stands and prevent resprouting and can significantly delay recovery by seeds.

Threetip sagebrush often grows intermixed, but

as separate stands, with mountain big sagebrush,

Wyoming big sagebrush, or alkali sagebrush. When

disturbances have occurred in these areas, threetip

sagebrush seedlings often invade into all circumstances. Natural sorting occurs, but rather slowly.

Threetip sagebrush demonstrates good compatibility

with many introduced and native grasses and broadleaf herbs. It survives and grows well with competitive

grasses including mountain brome, slender wheatgrass, intermediate wheatgrass, and orchardgrass.

Varieties and Ecotypes

Wyoming threetip sagebrush (Artemisia tripartita

ssp. rupicola) is a dwarf shrub with decumbent

branches that rarely grows over 6 inches (15.2 cm) tall.

It is frequently found layering and may have a crown

of 12 to 20 inches (30.5 to 50.8 cm) (Beetle 1960).

Leaves of the vegetative branches are often 3 cm long

and deeply divided into linear lobes, each at least 1 mm

wide (Beetle 1959, 1960). Flowerheads bear three to

11 disc flowers and are arranged into leafy, narrowly

racemose panicles. Flowers bloom in late August and

September. Seed ripens in October. Wyoming threetip

sagebrush has a rather limited range. It occurs on

rocky knolls from 7,000 to 9,000 ft (2,100 to 2,700 m)

in elevation in central and southeastern Wyoming

(Beetle 1960). Brunner (1972) reported this subspecies also occurs in southern Oregon, but had not yet

been found in Nevada. It typically grows on sites

adjacent to those of mountain big sagebrush.

Tall threetip sagebrush (A. t. ssp. tripartita) is a

freely branching shrub up to 6 ft (1.8 m) high. It can

layer easily when the conditions are right but is

seldom found layering in the field. After burning, it

may stump-sprout (Beetle 1960). Leaves of the vegetative branches are 1.5 to 4 cm long and deeply divided

into three linear lobes less than 1 mm wide. The lobes

may be further divided (Beetle 1959, 1960).

Flowerheads bear four to eight disc flowers and are

arranged into panicles that may sometimes be reduced to a spicate form. Flowers bloom in late August

and September. Seeds ripen in October. This subspecies occurs in dry, well drained soils at 3,000 to 7,500 ft

(910 to 2,300 m) elevation from British Columbia

south through Washington to northern Nevada and

eastward to northern Utah and western Montana.

Other Sagebrushes ______________

Coaltown sagebrush (Artemisia argillosa) is a threetip form endemic to a small area in Jackson County,

CO. This species is well adapted to heavy clay spoil

material and may have some utility in reclamation

activities (Beetle 1960; McArthur and others 1981).

Rothrock sagebrush (A. rothrockii) is similar in

many respects to subalpine big sagebrush and would

share the latter’s uses. However, Rothrock sagebrush

does not occur in the area of interest for this book. It is

confined to the high country of California (Goodrich

and others 1985; Shultz 1983).

Two other species—longleaf sage (A. longifolia)

and birdsfoot sage (A. pedatifida)—are not true sagebrushes in that they don’t belong to the subgenus

Tridentatae (table 2). Both, however, may prove useful

in revegetation efforts. Longleaf sage is the most

robust and woody member of the mostly herbaceous

Louisiana sage (A. ludoviciana) complex (see chapter

19 for treatment of the Louisiana sage or sagewort

complex). Longleaf sage may grow up to 3 ft (91.4 cm)

tall. It grows well on clay soils from Nebraska and

Wyoming north to Canada (Hall and Clements 1923).

Birdsfoot sage is a low perennial subshrub up to 6

inches (15.2 cm) tall with a tough woody root. It grows

on alkaline flats in Wyoming and Idaho and could be

used for rehabilitation of alkaline spoils (Hall and

Clements 1923).

General Rabbitbrush Culture ______

Table 4 presents distribution and differentation of

rabbitbrush taxa. Tueller and Payne (1987) and other

papers in Johnson (1987) provide additional information on rabbitbrush culture, management, and use.

The fruit of rabbitbrush is a spindle shaped achene

(seed). The terminal, or crown end, of each seed bears

a ring or crown of hairs, known as the pappus. Seed can

be wind and animal dispersed. Normally seed is produced annually. Seedlings naturally establish, especially on disturbed sites where competition is lacking

or slight. Thousands of seedlings can be found in bare

and disturbed soils, under and next to mature plants.

Most do not, however, develop beyond the seedling

stage. Seeds are relatively small (see chapter 20) with

considerable variation in size between taxa. White

rubber rabbitbrush seeds are among the smallest with

about 700,000 per lb (1.5 million/kg) (100 percent

purity). Mountain rubber rabbitbrush seeds are somewhat larger with just over 425,000 per lb (937,000 per kg).

Various rabbitbrush species have been successfully

seeded. Seeding success, based on successful establishment, has ranged from 0.001 to 4.0 percent (Stevens

and others 1986). The pappus has to be removed to

facilitate seed being seeded aerially, broadcast, or

drilled. Although seeding success theoretically could be
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Table 4—Species of Chrysothamnus with distribution, notes, and subspecific differentiation (drawn from Anderson 1986a; McArthur and Meyer

1987; McArthur and others 1979a; and references cited therein).

Section and Number of

species Distribution Notes subspecies

Chrysothamnus

C. albidus Scattered in alkaline soils W Utah to White-flowered very resinous —

EC California.

C. greeneia Often in sandy soils SC Colorado and N Mature plants have brittle white shiny stems 1

New Mexico to W Nevada and E California. Involuvral bracts are acuminate and recurved.

C. humilis Arid plains NW Nevada, NE California, Blooms earlier than rabbitbrush. —

SE Oregon, and isolated populations

further N in Oregon and Washington

C. linifolius Dry arroyos and other moist, deep but Large statured like rubber rabbitbrush. —

usually alkaline soil sites S Montana

to N Arizona and N New Mexico

C. viscidiflorus Broad distribution on dry open areas British Some populations have the only ray 5b

Columbia and Montana S to New Mexico, flowers in Chrysothamnus

Arizona, and E California; entire state of Utah.

Gramini

C. eremobius Known from only three populations in upland Related to genus Petradoria —

S Nevada.

C. gramineus Grows in mountain canyons in S Nevada Related to genus Petradoria. As opposed —

and adjacent California. to C. eremobius this species has longer

involucres, unkeeled phyllaries, and

glabrous achenes.

Nauseosi

C. nauseosus Broadly distributed in plains, foothills and Most common and widely distributed 22c

mountains British Columbia to Saskatchewan rabbitbrush with much diversity. Forms of

S to Western Texas, Sonora, and Baja ssp. albicaulis, hololeucus, and salicifolius

California. are preferred by browsing animals.

C. parryi Dry open places in mountains and foothills Shares stem tomentum character with

Wyoming and Western Nebraska W to C. nauseosus but has attenuate involucral 13d

California and S to New Mexico and Arizona. bracts as opposed to acute bracts of

C. nauseosus. Generally smaller and finer

structured than C. nauseosus.

Pulchelli

C. depressus Dry plains, hills, and rocky mountain slopes Dwarf shrub with conspicuously keeled, —

scattered over Western Colorado and New sharply ranked involucral bracts. Perhaps

Mexico W to SE California and S Nevada. the most palatable taxa to ruminants.

C. molestus Occurs only in a few scattered populations Rare species. Has glandular-hispid foliage. —

in uplands of NC Arizona.

C. pulchellus An undershrub in SW desert savannas SE Utah Has prominantly angled achenes. —

and SW in Utah, Kansas, to W Texas and N Coahuila.

C. vaseyi Scattered populations in mountain valleys and Resembles the more common C. viscidiflorus. —

foothills SE Wyoming to NC New Mexico W to

high plateaus of Utah.

Punctati

C. paniculatus Arid often stony slopes especially in dry arroyos Striate stems and terete leaves. —

SW Utah and SW Nevada to SC California.

C. teretifolius Arid stony slopes in S Nevada and SE California. Similar to C. paniculatis but more abundant. —

Has more strongly aligned involucral bracts

a

Formerly ssp. greenei and filifolius were recognized, but McArthur and others (1978a, 1979a) and Anderson (1986a) see no consistent distinctive differences between

the formerly recognized subspecies. b A total of 25 have been named at various taxonomic ranks. Hall and Clements (1923) recognized nine; Anderson (1986a) recognized five subspecies: ssp. axillaris,

lanceolatus, planifolius, puberulus, and viscidiflorus. c A total of 62 have been named at various taxonomic ranks. Hall and Clements (1923) recognized 20; Anderson (1986a) recognized 22 subspecies: ssp. albicaulis,

arenarius, bernardinus, bigelovii, ceruminosos, graveolens, hololeucus, iridis, junceus, latisquameus, leiospermus, mohavensis, nanus, nauseosus, nitidus, psilocarpus,

salicifolius, texensis, turbinatus, uintahensis, and wasinoensis. d

A total of 16 have been named at various taxonomic ranks. Hall and Clements (1923) recognized 10; Anderson (1986a) recognized 12 subspecies: ssp. affinis,

attenuatus, asper, howardii, imulus, latior, monocephalus, montanum, nevadensis, parryi, salmonensis, and vulcanicus.
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Chapter 21 Composite Shrubs

enhanced where the pappus is not removed (Stevens

and others 1986), there is presently no available equipment for seeding uncleaned seed.

Seeds are generally hand collected but can be harvested with various types of seed strippers, seed beaters, and vacuums. The pappus is removed by running

the seed through a barley debearder or hammermill

followed by screening and fanning. Seed is generally

collected between 10 and 15 percent purity. Seed

can be cleaned to 95 percent purity with persistent

screening and fanning. Barley debearding is preferred

over hammermilling. Seeds are somewhat brittle;

hammermilling can result in seed breakage and cracking in the balling of the collected material. Balling

hinders seeding.

Seed cleaned to less than 70 percent purity can only

be seeded aerially, by hand, or through a thimble

seeder. Seed cleaned to 70 percent or greater can be run

through a drill, seed dribbler, or browse seeder. Where

seeds are cleaned to high purities, they need to be

diluted with rice hulls or screened sawdust to facilitate seeding. Seed of most rabbitbrush species should

be surface seeded or covered not more than 1⁄32 inch

(0.8 mm) deep. If drilled, seed drop tubes and hoses

should be removed from between the disk furrow

opener and placed behind them. Seed will then be

deposited behind the disk furrow opener on a disturbed soil surface. Rabbitbrush seedlings are generally not competitive and may be suppressed by other

species. If drilled, thimble, dribbler, or browse seeder

seeded, rabbitbrush seed should be seeded individually or with less competitive species. Broadcasting

results in substantially less competition between species. Late fall and early spring seeding are preferred.

Rabbitbrush seed can be stored in an open, unheated,

uncooled warehouse for up to 3 years following collection without too much loss of viability.

Chrysothamnus depressus _______

Dwarf Rabbitbrush

Dwarf rabbitbrush is a small, irregularly branched,

depressed shrub or subshrub 1 ft (30.5 cm) high or less,

with numerous short branches arising from decumbent lower stems (fig. 34). The branches are covered

with a dense scabrid pubescence. The narrowly oblanceolate to spatulate leaves are 0.7 to 2 cm long, 1 to

4 mm wide, and finely puberulent or scabrous like the

branches. The heads contain five disc flowers that

are arranged into compact terminal cymes. Involucral

bracts have mucromate to attenuate tips, are 10 to

13 mm long, and are arranged into five distinct

vertical ranks. Achenes are 5 to 5.5 mm long and

glabrous or obscurely pubescent toward their apex.

The soft brownish-white pappus is slightly longer than

the corolla. Blooming occurs from May to October; seed

matures from October to December (see chapter 20).

This species occurs on dry plains, hills, and rocky

mountain slopes from 3,300 to 6,900 ft (1,000 to 2,100

m) in elevations scattered over western Colorado, New

Mexico, Utah, Nevada, Arizona, and southeastern

California (Anderson 1986a).

Dwarf rabbitbrush is often heavily browsed by sheep,

cattle, and wildlife (Hall and Clements 1923; Kufeld

and others 1973). It transplants readily, can be seeded,

and is useful for stabilizing depleted soils on which it

readily grows. This handsome shrub is a source of

protein when grasses and broadleaf herbs have dried

(McArthur and others 1979a).

Chrysothamnus linifolius _________

Spreading Rabbitbrush

Spreading rabbitbrush is a tall, robust shrub up to

7 ft (2.1 m) high that spreads underground (fig. 35) by

lateral roots that form adventitious shoots (Anderson

1964). Of all the rabbitbrushes only Parry rabbitbrush

shares the underground spreading trait and then not

nearly to the extent spreading rabbitbrush does. Leaves

are large, flat, green, glabrous, lanceolate to oblonglanceolate, 2 to 5 cm long, and 8 to 16 mm wide. Heads

contain four to six, usually five, disc flowers and are

arranged into broad, loose cymes. The involucral bracts

have obtuse tips with thickened green spots similar to

those of Vasey rabbitbrush. These spots, however, dry

to a brown color. Achenes are covered with dense, long,

soft hair. Blooming occurs during August and September with seed ripening from October to mid-December.

Spreading rabbitbrush occurs in Montana, Wyoming,

Colorado, New Mexico, Utah, and Arizona. It is most

abundant in alkaline soils along roadcuts, barrow pits,

ditches, streams, and washes in the Upper Colorado

River drainage.

Figure 34—Dwarf rabbitbrush growing near the

mouth of Ephraim Canyon, Sanpete County, UT.
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The species has aggressive underground spreading

characteristics coupled with an ability to spread by

seed that make it a valuable stabilizer of disturbed

alkaline soils (McArthur and others 1974; 1979a;

Plummer 1977). It transplants readily. Laycock and

Shoop (1986) found spreading rabbitbrush to be an

excellent living snowfence for the central Great Plains.

Spreading rabbitbrush receives little browsing use. It

does, however, provide year round cover for upland

game and small birds.

Chrysothamnus nauseosus _______

Rubber Rabbitbrush

Rubber rabbitbrush is a shrub usually 12 to 80

inches (0.3 to 2.0 m) high, but it varies from dwarf

forms to types over 10 ft (3.0 m) high (fig. 36). Usually,

several erect stems arise from the base and these

branch to form rounded bushes. Branches are covered

with a green, yellow-green, gray-green to white feltlike

tomentum usually infiltrated with a resinous gum, making the plant somewhat sticky. This coating is often

mistaken for part of the bark but can be discerned by

scraping with a knife edge or a fingernail. Leaves are

nearly filiform in some subspecies to broadly linear in

others. Leaves vary from 18 to 63 mm long and are

covered with a tomentose vestiture. They are not

twisted or gland-dotted. The heads of this species are

usually arranged into a cymose inflorescence. Each

head bears 20 to 25 glabrous to densely tomentose

involucral bracts arranged in up to five vertical rows.

Rubber rabbitbrush blooms from August to October

and are among the latest bloomers of the genus. Plants

at higher elevations bloom earlier than those at lower

elevations. Seeds mature, depending on location and

subspecies, and are collected from mid-October until

Figure 35—Root sprouts, foreground, of spreading rabbitbrush at the Snow Field Station.Figure 36—Large rubber rabbitbrush growing at

Palmetto, Esmeralda County, NV. The specimen

pictured is the viridulus form of ssp. consimilis.

the end of the year with over 500,000 cleaned seeds

per lb (1.1 million/kg) (see chapter 24). Rubber rabbitbrush is normally a heavy seed producer (fig. 37).

Ecological Relationships and

Distribution ____________________

We have recently discovered that galls induced by

tephritid flies are differentially distributed among the

subspecies of rubber rabbitbrush (McArthur 1986;

McArthur and others 1979b—See Varieties and

Ecotypes section later in this section for subspecies

descriptions). Over much of their range, the whitestemmed subspecies albicaulis and subspecies

hololeucus are infested with a persistent round stem

gall, 0.3 to 1.2 cm in diameter. This gall was found

restricted to these white-stemmed subspecies in our

observations. Green subspecies consimilis, graveolens,

Figure 37—Rubber rabbitbrush in seed, southcentral Utah.
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Chapter 21 Composite Shrubs

and turbinatus, on the other hand, have a less persistent, fluffy stem gall reminiscent of a ball of cotton

about 0.7 to 1.4 cm in diameter. This gall is found also

on ssp. hololeucus in southern areas near Kanab, UT,

and Colorado City, AZ. The round gall is absent in

these areas. In areas where these green-stemmed and

white-stemmed rubber rabbitbrush subspecies occur

together the galls are specific. We have observed only

a few cases of cross gall inoculation and then only on

putative hybrid plants. Wangberg (1981) independently observed some species and subspecies specificity of tephritid galls on rabbitbrush in Idaho. He found

less specificity for the round gall in Idaho than we did

in Utah. He identified the tephritid flies that induce

the galls as two different species of Aciurina. Dodson

and George (1986) confirmed the genetic and ecological integrity of the two Aciurina species.

Rubber rabbitbrush ranges from British Columbia

to Saskatchewan south to western Texas, Sonora,

Baja California, and eastern California (Anderson

1986a, c). It is a common plant on plains, valleys, and

foothills. It grows best in openings with the sagebrush,

juniper-pinyon, and ponderosa pine zones in sandy,

gravelly, or clay-alkaline soils. This species grows

at elevations ranging from 500 to 9,000 ft (150 to

2,700 m). Rubber rabbitbrush is a vigorous early

invader of disturbed sites such as roadcuts and overgrazed rangelands. On ranges where big sagebrush

has been destroyed by fire, insects, vehicular traffic, or

continued heavy grazing, rabbitbrush will increase

and often become the dominant vegetation (Evans and

others 1973; Rosentreter 1986). Nevertheless, in most

habitats, this species is not overly competitive with

herbaceous species, and on some sites, it does not

suppress grass. Production of herbaceous cover percentage have been notably greater (fig. 38) when

rabbitbrush is present than when it is not present

(Frischknecht 1963; Plummer 1959; Plummer and

others 1968). We find it of interest that rubber rabbitbrush has a high photosynthetic rate, not unlike

desert-adapted C4 photosynthetic pathway species

(Davis and others 1985).

Plant Culture

Rubber rabbitbrush can easily be transplanted in

the fall or preferably in the spring to avoid frost

heaving so long as the plants are physiologically inactive and moisture conditions are right and the transplants are small (10 inches tall; 25.4 cm or less). A less

successful method—which, nevertheless, yields some

results—is to divide dormant crowns (Hall and

Clements 1923). Tissue culture techniques have been

successfully employed to propagate rubber rabbitbrush (Upadhyaya and others 1985). Success was

obtained from direct seeding, especially when seeds

have been distributed on top of disturbed soil in the fall

Figure 38—Rubber rabbitbrush growing in association with smooth brome, Sanpete County,

UT. Note vigor of grass under shrub canopy.

or early winter. Once established, natural spread can

occur (Stevens 1986b).

Seed germination of rubber rabbitbrush is usually

quite high. Seeds maintain high viability (65 percent)

for 3 years under ordinary warehouse storage conditions (Stevens and others 1981a). With cool night

(40.5 ∞F; 4.7 ∞C) and warm day temperatures, rubber

rabbitbrush seed germinates in about 2 days (Weber

and others 1985). However, under standardized single

cool temperature germination conditions, seeds from

lower elevations and latitudes germinate in about

half the time (2 weeks) that it takes for seeds from

higher elevations and more northerly areas to germinate (McArthur and others 1987a; Meyer and McArthur

1987; Meyer and others 1989). Stevens and others

(1986) have shown that seeding rabbitbrush is more

successful when the pappus is left on the achene (the

fruit that holds rabbitbrush seed) and the achenes are

placed upright at the soil surface. Achene size in

rubber rabbitbrush subspecies appears to be correlated to habitat; subspecies adapted to sandy sites

have larger achenes (Meyer 1997).

Uses and Management

Rubber rabbitbrush is an excellent plant for controlling erosion because of its deep roots, heavy litter,

and ability to establish on severe sites (Aldon and Pase

1981; Luke and Monsen 1984; Monsen and Richardson

1984; Monsen and Stevens 1987; USDA Forest Service

1937). Once established, this species reproduces

easily and spreads fast from its light, plumed, winddisseminated achenes (Young and others 1984b). It

also grows vigorously when transplanted. The value of
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C. nauseosus as browse varies greatly between subspecies and ecotypes. The white to grayish subspecies

such as albicaulis, hololeucus, and salicifolius are

more palatable to livestock and big game than the

green subspecies, graveolens and consimilis (fig. 39)

(Hanks and others 1975; McArthur and others 1974).

Throughout much of the summer range, game and

livestock browse the plant lightly, if at all, except

under unusual conditions. During late summer and

fall when rubber rabbitbrush is in bloom, most livestock and game graze the flowers and occasionally a

few leaves and the more tender stems. Rubber

rabbitbrush is most heavily browsed on winter ranges

(fig. 40). Rubber rabbitbrush was found in 48 percent

of the deer stomachs examined on a portion of winter

range in Owens Valley, Inyo County, CA; however, it

never amounted to more than 6 percent of the total

food ingested (Sampson and Jesperson 1963). Crude

protein content ranged from 9 percent during the

dormant months to 11.8 percent in the spring after

new leaves had formed (Sampson and Jesperson 1963).

Antelope also make extensive use of rubber rabbitbrush (Yoakum 1986). Bhat and others (1990) found a

wide range of winter nutritive quality in rubber rabbitbrush accessions growing in a uniform garden.

Rubber rabbitbrush is resilient to browsing. Severely

browsed plants often obtain full regrowth by midsummer following heavy browsing. Rubber rabbitbrush

also provides cover for small mammals and birds.

The potential for this species to produce rubber and

other chemicals has long been recognized (Hall and

Figure 39—Differential winter-spring utilization of rubber rabbitbrush subspecies by deer

in Sanpete County, UT. Left, C. nauseosus

ssp. consimilis (10 percent use), right, C.

nauseosus ssp. hololeucus (50 percent use).

Figure 40—Heavy use of rubber rabbitbrush by

deer in the winter at an experimental plot in

Sanpete County, UT: (A) C. nauseosus spp.

salifolius plant in the late fall; (B) The same

plant after winter browsing. The stake in “A” is

1 ft tall; the mark on the stake in “B” is at 1 ft.

Goodspeed 1919). There has been a recent revival of

interest in this potential (Hegerhorst and others 1987;

Ostler and others 1986; Weber and others 1985). Some

accessions produce up to 6 percent stem rubber and

20 percent resin content. The plant also has other

possible industrial and horticultural uses (Weber and

others 1985).

Rubber rabbitbrush can become troublesome on

ranges when it invades and occupies disturbed lands

at high densities. It is a difficult plant to control

because of its resistance to herbicides and its crown

sprouting nature (Evans and others 1973; Whisenant

1986b, 1987; Young and others 1976a).

A

B
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Chapter 21 Composite Shrubs

Varieties and Ecotypes

Anderson (1986a,c) recognized some 22 subspecies

of rubber rabbitbrush. Often, two or more subspecies

will occur sympatrically. Each subspecies is essentially inbred and morphologically clean (Anderson

1986c; McArthur and Meyer 1987; McArthur and

others 1978a). However, reproductive isolation occasionally breaks down, especially at the limits of subspecies geographical ranges, and intermediate morphological forms are found (Anderson 1986c; Hanks

and others 1975; McArthur and others 1978a). Because of the mostly inbreeding, but potential for outbreeding, system of rubber rabbitbrush, many local

reproductively isolated populations of rubber rabbitbrush occur. We agree with Hall and Clements (1923)

that “improvement in any desired direction may be

brought about by selection, or by hybridization or by

both of these methods.” To date, no artificial improvement has been made, but individual populations

with desirable characteristics such as high rubber

content, palatability, or growth form have been identified. We discuss below the more common subspecies

of rubber rabbitbrush.

White stem rubber rabbitbrush—C. nauseosus ssp.

albicaulis (mountain), and hololeucus (basin)—are

shrubs from 2 to 6.5 ft (0.6 to 2.0 m) high with erect,

leafy branches and leaves covered with a permanent,

dense, white to grayish tomentum (fig. 41). The leaves

are 2.5 to 4 cm long, 0.5 to 1.5 mm wide or sometimes

to 3 mm wide in a few forms. The strongly keeled, acute

involucral bracts are white, more or less tomentose,

8 to 10 mm long, and arranged into five distinct

vertical ranks. The yellow corollas are 8 to 11 m long

and terminate in lobes 1 to 2 mm long. The achenes are

densely pubescent. The subspecies differ in the length

of their corolla lobes (ssp. albicaulis 1 to 2 mm long,

ssp. hololeucus 0.5 to 1 mm long) and style appendage

lengths for which ssp. albicaulis has style appendages

longer than stigmatic portions, but ssp. hololeucus

does not (Anderson 1986a). These subspecies tend to

overlap morphologically in their area of sympatry

(northern Utah, northeastern and western Nevada,

southwestern Idaho, and southeastern California;

Anderson 1986a,c). In these areas of sympatry, subspecies albicaulis occurs at higher elevations (over

6,000 ft; 1,800 m) whereas ssp. hololeucus is confined

to lower elevations. White rubber rabbitbrush subspecies are common and widespread and are found in

open places in plains, foothills, and mountains from

British Columbia, Alberta, and Montana southward

to northwestern Colorado, Utah, Nevada, and southern

California (Anderson 1986c). Subspecies hololeucus

grows in the Great Basin area whereas ssp. albicaulis

occurs on mountains and foothills, east, north, and

west fringes of the Great Basin and beyond (Anderson

1986a,c; Hall and Clements 1923). Subspecies

Figure 41—Four subspecies of rubber rabbitbrush: (A) C. nauseosus ssp. albicaulis; (B) C.

nauseosus consimilis; (C) C. nauseosus ssp.

graveolens; (D) C. nauseosus ssp. salicifolius

(McArthur and others 1979a).

hololeucus is often intermixed with ssp. graveolens

on foothill ranges and with subspecies consimilis in

valleys and plains. The striking white forms of both

subspecies have potential use as ornamentals

(McArthur and others 1979a; Weber and others 1985).

Another common white subspecies of the foothills and

plains of Colorado, Wyoming, Montana, and the western Dakoatas is ssp. nauseosus. This subspecies is

short statured, about 2 ft (61.0 cm) tall, and quite

palatable to game and livestock.

Threadleaf rubber rabbitbrush (C. n. ssp. consimilis)

may reach 10 ft (3.0 m) in height when mature. It has

leafy, erect branches covered with a green to yellowgreen tomentum. The narrow threadlike (linearfiliform) leaves are less than 1 mm wide and 2.5 to 5 cm

long (fig. 41). They are usually covered with a green to

yellow-green tomentum and are somewhat resinous.

The involucral bracts are acute, glabrous, and keeled

and are arranged in fairly distinct vertical rows. The

bracts are 6.5 to 8.5 mm long. The corollas are 7 to 9.5

mm long, with glabrous lobes 1 to 2.5 mm long. The

achenes are densely pubescent, suggesting that this

subspecies may be a connecting link between rubber

rabbitbrush and low rabbitbrush. Subspecies consimilis is most common in alkaline valleys and plains

of the Great Basin where it is often associated with

A

B

C

D
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Chapter 21 Composite Shrubs

various saltbushes and black greasewood. In less

alkaline areas, consimilis intermixes with ssp.

hololeucus and graveolens. Threadleaf rubber rabbitbrush also occurs in alkaline soil outside the Great

Basin from western Wyoming, Colorado, and New

Mexico to northeastern Oregon and eastern California.

A large ecotype, up to 12 ft (3.7 m) tall with treelike

trunks, occurring principally in the western parts of

its range, was formerly recognized as a separate subspecies (viridulus) (fig. 36). Perhaps ssp. viridulus

should be maintained as it is higher in rubber content

and in resins than ssp. consimilis (Ostler and others

1986). Another subspecies, pinifolius, has also been

reduced to be synonymous with consimilis (Anderson

1986a). Plants formerly referred to as pinifolius grow

in southern Colorado and New Mexico. Threadleaf

rubber rabbitbrush is one of the least palatable of the

C. nauseosus subspecies. It may, therefore, have value

in revegetating disturbed sites such as roadcuts where

attraction of browsing animals is not desired. This

subspecies helps control erosion on open alkaline soils

by providing ground cover and soil stabilization

(McArthur and others 1979a).

Green rubber rabbitbrush (C. n. ssp. graveolens)

ranges from 2 to 5 ft (0.6 to 1.5 m) high when mature.

Its leafy, erect branches are yellow-green to green or

sometimes gray-green and are covered with a compact

tomentum (fig. 41). The linear leaves are 1 to 3 mm

wide, 4 to 6 cm long, and only slightly pubescent. The

involucral bracts are 6 to 8 mm long, glabrous at least

on their backs, acute, keeled, and arranged in vertical

rows. Achenes are densely pubescent. Green rubber

rabbitbrush is widespread and sporadic from North

Dakota to Idaho and southward to western Texas,

New Mexico, and Arizona. It is most common on welldrained foothills but also extends up into the mountains and down into valleys and plains where it is

often intermixed with ssp. consimilis. Green rubber

rabbitbrush is generally less palatable than the white

or gray subspecies, hololeucus, albicaulis, and

salicifolius. Nevertheless, some forms of this subspecies have been found that are utilized to a moderate

degree by livestock and mule deer.

Mountain rubber rabbitbrush (C. n. ssp. salicifolius)

is a shrub from 1 to 6.5 ft (0.3 to 2.0 m) high. Its

ascending to erect twigs are leafy and covered with a

gray-green, fairly compact tomentum. The leaves are

broadly linear, ranging from 4 to 8 cm long and 3 to

10 mm wide, which makes them the largest leaves of

the species (fig. 41). The involucral bracts are 7 to 8 mm

long, mostly obtuse, nearly glabrous, and arranged

in rather obscure ranks. The yellow corollas are about

1 cm long and have a minutely pubescent throat. The

achenes are densely pubescent. Hall and Clements

(1923) reported that mountain rubber rabbitbrush is

“apparently rare and confined to Utah.” However,

McArthur and others (1979a) found it to be fairly

widespread at higher elevations in Utah and perhaps

extending to Nevada as part of the lower subalpine

vegetation. It extends down to the foothills in parts

of its range where it may be found intermixed with

ssp. albicaulis, hololeucus, and graveolens (Plummer

1977). Mountain rubber rabbitbrush appears to be

the most palatable subspecies for both livestock and

big game of the five common subspecies, salicifolius,

albicaulis, hololeucus, graveolens, and consimilis in

Utah, and it is often browsed heavily in the fall

(McArthur and others 1979a).

Other less common subspecies with locally large

populations include arenarius, junceus, leiospermus,

and turbinatus. Subspecies arenarius is a striking

white shrub adapted to sand dunes in the Four Corners area. It has large heads up to 20 mm long. Subspecies junceus is a nearly leafless, yellow-green form

adapted to sandy areas in the Colorado River drainage. Subspecies leiospermus is a low shrub usually

less than a meter in height. It has affinities to ssp.

consimilis. It occurs mostly on arid, rocky sites in the

southern half of the Intermountain area. A distinguishing feature of ssp. leiospermus is its glabrous or

nearly glabrous achenes. Subspecies turbinatus is a

yellow form adapted to sandy sites, mostly in western

Utah. It has large floral heads. A few additional

subspecies occur principally outside of the Intermountain area. These include bigelovii (mostly south and

west of the Four Corners area), nauseosus (mostly on

the western edge of the northern Great Plains), and

mohavensis (southern California) (Anderson 1986a,c;

McArthur and Meyer 1987; McArthur and others

1979a).

Chrysothamnus parryi ___________

Parry Rabbitbrush

Parry rabbitbrush (Chrysothamnus parryi) is a

shrubby species somewhat intermediate in height,

stem and leaf tomentum, and growth habit between

rubber rabbitbrush and low rabbitbrush. Parry rabbitbrush is a low, dense shrub similar in habit to

certain forms of low rabbitbrush (fig. 42). It is usually

from 8 to 24 inches (20.3 to 61.0 cm) in height with

numerous spreading to erect flexible branches. Like

rubber rabbitbrush, the branches of Parry rabbitbrush are covered with a feltlike white to green tomentum. The tomentum is neither as dense nor resinous

as in rubber rabbitbrush. Paulsen and Miller (1968)

and McArthur and others (1979a) observed that Parry

rabbitbrush spread from underground roots. The

glabrous to tomentose, somewhat viscid leaves are

narrowly linear to elliptic and range in size from 0.5 to

8 mm wide and 1 to 8 cm long. Flowerheads usually are

arranged in terminal leafy racemes that sometimes

form panicles. The involucral bracts are 9 to 14 mm
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Figure 42—Parry rabbitbrush (C. parryi ssp.

howardi) growing at Current Creek, Duchesne

County, UT.

high and terminate in acuminate to very attenuate

herbaceous tips. The yellow, tubular to funnelform

corollas are 8 to 11 mm long. Achenes are 5 to 6 mm

long and are covered with long, shaggy, appressed

hairs. Blooming occurs from July to September; seed

matures in October and November.

Ecological Relationships and Distribution

Parry rabbitbrush occurs in dry, open places in

mountains and foothills of Western North America

from Wyoming and western Nebraska west to

California and south to New Mexico and Arizona.

Like other species of rabbitbrush, this species tends

to increase on overgrazed and disturbed areas.

Populations of Parry rabbitbrush are usually smaller

and more scattered than those of the other common

species (rubber and low rabbitbrushes) of rabbitbrush.

Varieties and Ecotypes

Parry rabbitbrush is a diverse widespread group,

with 12 subspecies (Anderson 1986a; McArthur and

Meyer 1987).

Chrysothamnus parryi ssp. asper is a low shrub

6 inches (15.2 cm) or more high with slightly spreading to erect branches. Its green leaves, roughened

with short-stalked resin glands, are 2 to 5 cm long and

1 to 3 mm wide. The heads contain five to 10 disc

flowers and are subtended by somewhat ranked involucral bracts with straight tips. This subspecies

occurs on mountain sides bordering desert areas

from 6,900 to 8,500 ft (2,100 to 2,600 m) in elevation in

western Nevada and eastern California (Anderson

1986a; McArthur and others 1979a).

Chrysothamnus parryi ssp. attenuatus consists of

low shrubs with mostly erect stems up to 2 ft (61.0 cm)

high. It has green, slightly viscid, narrowly linear

leaves, 2 to 4 mm long and about 1 mm wide. The

leaves are erect but are not larger than the inflorescence. Heads contain five to seven disc flowers and

are subtended by involucral bracts with slender,

straight tips. The bracts are ranked into five vertical

rows. Blooming occurs from August to October. Subspecies attenuatus is found in the sagebrush, pinyonjuniper, and yellow pine vegetation types in Utah and

southwestern Colorado and northwestern New Mexico

(Anderson 1986a; McArthur and others 1979a).

Chrysothamnus parryi ssp. howardii is a low

shrub. Its spreading basal stems and erect branches

are up to 2 ft (61.0 cm) high. The narrowly linear,

tomentose leaves are 2 to 4 cm long, about 1 mm wide,

and the upper ones usually extend beyond the uppermost heads of the inflorescence. Flowerheads contain five to seven pale yellow disc flowers. The heads

are subtended by vertically ranked involucral bracts

usually with spreading tips. Blooming occurs from

July to September. The subspecies occurs on dry hills

and mesas associated with sagebrush, pinyon-juniper,

and ponderosa pine vegetational types in Utah, southern Wyoming, Colorado, New Mexico, and Nebraska

(Anderson 1986a; McArthur and others 1979a).

Chrysothamnus parryi ssp. monocephalus is a low

shrub from 4 to 24 inches (10.2 to 61.0 cm) high with

rigid, spreading branches. Its viscid, somewhat

tomentose leaves are linear-oblanceolate or spatulate,

1 to 3 cm long and 1.5 cm or less wide. The upper leaves

usually extend beyond the inflorescence. The flowering heads occur singly or in pairs on the end of short,

leafy branches. The heads contain five to six disc

flowers and are subtended by obscurely ranked involucral bracts with straight, attenuate tips. This

subspecies occurs in the high mountains of western

Nevada and eastern California at elevations between

2,600 and 11,000 ft (790 and 3,400 m) (McArthur and

others 1979a).

Nevada rabbitbrush (Chrysothamnus parryi ssp.

nevadensis) consists of low shrubs with ascending

to erect branches up to 2 ft (61.0 cm) high. The linear

to linear-oblanceolate leaves are 1.5 to 4 cm long,

0.5 to 3 mm wide, and sometimes green to resinous,

but usually gray-tomentose. The uppermost leaves

rarely extend beyond the inflorescence. The flowering

heads contain four to six yellow disc flowers and are

subtended by ranked involucral bracts with slender

recurved tips. Chrysothamnus parryi ssp. nevadensis

occurs between 4,300 and 8,900 ft (1,300 and 2,700 m)

in elevation on dry mountain sides from eastern

California to eastern Nevada, southwestern Utah,

and northern Arizona. It is most common along the

eastern flank of the Sierra Nevada (Anderson 1986a;

McArthur and others 1979a).
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Chapter 21 Composite Shrubs

Chrysothamnus parryi ssp. parryi consists of low

shrubs with erect branches 1 to 2.5 ft (30.5 to 76.2 cm)

high. The uppermost leaves usually extend beyond

the inflorescence. Flowering heads contain 10 to 20 disc

flowers and are subtended by obscurely ranked involucral bracts with straight attenuate tips. It blooms

during August and September. Subspecies parryigrows

in dry plains, valleys, and hillsides in central Nevada,

southern Utah, south-central Wyoming, western Colorado, and northern New Mexico (Anderson 1986a;

McArthur and others 1979a).

A few other subspecies of C. parryi, such as affinis,

glandulosa, imulus, and vulcanicus, occur outside of

or only on the fringe of the Intermountain area and in

small numbers (Anderson 1986a; Hall and Clements

1923). No efforts have been made to improve Parry

rabbitbrush. The species contains abundant genetic

variation. Its ssp. howardii is thought to be involved

in the parentage of C. nauseosus ssp. utahensis

(Anderson 1984).

Chrysothamnus vaseyi ___________

Vasey Rabbitbrush

Vasey rabbitbrush is a low, rounded shrub with

ascending to erect branches up to 1 ft (30.5 cm) high.

The bark on the young branches is pale green to

whitish and glabrous, becoming brown and fibrous

with age. Leaves are linear to linear-oblanceolate, 1 to

2.5 cm long, 1 to 3 mm wide, and glabrous. Heads contain

five to seven disc flowers each and are arranged into

small, compact cymes. The obscurely ranked involucral bracts are 5 to 7 mm high, oblong, obtuse to

rounded, and all but the innermost have a thickened

greenish spot near the apex. The achenes are about

5 mm long, terete, longitudinally 10-striate, and glabrous. Blooming occurs from July to September; seed

matures from October until mid-December (table 1).

Vasey rabbitbrush occurs scattered over plains, hillsides, and mountain valleys at altitudes of 5,600 to

8,500 ft (1,700 to 2,600 m) mostly in Utah and Colorado,

but also in New Mexico and Wyoming (Anderson

1986a; Hall and Clements 1923).

This shrub is browsed by sheep but is small and so

scattered that it provides little forage (McArthur and

others 1979a).

Chrysothamnus viscidiflorus______

Low Rabbitbrush

Low rabbitbrush varies in appearance and foliage

characteristics. It is usually 1 to 3.5 ft (0.3 to 1.1 m)

tall with many erect stems branching from a simple

base (fig. 43). The brittle, erect twigs are glabrous or

puberulent with pale green or white bark. Leaves are

narrowly linear to oblong or lanceolate, 1 to 6 cm long

and often twisted. Leaf vestiture is glabrous or pubescent and commonly viscidulous with usually

scabrous margins. Degree of pubescence may vary

tremendously in variants of this species. As Hall and

Clements (1923) pointed out, sometimes the pubescence among the subspecies of low rabbitbrush “will

occur as a fairly dense though minute puberlence in

certain plants, while others almost exactly duplicating these in every other respect will be perfectly

glabrous.” Furthermore, L. C. Anderson (n.d.) has

observed good correlation between plant stature and

leaf size of ssp. visicidiflorus and lanceolatus with

altitude and amount of precipitation. Flowerheads

containing approximately five perfect, fertile disc

flowers each are arranged in compact terminal cymes.

Involucral bracts number about 15 per head and are

arranged in poorly to well defined vertical ranks. The

bracts of some subspecies have a greenish or brownish

thickened spot near their apex (McArthur and others

1979a). Flowering occurs from August through October; seed matures from October to the end of December. Low rabbitbrush produces about 3.4 million seeds

per lb (7.5 million/kg) of cleaned seed (see chapter 24).

This species has strong basal sprouting tendencies,

especially following top removal and injuries (Wasser

1982).

Ecological Relationships and Distribution

Low rabbitbrush is one of the most widely distributed shrubs on rangeland throughout Western North

America (Anderson 1986a,b; McArthur and Meyer

1987). The species has great ecological amplitude. It

occurs in dry, open areas from British Columbia and

Montana, south to New Mexico, Arizona, and eastern

Figure 43—Low rabbitbrush growing at the Snow

Field Station, Sanpete County, UT. This specimen is a mountain low rabbitbrush (C. viscidiflorus

ssp. lanceolatus).
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Chapter 21 Composite Shrubs

California at elevations between 2,600 and 11,000 ft

(790 and 3,400 m). Low rabbitbrush is usually associated with sagebrush, snakeweed, and other species of

rabbitbrush. Anderson (1986b) has shown that polyploid races in those subspecies where polyploidy occurs (ssp. lanceolatus, puberulus, viscidiflorus) are

adapted to lower and drier sites than their diploid

counterparts (Anderson 1986b).

Plant Culture

Low rabbitbrush can be transplanted in the fall

and spring. When transplanting in the fall consideration needs to be given to frost heaving potential.

Fall seeding is preferred; seed can be drilled or broadcast, and seed should not be covered more than 1/8 inch

(3.2 mm). Seeds can be harvested by hand and by

vacuum harvesting techniques.

Uses and Management

This shrub may provide an important supply of

browse to both game and livestock, particularly during late fall and winter after more desirable forage

has been consumed. Throughout the Great Basin, low

rabbitbrush, especially the flowering shoots, provide

good sheep feed. In California small amounts of low

rabbitbrush were found in deer stomachs examined

between October and January (Sampson and Jespersen

1963). Much variation exists in palatability among the

different subspecies. Some may be heavily utilized,

whereas others are consumed little if at all (McArthur

and others 1974). Variety stenophyllus of ssp.

viscidiflorus on rocky foothills is often heavily used,

and sometimes destructively so, with animals preferring mature or partially mature plants to green immature ones. On a Utah winter range, this subspecies

averaged to 11.31 percent by weight of the diet of

sheep (Cook and Harris 1950; Cook and others 1954).

On another Utah winter range, mule deer diets consisted of over 80 percent low rabbitbrush in December

(Austin and Urness 1983). Substantial use of ssp.

lanceolatus in widely scattered areas of Utah and

Nevada has been observed (McArthur and others

1979a). Antelope, elk, and bighorn sheep, as well as deer

and livestock, show varying preferences for low rabbitbrush, depending on season, locality, and subspecies

(Kufeld 1973; Kufeld and others 1973). The species,

like rubber rabbitbrush, increases rapidly and vigorously on otherwise disturbed sites. Some subspecies

such as stickyleaf low rabbitbrush (ssp. viscidiflorus)

and mountain low rabbitbrush (ssp. lanceolatus) adapt

well to higher elevations, while other subspecies such

as hairy low rabbitbrush (ssp. puberlus) do best in

lower desert and foothill habitats. Low rabbitbrush is

valuable for revegetating depleted rangelands and

other disturbed sites such as strip mines and roadsides (Plummer 1977).

Varieties and Ecotypes

This species includes five subspecies and several

ecotypes within subspecies (Anderson 1980, 1986b;

McArthur and Meyer 1987). Most important among

these in the Great Basin are two glabrous subspecies,

viscidiflorus and axillaris, and two pubescent subspecies, lanceolatus and puberulus. Anderson (1980)

reduced ssp. stenophyllus to varietal status under ssp.

viscidiflorus. Hall and Clements (1923) believe numerous intergrades have been held together in one

rather close species through interbreeding where

their ranges meet or overlap. Abrams and Ferris

(1960) describe C. viscidiflorus as a highly polymorphic

species composed of several freely intergrading subspecies of overlapping distribution. Intermediates are

invariably found in the field. We believe forms of low

rabbitbrush may be improved for grazing and other

uses through selection and breeding. Because each

subspecies is self-fertilized and predominately selfpollinated, each maintains its identify despite occasional outcrossing (McArthur and others 1978a).

Mountain low rabbitbrush (Chrysothamnus viscidiflorus ssp. lanceolatus) is a small shrub from 8 to

20 inches (20.3 to 50.8 cm) tall (fig. 43, 44). Its branches

are straw colored or gray and are finely pubescent.

Flowerheads are borne in small compact cypes with

densely pubescent branches. Involucral bracts are

5.9 to 6.5 mm long, lanceolate to oblong, obtuse, and

Figure 44—Three subspecies and a variety of

low rabbitbrush: (A) C. viscidiflorus ssp.

lanceolatus; (B) C. viscidiflorus ssp. publerus;

(C) C. viscidiflorus ssp. viscidiflorus var. stenophyllus; (D) C. viscidiflorus ssp. viscidiflorus.

A

B

C

D
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Chapter 21 Composite Shrubs

glabrous to pubescent. Achenes are densely strigose.

On the basis of his systematic investigations in the

genus Chrysothamnus, Anderson (1980) recommended placing the former ssp. elegans in synonymy

with C. viscidiflorus ssp. lanceolatus. Chromatographic

work by McArthur and others (1978a) supports this

consolidation. Mountain low rabbitbrush is widespread and fairly common in dry foothill and mountainous habitats from 5,000 to 10,500 ft (1,500 to

3,200 m) ranging from British Columbia, east to

Montana, and south to New Mexico, Utah, and Nevada.

This subspecies may be found growing with such

shrubs as big sagebrush, snakeweed, various subspecies of rubber rabbitbrush, Greene’s rabbitbrush

(C. greenei) and Parry rabbitbrush (C. parryi)

(McArthur and others 1979a).

Hairy low rabbitbrush (Chrysothamnus viscidiflorus ssp. puberlus) is a small shrub up to 20 inches

(50.8 cm) high with yellowish to green, finely pubescent branches (fig. 44). Its linear-filiform to linear

leaves are sparsely to densely pubescent with scabridciliate margins and are usually twisted or revolute.

The leaves are up to 2 mm wide and up to 3 cm long.

Flowerheads are borne in small compact cymes with

densely pubescent branches. Involucral bracts are

about 5 to 6 mm long, lanceolate to oblong, acute to

obtuse, and are usually marked with a thickened

greenish spot near their tips. Hairy low rabbitbrush

occurs on dry plains, valleys, and foothills, especially

on poorer soils and disturbed areas. Its range is essentially the Great Basin area from southern Idaho and

Oregon south through Utah and Nevada to eastern

California and northern Arizona (Anderson 1986a).

This subspecies is most abundant in the big sagebrush

communities of western Utah, Nevada, and southcentral Idaho. However, it has been found growing in

one locality or another with most of the other subspecies of low rabbitbrush, shadscale, winterfat, halogeton, and occasionally with pinyon and juniper

(McArthur and others 1979a).

Stickyleaf low rabbitbrush (Chrysothamnus viscidiflorus ssp. viscidiflorus) is the largest subspecies of

low rabbitbrush. Mature shrubs are usually more

than 20 inches (50.8 cm) tall, whereas the other subspecies are normally under 20 inches (50.8 cm). Its

branches, leaves, and inflorescences are glabrous but

viscid (sticky). The broadly linear to narrowly lanceolate, bright green leaves are 1 to 5 mm wide, 2 to 5 cm

long, and flat to twisted (fig. 44). Leaf margins are

sometimes scabrid. Crushed foliage usually emits a

pungent odor. Branches of the stems are glabrous.

Involucral bracts are obtuse, oblong, not keeled, and

5 to 7 mm long. Stickyleaf low rabbitbrush is widely

distributed on dry plains and hills from Washington,

Idaho, Montana south to Colorado, Utah, Nevada,

northern Arizona, and eastern California. It occurs

primarily in sagebrush and pinyon-juniper communities at elevations between 5,000 and 8,500 ft (1,500 and

2,600 m). This subspecies often becomes dominant in

cleared or overgrazed areas. At lower elevations

stickyleaf low rabbitbrush may be associated with

such halophytes as shadscale, winterfat, and halogeton. Other subspecies of low rabbitbrush are also

often associated with it.

Anderson (1971) recommends placing the former

ssp. pumilis in synonymy with C. viscidiflorus ssp.

viscidiflorus because its specimens are “only environmentally modified variants of C. v. ssp. viscidiflorus.”

Chromatographic work (McArthur and others 1978a)

supports Anderson’s reduction of pumilus to synonymy.

Because of its distribution from moist to arid sites, the

species is well suited to a wide range of disturbed sites

over the Western States (McArthur and others 1979a).

Recently Anderson (1980) reduced ssp. stenophyllus

to varietal status under ssp. viscidiflorus. This variety

is known as narrowleaf low rabbitbrush (fig. 44). It is

a low, glabrous shrub up to 12 inches (30.5 cm) high

with white bark. Leaves are linear-filiform, often

twisted, viscidulous, 1 mm or less wide, 1 to 3 cm long,

and glabrous except for the usually scabrid and revolute margins. The branches of the small, compact

cymes are glabrous. Involucral bracts are 4 to 6 cm

long, not keeled, and lance-oblong.

Narrowleaf low rabbitbrush is rather common on

many desert ranges, particularly in the northern

Great Basin and adjacent areas (Anderson 1980,

1986c). On these ranges this variety is usually found

in the sagebrush type on poorer soils and disturbed

sites, but is also found growing with halophytes such

as shadscale, fourwing saltbush, greasewood, and

halogeton.

Anderson (1980, 1986a) recognized two other subspecies of low rabbitbrush. These are ssp. axillaris and

planifolius. The former subspecies is morphologically

quite similar to var. stenophyllus, but occurs mostly

in the southern Great Basin and on the Colorado

Plateau and appears by its somewhat attenuate

bracts to be introgressed by Greene’s rabbitbrush. It is

quite abundant in certain locales. Subspecies

planifolius is a local form with small heads and flat

leaves endemic to Coconino County, AZ (Anderson

1986a).

Other Rabbitbrushes ____________

Alkali rabbitbrush (C. albidus) and Greene’s rabbitbrush (C. greenei) are two other relatively important

species. Alakli rabbitbrush is a much branched, leafy

shrub. It has erect, brittle, glabrous, very resinous,

whitebarked branches up to 3 ft (91.4 cm) high. The

glabrous, filiform leaves are 1.5 to 4 cm long, 0.5 to

2 mm wide, and the margins become revolute. Their
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Chapter 21 Composite Shrubs

surface is covered with small pits and abundant resinous exudate. Heads with four to six white disc flowers

each are arranged in small compact cymes. Each head

is subtended by approximately 15 glabrous, resinous,

involucral bracts. These are 7 to 9 mm long and

terminate in attenuate to acuminate, usually curved

tips. The pappus is abundant and longer than the

corollas. Mature achenes are about 4 mm long and

densely covered with long soft hairs. Blooming occurs

from August to November. This species is a definite

halophyte. It occurs most commonly along the western

side of the Great Salt Lake desert but is also found

across Nevada to eastern central California in alkaline soils. Alkali rabbitbrush may invade strongly

alkaline areas as a pioneer plant. In less alkaline flats

it may be associated with threadleaf rubber rabbitbrush, basin wildrye, and greasewood. This species

has value as ground cover on alkaline soils (McArthur

and others 1979a).

Greene’s rabbitbrush is a small highly branched

shrub only 4 to 14 inches (10.2 to 35.6 cm) high. Its

glabrous brittle twigs are green at first but soon

become white and shiny. Bark on the lower branches

often peels off in sheets (Hall and Clements 1923). The

nearly glabrous or slightly scabrous-ciliate leaves are

narrow, linear, 1.2 cm or less wide, 1 to 3.5 cm long, and

are more or less viscidulous. Flowerheads normally

contain five disc flowers in rounded or flat topped

cymes. The involucral bracts are 5 to 7 mm long,

arranged in five poorly defined vertical ranks, and

terminate in narrowly acuminate, greenish tips. The

tubular to funnelform corollas may be whitish or

yellow and 4 to 4.5 mm long. Achenes are about 3 mm

long and are covered with dense, long, shaggy hairs.

Greene’s rabbitbrush has been divided into two subspecies, ssp. greenei and ssp. filifolius by some authors (Hall and Clements 1923; Harrington 1964;

Kearney and Peebles 1960). These authors separate

ssp. filifolius from greenei by its normally larger stature and shorter narrower leaves. McArthur and others

(1978a) believe no subspecies should be recognized

because the purported subspecies are (1) chromatographically similar and (2) often occur in mixed populations. The chromatographic data from McArthur

and others (1978a) further suggest that C. greenei

could be considered a subspecies of C. viscidiflorus

rather than a separate species. Chrysothamnusgreenei

closely resembles C. viscidiflorus, particularly ssp.

axillaris and ssp. viscidiflorus var. stenophyllus. All

have a low bushy habit, whitebarked stems, and short

narrow leaves. Furthermore, ssp. axillaris has involucral bracts with attenuate tips (Anderson 1964) that

closely resemble those of C. greenei. We concur with

Hall and Clements (1923) who felt the similarity

between C. greenei and C. viscidiflorus indicated a

close genetic relationship. Greene’s rabbitbrush occurs

on plains, valley, and foothills in Colorado, New Mexico,

Nevada, and Utah. Overgrazing allows it to greatly

increase, sometimes forming a subclimax community

(Hall and Clements 1923). This species provides cover

and emergency browse in areas where it is abundant.

There are several other rabbitbrush species

(Anderson 1986a). These are all less common than

the species discussed or are outside of the geographical area of concern for this treatment. These

are C. gramineus and C. eremobius both related to the

genus Petradoria and of limited distribution in uplands of southern Nevada (and for C. gramineus,

adjacent California). Chrysothamnus molestus is a

rare, distinctive species from Coconino County, AZ.

Both C. pulchellus and C. spatulatus are distinctive

species that occur southeast of the main distribution of

the genus Chrysothamnus. Chrysothamnus humilus

is similar in some respects to low rabbitbrush, but is

distributed on the northwest fringe of the species distributional range. Both C. paniculatus and C. teretifolius

are viscid hot desert southwest shrubs. The former is

more common in gravelly washes whereas the latter

occurs mostly on rocky slopes.

Other Composite Shrubs _________

Haplopappus species (goldenweed),

Tetradymia species (horsebrush),

Gutierrezia species (matchbrush or

snakeweed), Lepidospartum

latisquamatum (scalebroom)

Both goldenweed and snakeweed belong to the

same tribe (Astereae) of Compositae that rabbitbrush

does (McArthur and others 1978a). Horsebrush and

scalebroom belong to the tribe Senecionieae. Haplopappus is a large genus that is mostly herbaceous. Its

constituent species are widely distributed but usually

scattered over Western rangelands (Hall 1928; USDA

Forest Service 1937). The most notable shrubs are

goldenweed rabbitbrush (Haplopappus bloomeri)

and its near relatives (Anderson 1983). Goldenweeds

are generally of little forage value, but they are of

value in erosion control (USDA Forest Service 1937).

Other shrubby goldenweeds include H. macronema,

H. suffruticosus, H. resinosus, H. greenei, and H.

carthamoides (Hitchcock and others 1955).

The genus Tetradymia consists of rather low, stiffly

branched shrubs (fig. 45). The stems are uniformly

canescent or have glabrous to woolly streaks running

down the stem internodes from the primary leaves.

The tomentum may be permanent or deciduous on

both the stem and leaves. Spines may be present

(fig. 46). Tetradymia bears primary and secondary

leaves. Primary leaves develop alternately along
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Chapter 21 Composite Shrubs

elongated shoots and are usually long-lived. Secondary leaves develop in fascicles in the axils of the

primary leaves. They are generally short-lived and

often dry up and fall away within a few weeks

(Strother 1974). Horsebrush flowers are borne in

heads located singly or in pairs in the upper primary

leaf or spine axils or are clustered in short, dense

racemes or corymbs at the tips of branches. Each head

contains from four to nine yellow disc flowers. Ray

flowers are lacking. Four to six equal involucral bracts

with overlapping sides subtend each head. The pappus consists of numerous bristles or scales or may be

lacking. The ovary of each flower develops into a

glabrous to densely longhaired achene.

This genus blooms from April to August depending

on elevation, climatic conditions, and species. Numerous small moths, bees, flies, and beetles visit the Figure 45— Gray horsebrush growing at Wasatch

Station, Summit County, UT.

Figure 46—Horsebrush species (Strother 1974): (A) Tetradymia canescens; (B) T. filifolia; (C) T. glabrata;

(D) T. nuttallii; (E) T. stenolepis; (F) T. argyraea; (G) T. tetrameres; (H) T. comosa; (I) T. spinosa; (J) T. axillaris

var. longispina.

A B C

H

E

F G

D

I J 5 cm
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Chapter 21 Composite Shrubs

Figure 47—Broom snakeweed, right, growing

next to hairy low rabbitbrush, left, near Ephraim,

Sanpete County, UT.

flowers. Although numerous potential pollinators are

available and the flowers are highly fertile, seedlings

are not commonly seen in nature. This is probably due

to the harsh environment in which horsebrush is

usually found (Strother 1974). However, young plants

and sprouts may be found growing in burned-over

areas (McArthur and others 1979a). Horsebrush provides some critically needed ground cover in the dry,

sparsely vegetated desert ranges where it grows. Although several species are poisonous to sheep and

have caused losses of thousands of animals in Utah

and Nevada, horsebrush is browsed particularly

heavily during winter and early spring on desert

ranges and overgrazed areas where little else may be

available this time of year. In fact, Zimmerman (1980)

has built a successful winter range cattle operation in

central Nevada on horsebrush and other desert shrub

species that are often considered useless. Severe losses

have occurred when hungry animals have been trailed

from winter to summer ranges through stands of

horsebrush without allowing the animals a chance to

graze other plants (Kingsbury 1964). Considerable

variation in toxicity within and between species of

Tetradymia has been noted by Johnson (1974a,b), who

believed ingesting black sagebrush and horsebrush

under certain poorly understood conditions is responsible for toxicity. Four horsebrush species (T. canescens,

T. glabrata, T. nuttallii, and T. spinosa) are quite

common in the Intermountain area. The first two

species are spineless, and the latter two are spiny

(McArthur and others 1979a). Tetradymia axillaris,

another spiny species, occurs on the edge of the Intermountain area in scattered populations from southwestern Utah to southern California. Two other species are found southwest of the Intermountain area:

T. argyraea (southeastern California) and T. comosa

(southern California and Baja California) (McArthur

and others 1979a; Strother 1974).

The broom snakeweeds (Gutierrezia formerly

Xanthocephalum) consist of mainly perennial herbs

and low suffrutescent shrubs (subshrubs) with woody

roots, crowns, and stem bases (fig. 47). A few species

are annuals (Lane 1985). However, of these only,

G. sphaerocephala occurs in the Intermountain area

(in New Mexico). These perennials are ordinarily

short-lived as well. Snakeweed leaves are entire, linear to narrowly oblanceolate, and usually sticky

from resin exuded to the surfaces of both leaves and

young stems. Numerous small heads form in loose or

crowded terminal clusters. Resinous, imbricated involucral bracts within thin membranous margins and

green tips subtend each head. A few yellow ray and

disc flowers are both present (fig. 48). The ray flowers

are usually pistillate and fertile, while the disc

flowers are usually perfect and fertile or sometimes

staminate. Pappi of several small scales or awns are

Figure 48—Broom snakeweed, right, with its

showy ray flowers, growing next to Parry rabbitbrush, left, in Salina Canyon, Sevier County, UT.

generally pubescent. Blooming occurs from May to

October. The numerous common names of this genus

are perhaps indicative of its wide distribution. The

genus consists of about 25 species scattered throughout western North and South America (Hitchcock and

others 1955; Lane 1985). Only two of these species,

G. sarothrae and G. microcephala, are of importance

in the Intermountain area although there are two less

common endemics, G. petradoria and G. pomariensis

(Welsh and others 1987). Some additional North

American species include G. bracteata, G. californica,

and G. serotina (Lane 1985; McArthur and others

1979a; Solbrig 1971; . Snakeweed commonly invades

depleted ranges and is considered an indicator of
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Chapter 21 Composite Shrubs

overgrazed rangelands. It does occur, usually in a low,

but fluctuating density, in ranges in good to excellent

condition. Species in this genus are generally unpalatable and seldom grazed. When eaten in quantity,

this plant is more or less poisonous to sheep and

goats (Benson and Darrow 1945; Kearney and Peebles

1960).

Scalebroom (Lepidospartum latisquamatum) is confined to sandy washes and gravelly plains in the

Mojave Desert extending to the southwestern portion

of the Intermountain area (Kay and others 1977e).

This plant superficially resembles rubber rabbitbrush

but is a member of the tribe Senecioneae and hence

has diagnostic morphological as well as chemical differences from rabbitbrush (McArthur and others

1978a). Scalebroom has potential use as a revegetation plant in areas where it is adapted (Graves and

others 1975; Kay and others 1977e). These authors

demonstrated that seed should be planted less than

1 cm deep. Furthermore, seed viability is lost rapidly

(in less than 2 years) if not stored in sealed containers.
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Chapter 21 Composite Shrubs
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 539

Nancy L. Shaw

Stephen B. Monsen

Richard Stevens

22

Chapter

Important shrubs of the Rose Family (Rosaceae) in the Intermountain region are distributed from blackbrush and salt desert

shrub communities through high elevation forests and meadows.

Growth habits of this group vary from trailing brambles to

upright shrubs and small trees. Some species are evergreen

while others are deciduous. Many of these species are highly

valued for the cover, fruits, and forage they provide for wildlife

and livestock. Rosaceous species that develop spreading root

systems, root suckers, rhizomes, or stem layers provide soil

stabilization. Several species are capable of regenerating following fire. Those that produce fragrant flowers or colorful fall

foliage are prized for their ornamental value.

Because of their browse value, antelope bitterbrush and several

other rosaceous shrubs were among the first species to be used in

wildlife habitat improvement efforts. Members of the bitterbrush-cliffrose complex as well as serviceberries, chokecherries,

and the mountain mahoganies lend themselves to artificial

seeding; their seed are fairly large and relatively easy to collect,

clean, store, and plant. Additional species of this family are

seeing increased use in response to the growing emphasis on use

of native species for wildland revegetation and low maintenance

landscaping, community restoration issues, mitigation for endangered species, and a general shift to employ revegetation

when deemed necessary to conserve or restore ecosystem diversity and functionality.

Rosaceous Shrubs
540 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Use of rosaceous species often presents challenges

to those involved in revegetation efforts. Size and

quality of seed crops vary considerably from year to

year. Complex germination requirements, low initial

growth rates, sensitivity to competition, and rodent

and bird preference for seed and seedlings of some

species hinder their establishment and reduce survival. Appropriate planting practices and management of young seedlings or plants are required to

minimize these problems. Control of browsing by livestock and wildlife during the first 2 to 3 years is often

essential to improve establishment and early growth.

Bareroot or containerized planting stock is commonly

used to shorten the period of seedling vulnerability.

Chapter Contents by Species

Amelanchier alnifolia (Saskatoon serviceberry) ................................................................................ 541

Amelanchier utahensis (Utah serviceberry) ....................................................................................... 543

Cercocarpus betuloides (Birchleaf mountain mahogany) .................................................................. 544

Cercocarpus intricatus (Littleleaf mountain mahogany) .................................................................. 545

Cercocarpus ledifolius (Curlleaf mountain mahogany) ..................................................................... 545

Cercocarpus montanus (True mountain mahogany).......................................................................... 550

Coleogyne ramosissima (Blackbrush) ................................................................................................. 552

Cotoneaster acutifolia (Peking cotoneaster) ....................................................................................... 555

Cowania stansburiana (Stansbury cliffrose or cliffrose) ................................................................... 556

Crataegus douglasii (Douglas or river hawthorn) ............................................................................. 560

Fallugia paradoxa (Apache plume) .................................................................................................... 561

Holodiscus discolor (Creambush oceanspray).................................................................................... 563

Holodiscus dumosus (Rockspirea) ...................................................................................................... 565

Peraphyllum ramosissimum (Indian apple) ....................................................................................... 566

Physocarpus malvaceous (Mallowleaf ninebark) ............................................................................... 567

Potentilla fruticosa (Shrubby cinquefoil) ............................................................................................ 569

Prunus americana (American plum) .................................................................................................. 571

Prunus andersonii (Desert almond or Anderson peachbrush) .......................................................... 572

Prunus besseyi (Bessey cherry) ........................................................................................................... 573

Prunus emarginata (Bitter cherry) ..................................................................................................... 574

Prunus fasciculata (Desert peachbrush) ............................................................................................ 575

Prunus spinosa (Blackthorn) .............................................................................................................. 577

Prunus tomentosa (Nanking cherry) .................................................................................................. 577

Prunus virginiana (Chokecherry) ....................................................................................................... 577

Purshia glandulosa (Desert bitterbrush) ........................................................................................... 579

Purshia tridentata (Antelope bitterbrush) ......................................................................................... 581

Rosa woodsii ultramontana (Woods rose) .......................................................................................... 586

Rubus leucodermis (Black raspberry, blackcap) ................................................................................ 590

Rubus parviflorus (Thimbleberry) ...................................................................................................... 591

Sorbus scopulina (Greene’s mountain ash) ........................................................................................ 592

Sorbus sitchensis (Sitka mountain ash) ............................................................................................. 594

Spiraea betulifolia (Bridal wreath) .................................................................................................... 594

Spiraea densiflora (Subalpine spiraea) .............................................................................................. 596

Spiraea douglasii (Douglas spiraea)................................................................................................... 596

Seedlings of native or introduced species of Prunus

and Rosa are traditionally grown by many nurseries

for conservation plantings. Antelope bitterbrush, Saskatoon serviceberry, and curlleaf mountain mahogany

or true mountain mahogany seedlings are commonly

grown for wildlife habitat improvement. Seedlings of

specific ecotypes and most other species must generally be obtained through contract production.

Characteristics of the site of origin of seed or planting stock should be matched to those of the planting

site as closely as possible; ecotypes of individual species vary considerably in range of adaptability. Although few rosaceous shrub cultivars have been selected and released for commercial production of seed
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 541

Chapter 22 Rosaceous Shrubs

and planting stock by the USDA Soil Conservation

Service (Hassell 1988), a number of others are under

study. Site requirements and characteristics of released cultivars have been extensively studied and

documented. Publications are available describing the

attributes and site requirements of commonly collected antelope bitterbrush populations (Tiedemann

and Johnson 1983). Verification of seed origin of

wildland collections may be accomplished by monitoring contract seed collections from designated native

sites or plantings. Seed certifying agencies have developed a “source identified” classification for wildland

seed certification that provides verification of site of

origin as well as compliance with minimum seed

quality standards (Young 1995).

Seeding recommendations for major vegetative types

and conditions are discussed in chapter 17. Rosaceous

shrubs adapted to these vegetative types and conditions are included in the seeding recommendations.

Seed characteristics are found in chapter 24.

The following sections include a brief species description and discussions of ecological relationships,

distribution, culture requirements, use, improved varieties, and management for each of these Rosaceae

species.

Amelanchier alnifolia ____________

Saskatoon Serviceberry

Saskatoon serviceberry is a deciduous shrub or

small tree 1 to 20 ft (0.3 to 6 m) or more in height

(fig. 1). The root system varies from a strong taproot

to a branching lateral root system. New shoots develop

from rhizomes extending up to 5 ft (1.5 m) from the

parent plant (Hemmer 1975). Branches are reddish

brown and pubescent when young, but later became

glabrous. Leaf shape is extremely variable. They are

approximately 1 inch (2.5 cm) long and oval to suborbicular with serrate or dentate margins to the middle

or below. Perfect flowers with five showy white petals

appear in large terminal clusters. Mature pomes are

purple to black and 0.4 to 0.6 inch (1.0 to 1.5 cm) in

diameter. Each pome contains 10 locules, each with

one seed; one third to one half are commonly abortive

(Blauer and others 1975; Brinkman 1974a; Harrington

1964). Seed are dark purplish-brown with a leathery

seed coat (Belcher 1985). Flowers bloom from May

through June and fruits ripen in July and August

(Dittberner and Olson 1983).

Ecological Relationships and Distribution

Three species of serviceberry are distributed through

the western United States. Saskatoon serviceberry,

dwarf Saskatoon serviceberry, and Utah serviceberry.

Although different geographic races have not been

identified, they are expected to occur within these

widely distributed species (Brinkman 1974a). In addition, several hybrids have been reported (Cruise 1964).

Saskatoon serviceberry is distributed from Alaska

and the Yukon east to Hudson’s Bay and south to

California, New Mexico, and Nebraska (Harrington

1964; Welsh and others 1987). Plants occur from near

sea level to over 9,000 ft (2,770 m) in the Rocky

Mountains in areas with an annual precipitation of

12 to 25 inches (30 to 64 cm) or more (Jones 1946).

Saskatoon serviceberry grows on sites ranging from

dry, rocky, exposed slopes to riparian areas with deep

moist soils. At the lower edge of its range, plants are

often intermixed with mountain big sagebrush/grass

and to a lesser extent with Wyoming big sagebrush/

grass communities (Tisdale and Hironaka 1981).

Saskatoon serviceberry also occurs in mountain brush,

upper pinyon-juniper, and ponderosa pine communities (Blauer and others 1975). It sometimes grows in

nearly pure stands, dominating other shrubs, but

supporting a productive understory.

Plant Culture

A few small nurseries or conservation plantings of

Saskatoon serviceberry are managed to furnish seed

for planting stock production. Most seed currently

used in artificial revegetation is harvested from wildland stands. Large quantities of seed can be harvested

at reasonable costs during good production years.

Some stands produce consistently good seed crops, but

harvestable yields usually occur at intervals ranging

from 1 to 5 years, depending on the site. Plants

usually produce numerous flowers each year, but

Figure 1—Saskatoon serviceberry occurs as

single plants, thickets, and nearly pure stands on

sites ranging from riparian areas to dry, rocky

slopes.
542 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

spring frosts, rust infestations, and insect damage

often diminish seed production (Blauer and others

1975). Seed is disseminated by gravity and by birds

and other animals that pass the seed through their

digestive tracts (Hemmer 1975).

Pomes are fleshy when mature and dry slowly on the

shrub. They are normally collected by hand picking or

by beating the shrub to dislodge them. Seed can easily

be separated from the fresh pomes. However, once the

fruits dry, the pericarp remains closely attached to the

seed. Dry pomes are macerated in a blender or Dybvig

to separate the seed from the dried pulp. Dried material

is then extracted from the seed by screening. For seed

purchases recommended purity is 95 percent and

germination 85 percent. There are about 45,395 seed

per lb (100,078 per kg) at 100 percent purity (see

chapter 24). Seeds have varying degrees of embryo

dormancy that can sometimes be overcome by wet

prechilling (Brinkman 1974a). Seed prechilled at 34

to 41 ∞F (1 to 5 ∞C) for 3 to 20 months germinated from

84 to 100 percent; the time requirement varied considerably among seedlots (Hargrave 1937; Heit 1968b;

McKeever 1938). The seed coats of some serviceberry

species also prevent germination (Young and Young

1986). Weber and others (1982) found acid scarification for 30 minutes and imbibition in a mixture of

thiourea and benzyladenine released this form of dormancy. These authors found no differences in germination among different sized seed. Some selections

from central Montana appear less dormant than many

other populations.

Saskatoon serviceberry should be seeded separately

from rapidly growing herbs. Plantings should be made

in strips or scalps at least 25 to 30 inches (63 to 76 cm)

wide. Seed may be drilled or broadcast seeded in a

firm, prepared seedbed and should be covered with

0.25 to 0.50 inch (0.64 to 1.25 cm) of soil. Broadcast

seeding without soil coverage is not recommended.

Rodents quickly gather seed, even those placed in soil.

Consequently, seeding in late fall or early winter after

rodent activity has diminished is advised (Brinkman

1974a; Plummer and others 1968)

Seedling growth is dramatically affected by available soil water. Survival may be low if seedlings are

subjected to drought during the first few months of

growth. Stressed plants are often stunted, and although they may persist through one or more growing

seasons, they usually do not recover. In contrast,

healthy seedlings that survive the first growing season are usually very tenacious.

Saskatoon serviceberry can be grown and field

planted as bareroot or container stock. Bareroot seedlings reach adequate size for outplanting in one season, developing a heavy taproot (Shaw 1984). Container seedlings are propagated from seed, germinants,

or small transplants and require a 3 to 4 month

cropping period (Landis and Simonich 1984). The

shrub is not easily propagated from cuttings (Doran

1957). Komissarov (1938) found rooting occurred in

25 percent of all softwood cuttings taken in midsummer following treatment with 50 mg/L indole-3-

acetic acid (IAA) for 24 hours. Untreated cuttings did

not root. Field survival of planting stock depends on

use of healthy dormant or properly hardened stock

and reduction of competition on the planting site.

Uses and Management

Saskatoon serviceberry is an important browse

plant for game and livestock due to its fair to high

palatability (Blauer and others 1975). It is capable of

persisting with a diverse and productive understory

that is important for summer grazing. Saskatoon

serviceberry is used by livestock, particularly sheep,

in midsummer after full leaf development and after

more palatable species have been browsed (Blauer and

others 1975).

Big game animals make considerable use of the

shrub (Kufeld 1973; Kufeld and others 1973). Tueller

(1979) found serviceberry was utilized by deer during

the entire year, but it was most important during the

midsummer and fall months. New shoots are particularly palatable, but deer consuming large quantities of

new serviceberry growth in spring can be poisoned by

it. Problems also occur if Saskatoon serviceberry comprises more than 35 percent of the fall or winter diet of

stressed deer (Quinton 1985). Birds utilize the bark,

fruits, and leaves. The fruit is a key grizzly bear food

(Mace and Bissell 1986; Zager 1980). Seed are used by

rodents and small birds (Blauer and others 1975).

Saskatoon serviceberry is particularly valuable as a

minor species in sagebrush communities on open hillsides and ridges. Use should not be allowed to reduce

plant productivity at these locations.

Saskatoon serviceberry is a useful shrub for landscape plantings in recreation areas, roadways, and

other low maintenance areas. It persists in areas that

receive heavy traffic and can withstand wind and

severe storms. It can also be used as a background

screen or for focal plantings.

Saskatoon serviceberry should not be planted off

site. Sources from mountain brush communities are

not well suited to lower elevations. The species has

not performed well on road or mine disturbances.

Developing seedlings do not survive if heavily grazed

or subjected to drought stress. However, established

plants, those 5 or more years of age, can withstand

rather heavy use, particularly winter browsing. In

fact, mature plants may be stimulated by some

grazing. Plant vigor can be maintained with 60 percent fall or winter use or 40 to 50 percent spring use

(Young and Payne 1948).

Saskatoon serviceberry seedlings develop slower

than antelope bitterbrush seedlings. Young seedlings

are seriously affected by grazing, and, unfortunately,
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Chapter 22 Rosaceous Shrubs

game animals tend to seek these plants. If not heavily

grazed, 4- to 5-year-old plants often begin to grow

relatively rapidly. Such plants usually require 8 to 12

years to reach mature size, but some plantings of

adapted ecotypes established on dry sagebrush sites

have taken as much as 20 years to mature. Production

of good seedcrops may not occur until shrubs reach

full size. Moderate grazing of established plants may

keep bushes hedged, but this usually does not cause

mortality.

Mature plants tend to persist for long periods.

Hemmer (1975) reported ages of 50 to 85 years for

serviceberry growing in climax brush fields. Saskatoon serviceberry can remain suppressed in closed

conifers stands for long time periods. Reducing canopy

cover stimulates resprouting (Wright and others 1979).

Regeneration from seed is normally rare (Noste and

Bushey 1987).

Saskatoon serviceberry is most vigorous in seral

communities. Productivity may increase following

burning or mechanical control in such situations.

However, where density is low or succession far advanced, such efforts may be counterproductive

(Hemmer 1975; Howard 1997). Plants recover well

following fires and survive even severe burns if the soil

is moist at the time of burning (Fischer and Clayton

1983). Tops may be killed, but plants can resprout

from rhizomes (Frischknecht and Plummer 1955).

Those rhizome buds located immediately below the

burned tissue sprout most readily (Bradley 1984).

Varieties and Ecotypes

Selection work has focused on comparing seed germination characteristics and site requirements of

selected ecotypes. To date, studies have not resulted in

identification of ecotypes unusually well suited to

restoration and revegetation work, although some

populations seem to produce consistently higher seed

fill. For this reason, use of local material on sites

previously occupied by Saskatoon serviceberry is

recommended.

Amelanchier utahensis ___________

Utah Serviceberry

Utah serviceberry is similar in growth habit to

Saskatoon serviceberry. Plants vary from large upright shrubs to small trees 4 to 15 ft (1.2 to 4.6 m) tall

(fig. 2). Twigs are unarmed. The alternate deciduous

leaves are simple, about 0.8 to 2.0 inches (2 to 5 cm)

long, and are borne on a short petiole (Harrington

1964). Leaves are suborbicular, oval, ovate or obovate

with coarsely serrate-dentate margins to the middle

or below. Flowers are clustered on racemes borne on

Figure 2—Utah serviceberry is a highly variable species that usually occurs on drier sites

than Saskatoon serviceberry.

short leafy branchlets. They are perfect and regular

with five white petals. The stamens are numerous

and inserted on the rim of the calyx tube. The ovary is

inferior with three or four styles. Each locule contains two ovules. Fruits are purple to black pomes

0.2 to 0.4 inch (6 to 10 mm) in diameter (Harrington

1964; Welsh and others 1987).

Utah serviceberry differs from Saskatoon serviceberry in having a somewhat smaller stature, smaller

leaves that are pubescent at least on the lower surface,

three to four styles compared to four or five, and a

hard, dry fruit. Davis (1952) considered Utah serviceberry a subspecies of Saskatoon serviceberry, but

Jones (1946), Blauer and others (1975), and Welsh and

others (1987) considered differences in fruiting and

growth habits sufficient to warrant species status.

Ecological Relationships and Distribution

Utah serviceberry occurs at scattered sites from

eastern Oregon to Montana and south to Texas,

California, Baja California and Sonora (Blauer and

others 1975; Welsh and others 1987). It is generally

found on drier sites than Saskatoon serviceberry. It

occurs on foothill sites at elevations from 2,000 to

7,000 ft (600 to 2,600 m) where it grows on dry, rocky

outcrops and on shallow soils. It is associated with a

wide number of plant communities including aspen,

ponderosa pine, pinyon-juniper, mountain brush,

and big sagebrush. Although it may occur in dense

stands, plants are usually scattered and intermixed

with other woody species.
544 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Plant Culture

Fruits of Utah serviceberry ripen in late summer,

but may remain on the bush for up to 2 years. Utah

serviceberry plants generally produce fewer fruits

than Saskatoon serviceberries; this may be due to its

occurrence on drier sites. However, some stands in

southern Utah’s Iron County often yield extremely

heavy crops. Seed are dispersed by birds and small

mammals (Brinkman 1974a).

The pomes are somewhat more difficult to dislodge

than those of Saskatoon serviceberry but can be collected in fairly large quantities during years of good

production. To obtain seed, the dry pomes are usually

placed in a blender or a Dybvig with water and macerated to separate the seed from the dry pulp. Seeds

are smaller and often have lower fill than those of

Saskatoon serviceberry, but germination characteristics are quite similar.For seed pruchases, recommended

purity is 85 percent and germination 95 percent.

There are about 25,800 seeds per lb (56,900 per kg) at

100 percent purity (see chapter 24).

Utah serviceberry is best established by drill seeding. Seed should be planted at a depth of 0.5 inch

(1.3 cm) in a weed-free seedbed. Hand seeding disturbed sites or seeding with interseeders or similar

implements that can selectively place seed in the soil

are also effective. Seeding at rates of 2 to 4 lb per acre

(2.2 to 4.4 kg per ha) is required to produce acceptable

stands. Seedlings establish slowly and are sensitive to

herbaceous competition. Bareroot or container seedlings are reasonably easy to produce, but they are also

highly sensitive to competition.

Uses and Management

Utah serviceberry is adapted to low foothill sites,

and may be used to improve diversity in sagebrush

communities. Selections of Utah serviceberry have

been seeded in relatively arid situations such as

pinyon-juniper and big sagebrush communities where

serviceberry is not native, in expectation that it may

successfully establish and improve wildlife cover.

Successful plantings in such areas have persisted for

approximately 30 years to date, but natural recruitment has not occurred. Saskatoon serviceberry has

failed to persist in such situations. Utah serviceberry

is resistant to rust (Gymnosporangium spp.) diseases

that often attack Saskatoon serviceberry (Krebill 1972;

Plummer and others 1968) and is occasionally selected

for planting where the rust is prevalent.

Slow establishment and poor survival have somewhat limited the use of both serviceberry species, but

Utah serviceberry establishes and grows faster than

Saskatoon serviceberry (Plummer and others 1968).

Seedlings fail to establish when seeded in poorly

prepared, weedy seedbeds. Young seedlings require

protection from grazing until plants have fully established, but mature shrubs withstand considerable

browsing. Some early plantings have been productive

for 40 years, although little spread has been noted

(Monsen n.d.). Neither serviceberry species establishes or grows well on highly disturbed sites such as

roadways or mine spoils.

Utah serviceberry provides good forage for cattle

and good to excellent browse for sheep and goats in

early spring. Deer browse it year long, but use is

particularly heavy in winter. Birds and small mammals use the berries in winter and spring (Blauer and

others 1975). Plants survive fires and regenerate by

sprouting (Tisdale and Hironaka 1981; Wright and

others 1979).

Varieties and Ecotypes

There are no releases. Local ecotypes are recommended for on-site planting.

Cercocarpus betuloides __________

Birchleaf Mountain Mahogany

Birchleaf mountain mahogany is an open, erect

shrub or small tree varying in height from 6 to 23 ft

(2 to 7 m). Although evergreen, it is similar in other

respects to true mountain mahogany (Welsh and

others 1987) (fig. 3). It occupies dry slopes and washes

below 6,000 ft (1,846 m) from central California to

southwestern Oregon (Munz and Keck 1959).

Figure 3—Birchleaf

mountain mahogany

stem with new leaves

and flowers, just after

petal fall.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 545

Chapter 22 Rosaceous Shrubs

Collections of birchleaf mountain mahogany from

central California have been planted within the

Intermountain region on game and livestock ranges

because of their evergreen growth habit, seedling

vigor, and drought tolerance. Healthy birchleaf

mountain mahogany seedlings that survive the first

growing season are usually very tenacious. However,

young plants are often stunted when stressed and

although they may persist for 1 or more years, they

usually do not recover. The seedlings and young plants

grow slightly faster than most selections of true

mountain mahogany for about the first 5 or 6 years.

After this time, growth rates and growth habits of the

two species are quite similar.

Cercocarpus intricatus ___________

Littleleaf Mountain Mahogany

Littleleaf mountain mahogany is an intricately

branched shrub usually less than 8 ft (2.4 m) tall

(fig. 4), with strongly revolute linear leaves (Blauer

and others 1975). It occurs primarily in Utah, Nevada,

and southern Arizona, occupying harsh rocky sites,

particularly areas with high summer temperatures

and infrequent rain storms. There are low-growing

forms of littleleaf mountain mahogany in Oregon and

Washington (Hitchcock and others 1961). Stutz (1974)

suggests that littleleaf mountain mahogany is a segregant of curlleaf mountain mahogany that tends to

be associated with more xeric sites. Seed germination

and cultural requirements for littleleaf mountain

mahogany and other mountain mahogany species

are quite similar.

Littleleaf mountain mahogany is productive and

evergreen with a low shrubby growth habit. It hybridizes with both curlleaf and true mountain mahogany

(Blauer and others 1975; Stutz 1974).

Figure 4—Littleleaf mountain mahogany is evergreen, low-growing, and generally very palatable.

Cercocarpus ledifolius ___________

Curlleaf Mountain Mahogany

Curlleaf mountain mahogany is an erect shrub or

small tree sometimes attaining heights of 22 to 26 ft

(7 to 8 m) (fig. 5) (Harrington 1964; Munz and Keck

1959). It has a rather deep, well developed root system

(Noste and Bushey 1987) and one to several main

trunks up to 7 inches (18 cm) in diameter with deeply

furrowed reddish-brown bark. The shiny leaves are

thick and evergreen, 0.4 to 1.2 inches (1 to 3 cm) long

and 0.2 to 0.4 inch (0.5 to 1 cm) wide. Leaves are

resinous and aromatic, entire, lanceolate, usually glabrous above and white tomentose below. Leaf margins are highly revolute, hence the name curlleaf.

Flowers are solitary or in twos or threes with white to

greenish-yellow sepals and no petals. The fruit is an

achene 0.3 to 0.4 inch (0.8 to 1.0 cm) long with an

attached plumose style elongating to 1.6 to 2.8 inches

(4 to 7 cm) at maturity. The flowering period extends

from May to June; fruits ripen from May to August

(Blauer and others 1975; Harrington 1964; Munz and

Keck 1959).

Figure 5—Curlleaf mountain mahogany, note highlining by deer.
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Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution

Curlleaf mountain mahogany is distributed east of

the Cascades and Sierra Nevadas from Washington to

Baja California and east to Montana, Colorado, and

Arizona. Curlleaf mountain mahogany occurs at elevations from 2,000 to 9,000 ft (615 to 2,770 m)

(Harrington 1964; USDA Forest Service 1937).

Curlleaf mountain mahogany is usually found on

warm, dry, rocky ridges on southern or western slopes

but may occur on all exposures (Brayton and Mooney

1966; Martin 1950; Miller 1964; Tidestrom 1925). At

lower elevations in the Great Basin, it often grows

on limestone, decomposed granite, or other coarsetextured soils on steep north slopes and among cliffs

and ledges (Dayton 1931). Plants may be nodulated by

the nitrogen-fixing endophyte Frankia (Lepper and

Fleschmen 1977).

Curlleaf mountain mahogany characteristically

grows in isolated patches that are nearly pure stands.

These sometimes cover considerable acreages and

often form a band between conifer forests and sagebrush or other shrub communities. Curlleaf mountain

mahogany is not reported as a dominant or associated

species of principal forest habitat types in Montana

(Pfister and others 1977), northern Idaho, or

Washington (Daubenmire and Daubenmire 1968).

However, the plant is a codominant with Douglas-fir

in some drier forested areas in Idaho (Steele and

others 1981). Dealy and others (1981) described five

communities dominated by curlleaf mountain mahogany in southeastern Oregon. These include curlleaf mountain mahogany/mountain big sagebrush/

bunchgrass, curlleaf mountain mahogany/mountain

snowberry/grass, curlleaf mountain mahogany/Idaho

fescue, curlleaf mountain mahogany/bearded

bluebunch wheatgrass, and curlleaf mountain mahogany/pinegrass. Johnson and Simon (1987) described

a curlleaf mountain mahogany plant community type

for the Wallowa-Snake Province of northeastern

Oregon. Curlleaf mountain mahogany is the dominant

understory in some limber pine, white fir, and juniper

series (Dealy and others 1981; Hironaka and others

1983; Steele and others 1981; Youngblood and Mauk

1985). It also grows in association with other woody

species such as pinyon, oak, serviceberry, bitterbrush,

or gooseberry (USDA Forest Service 1937). Curlleaf

mountain mahogany occasionally occurs with ponderosa pine where both species overlap, but these

sites generally do not form large or distinct communities. Extremely tall curlleaf mountain mahogany plants

occur with Douglas-fir and Rocky Mountain maple in

canyon bottoms and narrow draws or drainages in

central and southern Utah. These small communities

provide important habitat for big game and livestock.

In most situations where curlleaf mountain mahogany dominates, its crown cover ranges from 35 to

60 percent. Dealy and others (1981) counted 300 to 800

stems per acre (120 to 325 per ha) with an average of

500 per acre (200 per ha) in eastern Oregon. Somewhat

higher numbers were reported from Montana, the

average figure being 1,375 stems per acre (550 per ha)

(Duncan 1975). In Idaho, Scheldt (1969) reported a

mean of 207 shrubs per acre (84 per ha).

Three principal species of mountain mahogany

(Cercocarpus ledifolius, C. intricatus, and C. montanus)

occur in the Intermountain region (Harrington 1964;

Welsh 1982). Holmgren (1987) divided C. ledifolius

into two taxa: C. ledifolius var. ledifolius and var.

intermontanus (Intermountain curlleaf mountain

mahogany). Intermountain curlleaf mountain mahogany is distinguished from variety ledifolius by its

broader leaves that are less densely pubescent ventrally, its usually longer petioles and hypanthium

tubes, and more treelike habit. Stutz (1990) concurred

with Holmgren’s revision, concluding that the taxa

growing in the Intermountain West apparently arrived from different origins. The evolutionary processes that produced the existing species, particularly

littleleaf mountain mahogany are still proceeding.

All three hybridize in areas where they overlap

(Blauer and others 1975; Plummer and others 1957;

Pyrah 1964; Stutz 1974). The resulting hybrid populations are frequently important components of local

vegetation.

Intermountain curlleaf mountain mahogany occurs

in southeastern Washington, eastern Oregon, northern, eastern, and southern California, central and

southern Idaho, western Wyoming, Utah, and adjacent

parts of Colorado and Arizona (Holmgren 1987). It

often exists in pure stands surrounded by open sagebrush or mixed with Gambel oak, bigtooth maple,

pinyon-juniper, ponderosa pine, Douglas-fir, or white

pine at elevations of 4,300 to 9,800 ft (1,300 to 3,000 m).

Variety ledifolius occurs in eastern Oregon, central

and southern Idaho, southwestern Montana, northcentral Wyoming, and northern Utah (Holmgren 1987).

It grows on talus slides, rock outcrops and rocky slopes

at elevations from 1,300 to 6,900 ft (400 to 2,100 m).

Plant Culture

Stands of curlleaf mountain mahogany produce good

seed crops about in 2 of every 5 years. Few years occur

when little or no seed develops. However, to be profitable for hand harvesting, a heavy seed crop with good

viability must be produced; seed must ripen fairly

uniformly; and most seed must remain on the bush

for a period of time to permit a collection period.

Elevational range and aspect affect the period of

flowering and seed development. Seed are sometimes

dispersed rapidly by stormy weather conditions. Consequently, ripening seed should be checked periodically if possible, and harvested immediately when

mature. Ripe seed are normally wind dispersed and

can be carried some distance from the parent plant

(Deitschman and others 1974a). Once the achenes fall
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Chapter 22 Rosaceous Shrubs

to the ground, rodents begin gathering them; a high

percent of all viable seed may be removed within 1 or

2 days. Field mice remove embryos from the achenes.

Consequently, fallen achenes should be carefully examined before they are collected. Twisting of the

plume in response to changes in water content plants

the seed by drilling it into the ground (Dealy 1978).

Seed are harvested by beating the shrub to dislodge

the achenes and catching them in lightweight tarps or

netting placed down wind. Plumes can be removed

with a Dybvig or barley debearder. The material is

then screened with a fanning mill to separate the seed

from the remaining debris. If harvested lots contain

numerous small twigs, they must be separated over a

gravity table to improve purity. Seeds are not removed

from the achenes. The ripened seed are hard and

brittle and can be fractured or broken by improper

cleaning procedures, thus hammermilling should be

avoided. Purity of at least 90 percent and germination

of at least 80 percent are recommended for seed

purchases. There are about 51,865 seeds per lb

(114,342 per kg) at 100 percent purity (see chapter

24). Curlleaf mountain mahogany seeds remain viable

for extended periods in storage. Stevens and others

(1981a) found little loss of viability after 7 years of

warehouse storage. Nearly 85 percent of seeds stored

for 15 years germinated.

Freshly harvested seed is dormant and requires wet

prechilling (Dealy 1975; Deitschman and others 1974a).

Dealy (1975) found seed prechilled at 39 ∞F (4 ∞C) for

170 or 270 days germinated to 98 and 100 percent,

respectively. He concluded that the membrane surrounding the embryo may block gas diffusion and be a

principal factor in controlling dormancy. Young and

others (1978) found that soaking seed at 41 ∞F (5 ∞C) for

21 days in an aerated solution of potassium nitrate

and gibberellin improved germination over a range of

temperatures. Drying the seed narrowed the temperature range permitting germination, but increased

germinability was retained at these temperatures.

Curlleaf mountain mahogany fruit size varies

among ecotypes and collection sites. To date, seed size

has not been closely correlated with site conditions.

Differences may be due, in part, to genetic traits;

sources producing large fruits tend to perpetuate this

trait when planted on different sites.

The oblong achenes of curlleaf mountain mahogany

are large enough that they are easily handled and

planted alone or in mixtures with other shrub seed

using conventional drills. They may also be hand

seeded in selected spots. Curlleaf mountain mahogany

establishes best from fall seeding because of its wet

prechilling requirement and ability to germinate at

low temperatures (Dealy 1975; Plummer and others

1968). In addition, late fall or early winter seeding,

when rodents are less active, reduces seed predation.

Concentrating seedings in areas with low rodent

populations can greatly enhance seeding success.

Curlleaf mountain mahogany seedlings are not

competitive with exotic annuals, particularly

cheatgrass. Attempts to plant curlleaf mountain mahogany at lower elevations of mountain brush and big

sagebrush communities infested with these weeds

have generally not been successful. Proper seedbed

preparation and control of competing herbs are essential for shrub establishment. Phillips (1970) reported

high seed germination occurred on mineral soils cleared

of litter and plant competition.

Seedlings emerge early in spring following ground

thaw when soil water is likely to be favorable. However, seedlings are sensitive to late frosts. Dealy (1975)

found new seedlings exhibit rapid root growth in

relation to top growth, providing some resistance to

drought and codeveloping competition. However,

Scheldt and Tisdale (1970) did find heavy seedling

mortality occurred during the first summer of establishment, often resulting from drought and predation

by rodents and rabbits.

Curlleaf mountain mahogany can be propagated

and field planted as bareroot or container stock

(Butterfield and Tueller 1980; Landis and Simonich

1984; Shaw 1984). Seedlings develop rapidly in bareroot

nurseries. They must be lifted when dormant. Evergreen plants are particularly sensitive to lifting and

handling once growth has begun. Damage to the taproot should be avoided during lifting. Container stock

is easier to handle and often provides better establishment (Ferguson 1983; Landis and Simonich 1984).

Propagation of curlleaf mountain mahogany from

stem cuttings is difficult (Everett and others 1978).

Uses and Management

Because of its evergreen habit and occurrence between forest and shrub communities, curlleaf mountain

mahogany provides important cover and forage for a

wide variety of wildlife species. Curlleaf mountain

mahogany produces highly palatable winter browse

and thermal and hiding cover for elk, deer and other

big game (Dealy 1971, 1975; Kufeld and others 1973;

Mueggler and Stewart 1980; Smith 1952). Tueller

(1979) reported curlleaf mountain mahogany was one

of six shrub species that comprised the largest single

fraction of deer diets on winter ranges in Nevada. It is

also ranked as a principal winter browse for big game

in Utah (Richens 1967), Oregon, and Washington

(Mitchell 1951). Curlleaf mountain mahogany provides food, hiding cover, and nesting sites for many

birds (Dealy and others 1981).

Blauer and others (1975), Duncan (1975) and

Mueggler and Stewart (1980) reported livestock make

light use of mature curlleaf mountain mahogany

shrubs. However, seedlings and young stands can be
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Chapter 22 Rosaceous Shrubs

heavily browsed. Many healthy stands support a diverse herbaceous understory that provides good summer forage for sheep and cattle (Dealy and others

1981), but they are often inaccessible to livestock in

winter. Curlleaf mountain mahogany has good nutritive value (5.9 percent digestible protein) and digestibility (64.8 percent) in winter (Bissell and Strong

1955; Smith 1952; Welch 1981). It is one of only a few

shrubs to exceed the protein requirements of wintering mule deer; its leaves are evergreen and retain high

protein levels throughout the winter (Welch and

McArthur 1979b). Winter phosphorus level (0.18 percent) exceeds the lower range of phosphorus required

for wintering mule deer (Trout and Thiessen 1973;

Tueller 1979).

Curlleaf mountain mahogany is a useful species

for revegetating roadway and mine disturbances

(Hungerford 1984; Stark 1966) and it is an excellent

ornamental. It requires little maintenance, the leaves

are attractive and evergreen, and it withstands considerable traffic and browsing. Compared to other

wildland shrubs, it responds well to supplemental

watering. Curlleaf mountain mahogany wood is extremely dense as a result of its low growth rate. It has

been used for charcoal, fenceposts, and wood carving

(Blauer and others 1975). Stands are occasionally

logged for firewood.

Curlleaf mountain mahogany exhibits three characteristics that should be recognized and understood if

the species is to be used in revegetation projects. First,

seedlings can perform well on exposed sites if protected from competition. Second, young plants can

persist and furnish considerable high quality forage

even when moderately to heavily grazed. Under such

conditions plants will not reach mature stature and

could be weakened or killed if subjected to other forms

of stress. Third, although seedlings are not compatible

with herbaceous competition, established plants do

very well in conjunction with some highly productive

herbs. Curlleaf mountain mahogany could be more

widely planted with some herbaceous species to improve total and seasonal forage production if the shrubs

were planted in separate strips or spots from the

herbaceous species.

Many stands of curlleaf mountain mahogany exist

with little herbaceous understory. Shrub/grass

plantings established with intermediate wheatgrass

and other introduced herbs can be very productive.

However, the ability of the shrub to regenerate naturally under such conditions is reduced, particularly on

deep, productive soils. On shallow, rocky soils, where

weedy grasses and herbs have not gained dominance,

native forbs can usually be seeded along with the

shrubs.

Curlleaf mountain mahogany requires a longer period to attain mature stature than most other shrubs.

Plantings and stand rejuvenation treatments should

be extensive to reduce the impact of browsing during

establishment or recovery (Gruell and others 1985).

Plantings established in south central Idaho and protected from livestock and deer use grew relatively

slowly for about 6 years, but then grew dramatically

during the next 4 to 6 years. This growth pattern has

been noted in other protected study locations (Monsen

n.d.). Because the species is highly palatable (Mueggler

and Stewart 1980; Smith 1952), the naturally slowgrowing plants are continually browsed and, if not

protected, remain in a stunted condition, assuming a

rounded, hedged form (Blauer and others 1975; Garrison 1953). Plantings established in central Utah in

1950, for example, have survived continuous heavy

browsing, and the shrubs have remained quite small.

Once released from grazing, hedged plants can recover

quickly. Ferguson (1983) recommended browsing be

limited to 50 to 60 percent of current annual growth to

maintain productivity and a vigorous, shrubby growth

habit.

Many extensive stands of curlleaf mountain mahogany have been highlined. Most plants are mature

to decadent. New growth is out of reach of browsing

animals. Growth from lower branches diminishes as

a result of hedging and shading. Little or no reproduction occurs in these closed stands (Phillips 1970).

Various treatments have been investigated to regenerate them. Stands with live branches within reach of

browsing animals have been treated by top pruning in

Utah and Montana. Pruning in spring or early fall

prompted regrowth and increased forage availability

(Austin and Urness 1980; Garrison 1953; Ormiston

1978; Thompson 1970), but such treatments are expensive. Chaining or bulldozing kills most older or

highlined, single stemmed plants (Christensen and

others 1966; Dealy 1971), but can be used to open

areas within closed stands for natural regeneration or

planting.

Plants growing on shallow soils and exposed sites,

usually midwinter ranges for game, are difficult to

manage. Curlleaf mountain mahogany growing in

monotypic stands or in mixed stands with other shrubs

such as antelope bitterbrush, skunkbush sumac, and

green ephedra often receives heavy browsing year

after year. Most of these species are capable of withstanding considerable use, but may be weakened and

killed over time. Natural recovery is slow and often

prevented by continued use. Artificial revegetation is

also very difficult on these sites.

Curlleaf mountain mahogany plants are not highly

fire tolerant. The plants and onsite seed banks can be

killed by high intensity burns (Noste and Bushey

1987; Wright and others 1979). The thick lower bark

protects plants from low-intensity fires and although

the species is a weak sprouter, some surviving plants
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Chapter 22 Rosaceous Shrubs

can resprout from stem buds. In addition, new seedlings may emerge from unburned seed (Bushey and

Noste 1987; Gruell and others 1985). Some stands are

now restricted to rocky slopes due to high wildfire

frequency. In other areas, intense fire suppression and

understory reduction with grazing have permitted

curlleaf mountain mahogany to invade sites with

deeper soils (Dealy 1975; Gruell and others 1985).

Low intensity fires can be used to stimulate communities with low vigor or maintain curlleaf mountain

mahogany where it occurs as a seral species with

conifers (Gruell and others 1985). Burning can also

improve age class structure of curlleaf mountain mahogany stands. Natural recovery can occur following

prescribed fires, but it is often significantly enhanced

by mechanical site preparation that reduces herbaceous competition and creates a weed free seedbed

(Phillips 1970). Stands may also be rejuvenated by

thinning them to 40 to 60 clumps per acre (99 to

148 per ha) and mechanically clearing seedbed areas

within the stand. Protection of stands less than 50

years of age from fire is recommended if curlleaf

mountain mahogany is the climax species. Reduction

of browse use is essential for all stand improvement

projects.

The mountain mahoganies are relatively free from

infestation by insects and disease (Ferguson 1983).

However, serious outbreaks have destroyed sizeable

stands in southwestern Idaho. Furniss and Barr (1967)

reported the looper destroyed 46 percent of a 5,900

acre (14,580 ha) stand of curlleaf mountain mahogany

over 3 years. Tent caterpillars have been responsible

for severe defoliation in the Great Basin (Furniss and

Barr 1967).

Varieties and Ecotypes

Numerous accessions of curlleaf mountain mahogany

have been collected and evaluated for forage value,

seedling establishment, and vegetative growth characteristics. Attempts have been made to select and

develop ecotypes with improved seedling vigor and

high growth rates, but to date no major improvements

have been achieved. In general, selections appear

universally adapted to all areas where the species

occurs. However, it is possible that differences exist,

but have not yet been fully recognized.

Differentiation of the ancestral mountain mahogany

may have resulted in the appearance of some important traits. Stutz (1974) proposed that the highly

plastic true mountain mahogany, a plant better adapted

to mesic conditions, may have evolved from curlleaf

mountain mahogany. In contrast, the smaller, narrower leaves of littleleaf mountain mahogany may

have evolved as a response of curlleaf mountain

mahogany to more xeric circumstances. Because the

three species are closely related, each with chromosome number 2n = 18 (McArthur and Sandersen

1985), artificial hybridization is possible and could be

employed to incorporate specific traits into selected

populations.

Natural hybrids between Intermountain mountain

mahogany and true mountain mahogany and later

generation backcrosses are quite common where both

parents are found growing together (Blauer and others 1975; Plummer and others 1957). Hybrids are

somewhat fire tolerant, a characteristic of true

mountain mahogany. They often have a wider area of

adaptation than curlleaf mountain mahogany. Hybrids are easy to recognize; they are tall and treelike

with evergreen leaves and always occur in stands of

true mountain mahogany where true mountain mahogany is the female parent (Stutz 1990). These hybrids rarely reproduce. Few hybrids of C. ledifolius

var. ledifolius and true mountain mahogany have

been reported. This is perhaps due to the fact that

these taxa occur together only infrequently and hybrids are less easily recognized. They are highly fertile. Hybrids between littleleaf mountain mahogany

and true mountain mahogany are rare (Pyrah 1964),

perhaps due to reproductive barriers (Stutz 1990).

They are usually sterile. Hybrids between littleleaf

mountain mahogany and Intermountain mountain

mahogany are common in some areas where the species overlap (Blauer and others 1975; Plummer and

others 1957; Stutz 1990).

Cercocarpus ledifolius var. ledifolius is a small shrub

or tree occurring in the northern part of the Intermountain region, and is most common in Wyoming and

Montana (Cary 1917; Miller 1964). It also occurs in

Oregon, Idaho, California, Nevada, Utah, Colorado,

and Arizona. It is taller in stature than littleleaf mountain mahogany with larger leaves and flowers. Hybridization with curlleaf mountain mahogany is common.

Some C. ledifolius var. ledifolius populations express unique features of value in range and wildlife

habitat improvement. Ecotypes occurring in eastern

Wyoming and western Montana include low decumbent and semi-erect forms (Miller 1964). They are

evergreen, propagate by stem layering, and produce

profusely branching root systems (Miller 1964). Some

plants are also capable of resprouting, a feature uncommon in curlleaf mountain mahogany. The decumbent ecotypes exist primarily on shallow soils developed from a fractured limestone substrate.

Members of these ecotypes vary in stature and

growth form. Most develop multiple stems and are

semi-erect with leaves of varying sizes. The decumbent growth form and the hybrids provide an extremely valuable base for future plantings. If adapted

to typical curlleaf mountain mahogany sites, the
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Chapter 22 Rosaceous Shrubs

hybrids could be used to decrease the size of the treelike curlleaf mountain mahogany growth forms. In

addition, the stem layering and root sprouting features could enhance the fire tolerance and soil stabilization values of this important shrub.

Cercocarpus montanus __________

True Mountain Mahogany

True mountain mahogany also known as blackbrush,

deerbrush, alder, sweetbriar, and tallow brush, is an

erect shrub or small tree 2 to 13 ft (0.6 to 4 m) in height.

Stems of main branches have smooth, grayish-brown

bark (fig. 6). Young twigs are often villous. The short

petiolate leaves are oval to broadly ovate, serrate

above the middle, 0.4 to 2.0 inches (1 to 5 cm) long, and

0.4 to 1 inch (1 to 2.5 cm) wide. Leaves are deciduous

to semi-evergreen, dark green and glabrous above

with prominent veins and a fine whitish tomentum

beneath. Flowers are perfect and regular, borne in

small clusters in the axils of short spurs. There are

five sepals extending from a trumpetlike hypanthium, no petals, 25 to 40 stamens in two to three

whorls, and a single pistil. The fruit is an elongate

achene with a plumose style, 2.4 to 3.9 inches (6 to

10 cm) long at maturity. Flowering occurs from midMay to late June. Fruits mature from July into September depending on elevation. They are wind dispersed (Harrington 1964; Hitchcock and others 1961;

Welsh and others 1987).

Chromosome number of true mountain mahogany is

n = 9 (McArthur and Sanderson 1985). Natural hybridization between species of mountain mahogany in

Utah has been observed in areas of overlapping ranges.

Curlleaf mountain mahogany x littleleaf mountain

Figure 6—True mountain mahogany post-flowering, but prior to development of the elongate plumose styles.

mahogany hybrids are most abundant because oftheir

overlapping flowering periods. True mountain mahogany x curlleaf mountain mahogany hybrids are

less common, and true mountain mahogany x littleleaf

mountain mahogany hybrids are rare (Pyrah 1964).

Ecological Relationships and Distribution

The range of true mountain mahogany extends from

Montana to South Dakota and south to New Mexico,

Arizona, and Colorado (Harrington 1964; Martin 1950)

in areas receiving 10 to 23 inches (25 to 58 cm) of

annual precipitation. It occupies bluffs, mountain

slopes, and foothills from 3,500 to 10,000 ft (1,070 to

3,050 m) in elevation. It commonly grows on rocky or

well-drained soils on all exposures. It is also common

on moist fertile sites, canyon bottoms, and north and

east slopes. It usually grows in association with other

shrubs and trees including ponderosa pine, Gambel

oak, Utah juniper, pinyon, and other woody species

of mountain brush communities (Blauer and others

1975). It frequently grows with mountain big sagebrush. Pure stands do exist, but are limited in extent.

In Utah the species is widespread, occupying diverse

situations, but most commonly occurring on shallow

soils and slickrock sites (Greenwood and Brotherson

1978). In Colorado, Medin (1960) found soil depth to be

the most important factor influencing true mountain

mahogany production on both shale and sandstonederived soils. Clay content of the A horizon was very

important in sandstone-derived soils; greater clay

content was beneficial. Factors influencing water availability to the plant were key in influencing production.

Brotherson and others (1984) also found factors

affecting water availability had a significant impact

on the development of true mountain mahogany

communities. In Utah, Anderson (1974) reported a

combination of site factors including soil texture, percent litter, soil pH, and exposed rock influenced the

distribution of the shrub. However, no specific combinations of factors could be used to predict its occurrence. Stands on north-facing slopes of central Utah

study sites were transitional in succession to other

mountain brush types (Christensen 1964). Succession

progressed much more slowly on more xeric southerly

sites, and perhaps would never progress beyond the

true mountain mahogany stage (Brotherson and others 1984).

Plant Culture

Reproductive phenology of true mountain mahogany

is related to aspect and elevation (Blauer and others

1975). Stands of true mountain mahogany growing on

fertile sites usually produce an abundant seed crop.

Seeds are more easily harvested than those of curlleaf

mountain mahogany; plants are smaller in stature.
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Chapter 22 Rosaceous Shrubs

Seed is usually hand collected from wildland stands;

no commercial nurseries are being maintained.

Plumes are removed with a Dybvig or barley debearder,

followed by screening. Piatt (1973) reported that the

larger seed of true mountain mahogany germinate

better than small seed, but relative size may not be a

reliable index of differences in source germinability.

Seed may be stored for up to 7 years in an open

warehouse without a major decrease in viability

(Stevens and others 1981a). Purity of 90 percent and

germination of 80 percent are recommended for seed

purchases. There are approximately 59,030 seeds per lb

(130,138 per kg) at 100 percent purity (see chapter 24).

Ecotypes of this species appear much more sensitive to site conditions than do collections of curlleaf

mountain mahogany. Utilization of ecotypes acquired

near the proposed planting site or from similar sites is

recommended.

Seed must be fall sown at a depth of approximately

0.5 inch (1.25 cm). Surface plantings are not successful and should be avoided. Growth of emerging

seedlings is seriously depressed if they are subjected

to competition from herbs. Consequently, the shrub

should be seeded in strips or plots separate from

grasses. Seeding this species with a Hansen seed

dribbler attached to a caterpillar tractor has been very

successful on pinyon-juniper chainings. This method

also serves to prevent development of such a dense

shrub overstory that herbaceous species are eliminated. Interseeding the shrub into established stands

of perennial grasses can be successful if strips or

patches of understory vegetation are first removed.

True mountain mahogany is frequently scattered

among the upper big sagebrush communities. In those

situations it has sometimes been eliminated or seriously reduced by browsing. Natural recovery is slow,

and artificial seedings are difficult to establish if the

sites are infested with weedy annuals. If populations

of exotic annuals are low, natural and artificial seedings

are quite successful.

Germination requirements are similar to those of

curlleaf mountain mahogany; seeds require afterripening and wet prechilling (Deitschman and others

1974a). Seedlings emerge shortly after the ground

thaws. Many of the small seedlings are lost to drought

and early spring frost (Plummer and others 1968).

Mortality also results from severe browsing by rodents, rabbits, and big game. Consequently, seedings

should be protected from browsing for 3 years or more.

Seedlings are easily grown as either container or

bareroot stock. Planting stock must be dormant or

properly hardened to ensure success. Bareroot seedlings should not be root pruned during lifting or

outplanting. Transplants should be planted in strips

or scalps cleared of competing vegetation. Field survival of container or nursery grown transplants of

all mahogany species is only fair to good. However,

true mountain mahogany usually transplants better

than curlleaf mountain mahogany.

Uses and Management

Medin and Anderson (1979) and Tueller (1979) reported true mountain mahogany is one of the principal browse species used by wintering deer within its

range. The widespread occurrence, nutritive value,

and high palatability of this species ensure its rank

as a valuable shrub for big game habitat (Plummer

and others 1968; Young and Bailey 1975). The plant

also receives considerable spring and summer use by

big game and livestock and provides cover for numerous wildlife species. Compared to curlleaf mountain

mahogany it is a superior browse for livestock, particularly sheep (Ferguson 1983). True mountain mahogany is fairly tolerant of browsing. A Colorado

clipping study indicated plants can withstand 60 to

80 percent browsing (Shepherd 1971). However,

Brotherson and others (1984) found that about onehalf of the true mountain mahogany in central Utah

stands was highlined and out of reach of browsing

animals. Some stands have been severely reduced by

heavy browsing (Ream 1964).

Nutrient value of true mountain mahogany in winter is not as high as that of curlleaf mountain mahogany. Digestibility of true mountain mahogany

has been reported as 48.4 percent. Digestible protein

is 3.4 percent (Smith 1957; Urness and others 1977;

Welch 1981); adequate winter digestible protein content is 5.2 percent for wintering deer.

True mountain mahogany also has value as an

ornamental. Plants are used for landscaping along

the West coast. In inland areas they are very hardy,

requiring low maintenance. Its semi-evergreen habit,

shiny plumose fruits, and growth habit are unique

and attractive. It grows rapidly with limited irrigation.

Burning is a good technique for rejuvenating stands

and increasing plant vigor. True mountain mahogany

can resprout vigorously from surviving root crowns

following burning. Recovery may be rapid even following rather intense burns (Blauer and others 1975;

Wright and others 1979). Survival after burning may

be high at all seasons, but response is greater following dormant season burns in Colorado (Young 1983).

Current annual growth measured in late winter

was 475 to 650 percent greater on burns than on

controls. Production was greater on low intensity than

on high intensity burns. Seed dispersed long distances

by winds from unburned sites also contributes to

recovery of burned or otherwise disturbed sites.

True mountain mahogany has been established

from direct seedings on mine disturbances in Utah and

Idaho (Ferguson and Frischknecht 1985). Seedlings
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Chapter 22 Rosaceous Shrubs

survive on infertile soils; on spoil wastes they appear

to benefit by the addition of even small amounts of

topsoil. Plants may be nodulated by the nitrogenfixing endophyte Frankia (Hoeppel and Wollum

1971). Transplants grow slowly and are sensitive to

competition.

Varieties and Ecotypes; Other Species

Attempts have been made to enhance seed germination, seedling vigor, and growth rate of true mountain mahogany by selection of ecotypes. A selection

referred to as “Montane” was released by the U.S.

Department of Agriculture, Natural Resource Conservation Service, Los Lunas Plant Materials Center

(Thornburg 1982). This cultivar, collected on the Santa

Fe National Forest near Coyote, NM, has nondormant

seed and excellent seedling vigor. Its adaptation to

sites outside its area of collection has not been fully

documented.

Coleogyne ramosissima __________

Blackbrush

Blackbrush, the only member of its genus, is a longlived densely branched shrub usually about 3 ft (0.9 m)

in height, but sometimes reaching 6 ft (1.8 m) or more

(fig. 7). The root system is shallow and spreading with

most roots less then 16 inches (40 cm) deep. Roots are

associated with the nitrogen fixing actinomycete

Frankia (Bowns and West 1976; Dittberner and Olsen

1983). The tangled ash-gray branches give the shrub

a dark appearance, particularly when wet; hence the

name blackbrush (Benson and Darrow 1945). Main

Figure 7—Blackbrush community in southwestern Utah.

stems proliferate by stem splitting. The short branches

die back from the apex after a few years of growth

leaving a spiny tip. Lateral branches then develop

producing a rounded shrub (Bowns and West 1976).

The small persistent leaves are clustered along the

branches. They are simple, leathery, about 0.2 to

0.6 inch (5 to 15 mm) long, and 0.04 to 0.12 inch (1 to

1.5 mm) wide. The flowers are terminal on new

branchlets. They have four yellow to brownish sepals

and are usually apetalous. Numerous stamens are

inserted at the base of the disk that encircles the

ovary. The fruit is a glabrous, brown achene 0.2 to

0.3 inch (5 and 8 mm) in length (Harrington 1964;

Munz and Keck 1959; Welsh and others 1987). Plants

normally flower as early as March at lower elevations

(McMinn 1951) and as late as June at higher elevations (Munz and Keck 1959). Seeds mature in midsummer. Blackbrush has n = 8 chromosomes, which is

unusual for the subfamily Rosoideae (McArthur and

Sanderson 1985). The 4-merous apetalous flowers

and opposite leaves are also unusual for the Rosaceae.

New growth and flowering are dependent on the

quantity and distribution of precipitation. In the absence of adequate fall or winter/spring rain, vegetative and reproductive growth are curtailed (Ackerman

and others 1980). Blackbrush enters a period of summer inactivity in response to low soil moisture. Flowering occurs from April to August depending on location.

Fruits ripen from April to June and are dispersed

shortly thereafter (Ackerman and others 1980).

Ecological Relationships and Disribution

Considered a relict endemic species of arid and semiarid areas of western North America (Stebbins 1972),

blackbrush’s present distribution is thought to represent a restriction of a once more extensive range

(Stebbins and Major 1965). Blackbrush is distributed

from southern California, east through southern

Nevada, northern Arizona, southeastern Utah and

southwestern Colorado in areas with annual precipitation in the 6 to 12 inch (150 to 300 mm) range

(Harrington 1964; Munz and Keck 1959; Turner 1982;

USDA Soil Conservation Service and USDI Bureau of

Land Management 1981; Welsh and others 1987).

Blackbrush occurs as a dominant species over extensive areas and is particularly common in the

southwestern Great Basin Desert along the Colorado

River drainage system (Turner 1982). Blackbrush is a

major shrub along the Transition Zone between the

Great Basin and Mojave Deserts (Tidestrom 1925).

Monotypic stands within this Zone intergrade into

mixed communities of Mojave Desert, Colorado Plateau, and Great Basin vegetation.

Monotypic stands of blackbrush are characterized

by low understory species diversity and greater total
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Chapter 22 Rosaceous Shrubs

cover than most desert shrub communities. Plants are

long-lived and slow growing; their communities are

highly stable. Blackbrush often replaces itself following disturbances, but reestablishment of shrub cover

occurs very slowly due to extremely erratic seedling

establishment and low growth rates (Webb and others

1987). In southern Nevada, Beatley (1974) characterized blackbrush as requiring Great Basin precipitation and Mojave Desert temperatures and existing

primarily between creosotebush and pinyon-juniper

or sagebrush communities (Wells and Jorgensen

1964). In Arizona, blackbrush occurs in areas having

a mean annual soil temperature of 59 ∞F (15 ∞C)

(USDA Soil Conservation Service and USDI Bureau

of Land Management 1981).

In general, blackbrush stands occur on upland

sites, broad terraces, and gently rolling hills (USDA

Forest Service and USDI Bureau of Land Management 1981). Soils are usually shallow, calcareous, and

coarse textured, ranging from gravelly to cobbly loam

or gravelly clay loam (Foster 1968; Kearney and Peebles

1960; Shreve 1942). The shrub is less prevalent and

occurs with an understory dominated by grasses in

drainage bottoms, where soils are moderately deep to

deep with fine sandy loam surface textures and calcareous underlying layers of very gravelly sandy loam or

gravelly clay loam (USDA Soil Conservation Service

and USDI Bureau of Land Management 1981). Other

blackbrush sites with good fertility and soil water also

support a herbaceous understory (Bowns 1973).

Blackbrush normally occurs on soils low in salt content (McCleary 1968). Seedlings are sensitive to salinity, which may limit its distribution and spread

(Wallace and Romney 1972).

Blackbrush is often a component of many transitional shrub communities that provide important forage and habitat for game and livestock. These include

spiny hopsage, Nevada ephedra, rabbitbrush, big sagebrush, desert bitterbrush, cliffrose, and pinyon-juniper

communities. It is considered a potential candidate for

use in revegetation; it occupies or once occupied extensive areas that have suffered severe abuse by destructive grazing practices.

Plant Culture

Blackbrush seed production is highly erratic. Most

stands produce a harvestable seed crop in perhaps

1 of every 6 to 10 years. Ripe seed readily detach from

the plant and are dispersed by gravity. Seed collection is slow and tedious; consequently, commercial

supplies are rarely available. Fruits are cleaned

using a fanning mill or gravity separator. Purity of

95 percent and germination of 70 percent are recommended for seed purchases. There are approximately

27,015 seeds per lb (59,557 per kg) at 100 percent

purity (see chapter 24).

Seeds require a short wet prechilling period to relieve dormancy. Bowns and West (1976) reported

prechilling at 39 ∞F (4 ∞C) for 23 days and incubation

at 57/40 ∞F (14/4 ∞C) provided 80 percent germination

in 6 days and 90 percent in 13 days. Natural seedlings

are uncommon. Their appearance may be associated

with dry autumns and winter or spring precipitation

totaling 2 to 5 inches (50 to 130 mm) (Beatley 1974).

Plants can be established by artificial seeding, but

sufficient plantings have not been attempted to adequately determine appropriate seeding rates or seedbed requirements. Local seed sources should be used;

populations are not adapted to areas other than their

sites of origin. Seed can be planted with most conventional seeders. A planting depth of about 0.5 inch

(1.3 cm) is recommended. Rodents may aid in natural

seed dispersal and planting, but rodent predation of

planted seed has been observed (Bowns and West

1976). Fall planting is recommended to provide adequate wet prechilling and reduce rodent problems

(Bowns and West 1976). Stark (1966) reported good

success from seedings in Nevada, but wildland seedings

in Utah have not been as successful. Seedlings developing from plantings in central Utah, north of the

species natural range, have failed to persist for more

than 2 to 3 years due to cold winter temperatures.

Seedlings developed slowly and some were suppressed

by herbaceous competition. In general, most attempts

to plant blackbrush offsite have not been successful.

Only fair survival has resulted from planting

projects using either container or bareroot stock. Container stock grows moderately well in a buffered peatvermiculite medium, but seedlings are sensitive to

overwatering. Four to 6 inch (10 to 15 cm) tall seedlings can be produced in 6 to 10 months under greenhouse conditions. Survival has not exceeded 30 percent from plantings conducted in southeastern Utah.

Shading the small transplants improved survival

(Kitchell n.d.).

Uses and Management

Dayton (1931) and Allen (1939) reported blackbrush

provides fair winter forage for cattle, sheep, goats,

deer, desert bighorn, and other game animals. Blackbrush communities are commonly used as winter

range for sheep. However, blackbrush is often regarded as a poor forage species with low palatability

(Bates 1984; Sampson and Jesperson 1963). Although

used by big game and livestock (Gullion 1964), it is

usually not a major component of their diet. It may

become an important forage during severe winters

(Kufeld and others 1973; Sampson and Jesperson

1963). Blackbrush is deficient in phosphorus for sheep

and cattle and low in protein during fall and winter

(Bowns and West 1976). In addition, the low growing,

intricately branching habit of the shrub reduces its
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Chapter 22 Rosaceous Shrubs

attractiveness to livestock (Provenza and others 1983).

Light to moderate grazing may reduce woody materials and spininess and increase production of palatable

new growth (Provenza 1977; Provenza and others

1983). Mule deer make moderate use of this shrub in

winter and spring (Leach 1956). Its seeds are used by

birds and rodents (Mozingo 1987; USDA Forest Service

1937) and it provides cover for many small animals.

Blackbrush is adapted to harsh sites, but its slow

growth and limited area of adaptation have restricted

its use in revegetation projects on non-native sites. All

blackbrush sites do not respond similarly to restoration and management practices. Bates (1984) and

West (1969) concluded that blackbrush occupies two

different situations, each responding differently to fire

and other improvement techniques. In the first situation, blackbrush occurs in monotypic stands as a

climax species supporting few native understory species. Shrubs are widely spaced and cover provided by

grasses and forbs may be extremely low. Cryptogamic

crusts covering interspaces reduce soil erosion; those

with a blue-green algae phycobiont provide an important source of soil nitrogen (West and Skujins 1977).

These sites are frequently considered for range improvement projects. Burning has not been a successful

means of releasing the existing understory due to the

unpredictable recovery of native herbs, invasion of

annual weeds, and loss of cryptogamic crust that

recovers very slowly (Jensen and others 1960). In the

absence of annual weeds, these sites often revert back

to blackbrush without any intermediate stage (West

and Skujins 1977). Attempts to plant substitute

shrubby species have not been successful.

Monotypic stands of blackbrush are often associated

with shallow soils and soil water conditions that do

not favor seeding success. Annual weeds, principally

red brome, compete with the shrub seedlings. Attempts to replant blackbrush back onto these sites

following wildfires or other disturbances often meet

with failure.

The second common revegetation situation occurs

on sites where blackbrush invades other plant communities or is naturally present in low density. Invasion normally occurs on sites with deeper soils and

better water availability than is found on monotypic

blackbrush sites. Blackbrush increases on these sites

when heavy grazing suppresses other plants. Under

these conditions, burning or chaining can be used to

reduce blackbrush density and allow release of existing natives or aid in the establishment of seeded

species. Success is dependent on edaphic and climatic

conditions. Sites with deeper soils generally respond

more favorably (Bates 1984; Halliday 1957).

The successional patterns of recovery following fires

are somewhat unpredictable in blackbrush communities. The shrub is reported by some to be a nonsprouter

(Bowns 1973; Bowns and West 1976). Most plants are

killed if the understory is dense enough to carry a fire.

Low intensity burns may not kill mature plants.

Bates (1984) reported considerable resprouting following fires in California. Blackbrush seed may remain viable in the soil for several years; new seedlings

may emerge from seed banks many years after mature

plants are destroyed by fires or other disturbances

(Bates 1984). Blackbrush seedlings establish very

slowly following fires (Anderson 2001; Wright and

Bailey 1982); red brome and other annuals develop

quickly and dominate burned sites, often precluding

blackbrush recovery (Rickard and Beatley 1965). However, Bates (1984) observed blackbrush reestablishment from seed on sites occupied by annual and

perennial grasses following wildfires. Thus, the

species may exert some competitiveness with herbaceous species in some locations.

Jensen and others (1960) found burns in Nevada

effectively killed blackbrush plants. Unless native

perennial grasses and broadleaf herbs recovered rapidly following burns on these sites, annual weeds

quickly dominated and prevented blackbrush recovery. Some shrubs, including turpentine bush, desert

bitterbrush, and desert peachbrush, are capable of

recovering following fires, and if present prior to burning, may soon dominate burns where blackbrush was

once common (Bowns 1973).

In California, annuals quickly invaded blackbrush

burns, but were replaced by perennial herbs within

10 to 15 years (Bates 1984). A mixed shrub community

then appeared in about 30 years, eventually reverting

back to solid stands of blackbrush if the stands were

poorly managed. Not all sites responded so favorably.

Once annual weeds established, the reentry of more

desirable herbs and shrubs was often limited. Burning

or chaining increased the density of other shrubs

present after treatment.

Blackbrush communities provide desirable ground

cover and soil protection and should not be removed or

converted to other species unless the treatments are

assured of some success. Anchor chaining has been

effective in reducing blackbrush density where it exists in pure stands or intermixed with pinyon-juniper.

Brush beating has also been used to decrease shrub

density, remove older wood, and stimulate new growth

(Bowns and West 1976). Neither chaining nor brush

beating kills all shrubs. Double chaining for example,

initially killed about 25 percent of the plants on two

large blackbrush chaining projects in southern Utah

(Monsen n.d.). Crested, intermediate, and pubescent

wheatgrass were seeded at both sites, producing good

understory stands. Blackbrush resprouted and regained its earlier productivity in about 10 years. Its

productivity was not suppressed by the presence of the

seeded grasses. Indian ricegrass has also responded
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Chapter 22 Rosaceous Shrubs

favorably following chaining of certain blackbrush

sites. Blackbrush is particularly compatible with Indian ricegrass, and when protected from livestock

grazing these sites often support a mixed grass-shrub

community (Jeffries and Klopatek 1987).

Varieties and Ecotypes

Native stands in the Colorado River drainage in

southeastern Utah provide the seed for most blackbrush

revegetation projects. However, collections acquired

from this area have not demonstrated adaptation to

other locations. If artificial plantings are contemplated, seed should be acquired from native stands

growing near the planting sites. Different ecotypes

demonstrate specific attributes that appear to be genetic traits. Certain ecotypes demonstrate adaptability to arid desert conditions; others express compatibility with herbaceous understory plants, and some

appear better able to recover from burning.

Cotoneaster acutifolia ___________

Peking Cotoneaster

Peking cotoneaster is an unarmed, low-spreading

shrub, often reaching 12 ft (3.7 m) in diameter (fig. 8).

Plant height is usually less than 5 ft (1.5 m). The shrub

is described as deciduous (Welsh and others 1987), but

numerous leaves often remain on the plant until late

winter. Branches are numerous, reddish brown, and

arcuate to horizontal. Flowers are small, perfect, regular, and solitary or in cymes terminating lateral

branches. Flowers are white or pink. The fruit is a

Figure 8—Peking cotoneaster, an introduction

from China, has received considerable use in

midwestern windbreaks and shelterbelts where

it provides winter cover and food for birds and

big game.

black, two-celled pome about 0.4 inch (1 cm) long

(Johnson and Anderson 1980; Welsh and others 1987).

Ecological Relationships and Distribution

Peking cotoneaster was originally introduced from

northern China in 1883 as an ornamental shrub

(Hoag 1958; Leslie 1954; Slabaugh 1974). It has been

planted for conservation and shelterbelt purposes

throughout the United States, particularly in the

northern Great Plains and upper Midwest. It is also

used in the southern Canadian prairie provinces

(Johnson and Anderson 1980; Slabaugh 1974). Like

other Cotoneaster species, Peking cotoneaster is commercially cultured and sold for ornamental plantings.

A number of varieties have been developed and are

widely used. Plants have demonstrated a wide range

of adaptation when planted in shelterbelts and conservation plantings in the West and Midwest.

Plant Culture

Peking cotoneaster normally flowers in early May;

fruits ripen in September or October (Slabaugh 1974).

Plummer and others (1968) reported that fruits do not

ripen until mid November in Utah. Fruits remain on

the shrub until mid to late winter. Abundant crops are

often produced on cultivated hedges and shelterbelt

plantings. Fruits become rather dry as they ripen.

Mature fruits are easily dislodged from the shrub and

are hand collected in late fall or early winter after

some leaves have fallen. Seed are removed by macerating the fruits and flushing the pulp with water.

The remaining seeds and pulp are then dried and

screened. Empty seeds are removed by flotation

(Slabaugh 1974). Conditioned seedlots normally are

very pure with little debris. The average number of

cleaned seeds per lb varies from 21,984 to 32,310

(48,465 to 71,231 per kg) (Plummer and others 1968;

Slabaugh 1974). Purity of 98 percent and germination

of 80 percent are recommended for seed purchases.

Germination of Peking cotoneaster is restricted

by the presence of a hard, impermeable seedcoat

and embryo dormancy (Slabaugh 1974). Scarification

with sulfuric acid and wet prechilling pretreatments are recommended to hasten germination

(Fordham 1962). However, fall nursery or wildland

plantings usually provide sufficient wet prechilling.

An alternating temperature regime of 50 ∞F (10 ∞C)

for 15 hours and 77 ∞F (25 ∞C) for 9 hours, with light

supplied during the warm cycle, usually provides

optimal germination conditions for pretreated seed

(Slabaugh 1974).

Peking cotoneaster is commonly grown as nursery

stock for outplanting and is usually lifted after the

second growing season. Cuttings are easily propagated; the plants layer readily. Wildings or rooted
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Chapter 22 Rosaceous Shrubs

stem layers can be harvested early in the spring for

field plantings. Doran (1957) reported softwood cuttings of cotoneaster taken in early summer root well.

Wildland seedings have been only marginally successful. Seed germination is erratic. Seedlings grow

slowly and are easily suppressed by herbs. Planting

success is improved if the seedbed is kept weed free.

Established plants are more resilient and persist well.

This shrub rarely suffers heavy damage from insects,

disease, or use by small mammals. Little damage has

been noted during heavy grasshopper infestations.

Uses and Management

Peking cotoneaster has been most widely used for

windbreaks, shelterbelts, conservation, and ornamental plantings (Hoag 1958; Johnson and Anderson 1980;

Leslie 1954; Slabaugh 1974; Wyman 1949). The shrub

is used in multi-row windbreaks and cover plantings

to protect soils and reduce wind erosion. It furnishes a

low, dense groundcover that provides year-long protection. The plants are also commonly used in roadway, recreation, or conservation plantings where

ground cover, low maintenance, and aesthetics are

important. Plants are attractive and provide glossy

green foliage with persistent red or black berry-like

pomes. The shrub withstands snow deposition and is

used to trap snow and protect roadways and buildings.

Peking cotoneaster is commonly used to control

erosion in residential areas, recreation sites, and wildlands. It is particularly useful in these areas as plants

establish well from transplanting and quickly improve ground cover. Transplants can be planted at

various spacings depending on the availability of

seedlings. Planted shrubs will spread to occupy the

planting site, but do not seed into adjacent areas.

Peking cotoneaster is adaptable to many ranges

where native species have been lost and are difficult to

reestablish. It has been used to provide winter food for

various birds and big game (Johnson and Anderson

1980; Miller and others 1948). The plant exhibits high

palatability to game and livestock and is moderately

tolerant of grazing (Plummer and others 1968). It is

readily grazed in midsummer by mule deer (Kufeld

and others 1973; Leach 1956). Winter browsing has

been heavy in Utah. Spring and fall use is light. Game

animals are attracted to the shrub and can damage

landscape plantings. Peking cotoneaster should not be

used in roadway or other plantings where wildlife

concentrations create hazards.

Peking cotoneaster may be used to improve areas

where game animals concentrate and more productive

forage and cover is essential. This shrub can be planted

to enhance depleted native communities and attract

and reduce animal use of native species. It can also be

planted to reduce erosion on disturbed sites and to

protect watershed and soil values.

Varieties and Ecotypes

There are no releases.

Cowania stansburiana ___________

Stansbury Cliffrose or Cliffrose

Stansbury cliffrose has often been identified as a

variety of C. mexicana var. stansburiana [Torr.] Jeps.

However, we have elected to accept it as a separate

species as proposed by McArthur and others (1983b).

Plants are long-lived, erect, evergreen, aromatic

shrubs or small trees 3 to 18 ft (1 to 5.4 m) tall with

stiff, often somewhat brittle stems and gray shreddy

bark (Cline 1960; Welsh and others 1987). New growth

is flexible with glandular bark. Stansbury cliffrose has

an extensive root system with long, spreading, and

descending roots (Sutton and Johnson 1974). Roots

may form symbiotic nitrogen-fixing associations with

Frankia (Nelson and Schuttler 1984). Plants are resinous and strong smelling. The light to dark green

leaves are clustered along the branches and are 0.5 to

1 inch (1.2 to 2.5 cm) in length with revolute margins

and five to seven teeth at the apex. Leaf surfaces are

glandular dotted and resinous above with a white

tomentum beneath (Blauer and others 1975; Munz

and Keck 1959). About 90 percent of the showy, fragrant, cream to yellow flowers appear during the first

flush of flowering and are formed on lateral spurs of

the previous year’s wood. Later flowers appear sporadically throughout the summer near the tip of the

current year’s leaders (Shaw and Monsen 1983b).

Later flowers are particularly abundant following summer rains. Flowers are about 0.8 inches (2 cm) in

diameter and are normally perfect with two series of

stamens enclosing 5 to 12 hairy pistils. Fruits are

plumose achenes (fig. 9) dispersed by gravity and wind

(Blauer and others 1975).

Figure 9—Stansbury cliffrose branch with

mature achenes, note the plumose styles.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 557

Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution

Stansbury cliffrose is distributed from southern

California east to Colorado and south to Sonora and

Chihuahua (USDA Forest Service 1937; Welsh and

others 1987). It does not occur north of about the 42 ∞N

parallel (McArthur and others 1983b). It is found

primarily on dry, rocky, well-drained silty to gravelly

soils of foothills and mesas growing in blackbrush,

chaparral, oak, desert shrub, mixed grass-desert shrub,

mountain brush, and pinyon-juniper woodlands up to

the edge of ponderosa pine forests at elevations from

3,000 to 9,000 ft (900 to 2,700 m) (Blauer and others

1975; McArthur and others 1983b; Welsh and others

1987).

Cowania and Purshia are closely related genera but

differ in many physical characteristics (table 1) and

geographical distribution (McArthur and others 1983b;

Stutz and Thomas 1964; Thomas 1957). Because of

these differences, we prefer to retain the genus name

Cowania even though Welsh and others (1987) recently transferred its species to Purshia.

Disjunct flowering periods normally prevent hybridization between antelope bitterbrush and

Stansbury cliffrose. Antelope bitterbrush usually flowers earlier than Stansbury cliffrose (McArthur and

others 1983b; Shaw and Monsen 1983b), but aspect or

site conditions may result in overlapping periods of

anthesis and pollination. Stutz and Thomas (1964)

and Thomas (1957) reported that hybridization is so

common that no populations of antelope bitterbrush

in Utah have been examined that are free from the

influence of Stansbury cliffrose. Introgression of antelope bitterbrush into Stansbury cliffrose is less common, but does occur. McArthur and others (1983b)

Table 1—Morphological characteristics separating Cowania

and Purshia (after Stutz and Thomas 1964).

Character Purshia Cowania

Twig pubescence heavy glabrous

Dorsal leaf pubescence heavy glabrous

Hypanthium glands none 3+/mm2

Stalked glands absent present

Pistils 1 or 2 4+

Style length <1.5 cm 1.6 - 3.4 cm

Style pubescence puberulent plumose

L/W ratio of achenes <3.1 >3.3

Stamen insertion hypanthium hypanthium

 margin throat

Leaf margin revolution slight pronounced

Hypanthium pubescence pubescent glabrous

Stamen series one two

Lobes/leaf three five+

Leaf glandulosity none 3+/mm2

Leaf retention deciduous evergreen

concluded that two adaptive products have evolved

through natural hybridization. Desert bitterbrush,

the southern adaptive derivative, is one product. The

second is less distinct. It occurs in areas north of

Utah on sites not occupied by Stansbury cliffrose and

consists of antelope bitterbrush populations that express Stansbury cliffrose characteristics, particularly

low palatability. Stutz (1972) suggested selection pressure resulting from heavy browsing has favored the

less palatable plants.

Both Purshia and Cowania were present in the

Madro-Tertiary geoflora, which evolved from the

Arcto-Tertiary and Neo-Tropical geofloras in response

to increasingly drier conditions in the Southwest

(Axelrod 1958, McArthur and others 1983b). McArthur

and others (1983b) postulated that the weak isolation

barriers now existing between the two genera indicate

that the present zone of contact of the two species

developed after a long separation. Much of the area

where they now coexist was occupied by Pleistocene

lakes and glaciers as recently as 12,000 years ago.

Plant Culture

Although Stansbury cliffrose grows in semiarid regions it usually produces at least some seed each year.

Abundant or commercially harvestable seed crops

develop in about 2 out of every 3 to 4 years. Although

flowering usually occurs after the last spring frost,

unusually late frosts can diminish seed yields.

The period of floral development, anthesis, and fruit

development lasts from 68 to 78 days in southern

Idaho (Shaw and Monsen 1983b). Anthesis and pollination occur over a 10-day period. Fruits then mature

about 37 days later. The plants are insect-pollinated

and self-incompatible, consequently seed production

is significantly impacted by high winds, rainy periods,

unusually hot temperatures, and other factors that

influence insect pollination.

Fruit (achene) ripening is less uniform for Stansbury cliffrose than for antelope bitterbrush, but the

bulk of the fruit can generally be harvested on one

date. Fruits are hand collected by striking the limbs

with a paddle or stick to knock them into collection

hoppers or trays. The brittle styles are removed with

a Dybvig or barley debearder and separated by fanning. Seeds are not removed from the achene to avoid

damaging the embryo (Young and Evans 1983). Generally about one-third of the harvested material by

weight consists of viable seed (achenes) (Alexander

and others 1974). There are about 64,600 seeds per lb

(64,500 per kg) (Alexander and others 1974; Plummer

and others 1968). Purity of 95 percent and germination of 85 percent are recommended for seed purchases (see chapter 24).

Seed can reportedly be warehouse stored for 5 to

7 years (Alexander and others 1974; Plummer and
558 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

others 1968; Springfield 1973), or even as long as 15

years (Stevens and others 1981a). Germination tests

conducted on seed lots harvested from the same area

in different years exhibit similar germination patterns. However, some differences do occur. Poorer and

slower germination results from unfavorable growing

conditions during the year of seed production; more

rapid and complete germination characterize seed

produced during years of abundant production

(Jorgensen n.d.). Shaw and Monsen (1983b) reported

the loss of achenes prior to the time of maturation

was 13 percent in 1979 and 52 percent in 1980.

Although insects were a major factor, losses could not

be attributed to insect damage alone.

Stansbury cliffrose seed requires a wet prechilling

treatment to release dormancy. The optimal treatment duration varies with population and year of

harvest. Consequently, seed should be planted in fall

or winter; spring seeding should be avoided. Meyer

and Kitchen (n.d.) found that the rate and total germination percentage increased from 30 percent after 2

weeks of wet prechilling at 33 ∞F (1 ∞C) to 100

percent in 7 days after 6 weeks of wet prechilling.

Seed maintained at 33 ∞F (1 ∞C), began germinating

during prechilling within 8 weeks and completed

germination in 11 weeks. Plummer and others (1970b)

reported 89 percent germination after 90 days at

alternating temperatures of 32/85 ∞F (0/31 ∞C) and 99

percent germination after 90 days of alternating

temperatures of 32/38 ∞F (0/3 ∞C). Young and Evans

(1981b) recommended prechilling at 32, 36, or 39 ∞F (0,

2, 4 ∞C) for approximately 6 weeks. Seeds do not

require light for germination (Sabo and others 1979).

Stansbury cliffrose is generally fall seeded in a

mixture with other shrub seed. Seed can be planted

with most conventional drills or seeders. Cleaned

seed must be incorporated into the soil to a depth of

about 0.25 to 0.50 inch (6 to 12 mm) (Alexander and

others 1974; Plummer and others 1968). Surface broadcasting is generally unsuccessful if the seed is not

subsequently covered. Seedling emergence is similar

to that of antelope bitterbrush, occurring early in

spring. Small seedlings are sometimes subjected to

late frosts. During the establishment period, seedlings are quite sensitive to competition and water

stress. However, once established, Stansbury cliffrose

seedlings tolerate drought, heat, and soil salinity

better than antelope bitterbrush or Apache plume

(Plummer and others 1968). Seedlings are not capable

of competing with cheatgrass, red brome, or dense

stands of some perennial grasses.

Stansbury cliffrose is easily grown as bareroot (Shaw

1984) or container stock (Landis and Simonich 1984);

it develops rapidly if properly cultured. One-year-old

bareroot seedlings generally attain satisfactory size

for field transplanting. Container plants are usually

propagated from seed, but stem cuttings can be rooted

(Shaw and Monsen 1983b). Because Stansbury cliffrose

is an evergreen shrub, plants are sensitive to lifting

and transplanting if not properly hardened or dormant. Survival of actively growing plants is generally

low.

Grasshoppers utilize Stansbury cliffrose (Edgerton

and others 1983) but have not been observed to seriously damage or eliminate new plantings. Weedy

competition on the planting site should be controlled

by burning, mechanical, or other means of seedbed

preparation prior to transplanting (Giunta and others

1978; Shaw and Monsen 1983b).

Uses and Management

Stansbury cliffrose is regarded as a desirable winter

forage for big game as well as cattle and sheep (Blauer

and others 1975; Plummer and others 1968; Robertson

1974; Tueller 1979; USDA Forest Service 1937). Kufeld

and others (1973) reported that mule deer make heavy

use of it in fall, winter, and spring but only moderate

use in summer. Some Stansbury cliffrose stands are

grazed by big game and livestock during all seasons.

However, the species is not as universally palatable as

antelope bitterbrush (McArthur and others 1983b;

Stutz and Thomas 1964). Plants often receive negligible use in spring and summer in areas where other

forage is available (USDA Forest Service 1937). Certain ecotypes, including some southern Arizona populations, evidently have low palatability in all seasons.

Two dimensional chromatograms of their phenolic

compounds can be distinguished from those of more

palatable populations (Blauer and others 1975).

Like other evergreen species, Stansbury cliffrose

maintains its protein levels and energy values in

winter (Dittberner and Olsen 1983; Smith and Hubbard

1954; Welch and others 1983a). In a common garden

study in southern Idaho, Stansbury cliffrose leaders

were found to contain a significantly higher percentage of crude protein in winter (8.4 to 8.6 percent, dry

weight basis) than those of antelope bitterbrush or

Apache plume, but slightly less than those of desert

bitterbrush (Welch and others 1983a). The in vitro

digestible dry matter of Stansbury cliffrose exceeded

that of all species tested (37.6 percent).

Stansbury cliffrose can withstand heavy grazing

and does occur in areas where winter use is quite

intense. Flowering and production of new growth may

be stimulated by 65 percent use; browsing in excess of

80 percent of new growth decreases vigor (Wasser

1982). Stands frequently sustain game animals during years of heavy snow accumulation and should be

managed to ensure forage availability during the

critical midwinter period.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 559

Chapter 22 Rosaceous Shrubs

Stansbury cliffrose has been seeded on a variety of

sites to improve diversity and winter wildlife habitat

value of vegetation (Edgerton and others 1983). Good

stands can be established in the mountain brush zone

and in most pinyon-juniper communities (Davis 1983;

Giunta and others 1978; Monsen and Christensen

1975). Stansbury cliffrose is frequently intermixed

with Wyoming big sagebrush, blackbrush, and salt

desert shrubs and can be successfully reseeded on

such sites. Although Stansbury cliffrose frequently

grows on soils derived from limestone, it is not limited

to such sites. Establishment and survival are less

certain when the species is planted in shrub communities where it does not occur naturally. However,

plantings have established and survived successfully

for nearly 40 years in antelope bitterbrush communities in south central Idaho and for over 10 years in

northeastern Oregon and eastern Washington (Blauer

and others 1975; Edgerton and others 1983; Monsen

and Shaw 1983c). Stansbury cliffrose is not fire tolerant and should not be planted in areas subject to

repeated burning.

On native sites Stansbury cliffrose may function as

a pioneer species following disturbances (Wright and

others 1979). It has been considered a candidate plant

for mined areas, roadways, and other disturbed sites.

It provides only moderate erosion control due to its

upright habit and low growth rate (Dittberner and

Olsen 1983; Smith and others 1978). Its frequency of

nodulation by the nitrogen-fixing endophyte is generally low and varies with collection date and location

(Nelson 1983). Plants have been successfully established on mine sites near Alton, UT, and on the

phosphate mines of southeastern Idaho (Ferguson and

Frischknecht 1985; Monsen, data on file, Shrub Science

Lab., Provo, UT). Replacement of topsoil is required to

attain satisfactory stands. Results of roadway seedings

and transplanting trials in Utah and Idaho have been

erratic, even when plantings are conducted in areas

where Stansbury cliffrose naturally occurs.

Stansbury cliffrose is an attractive, drought-tolerant

ornamental. The leaves are evergreen, the flowers and

fruits are showy, and flowering occurs over a prolonged period. Mature shrubs can become rather ragged

in profile, but respond well to pruning.

Stansbury cliffrose seedlings grow slowly and may

require 3 to 4 years to reach a height of 6 to 12 inches

(15 to 30 cm) (Monsen and Christiansen 1975). Growth

thereafter is much more rapid. In a big game exclosure

near Boise, ID, Stansbury cliffrose planting stock

grew more slowly than several sources of antelope

bitterbrush for the first 4 to 5 years, but exceeded

their growth rates thereafter. Both species produced

seed 5 to 6 years after planting (Shaw and Monsen

1983b).

Seedlings of Stansbury cliffrose must be protected

from excessive competition and heavy grazing until

plants are 3 to 5 years of age. In addition, Stansbury

cliffrose stands have been and are being lost due to the

invasion of weedy annuals that restrict natural or

seeded shrub establishment and expansion and ultimately contribute to the loss of the shrub overstory

(Cline 1960; Price 1982). Sites should be managed or

treated to create a weed-free seedbed and maintain a

suitable native understory after planting.

Stansbury cliffrose frequently does not recover from

burning, chaining, or other methods of mechanical

treatment. Multiple stemmed shrubs are more tolerant than older single-stemmed shrubs (Plummer and

others 1968). Extreme variability in response to fire

has been reported (Ferguson 1983; Howard 1995;

Wright and others 1979), perhaps reflecting differences among ecotypes. Attempts to improve existing

stands using these treatments have not been successful unless the sites are also seeded.

Varieties and Ecotypes

The extensive natural hybridization of Stansbury

cliffrose with antelope and desert bitterbrush, and to

a lesser extent Apache plume, provides an immense

array of material for improving palatability; increasing drought, heat, and fire tolerance; broadening areas

of adaptation; enhancing evergreen growth characteristics and forage traits; and manipulating growth

habits of these taxa (McArthur and others 1983b;

Monsen and Davis 1985; Stutz 1972). Improvement of

seedling vigor, establishment, and initial growth rates

would enhance the use of Stansbury cliffrose and its

hybrids.

Stansbury cliffrose selections from central Utah

have been among the most successful and productive

populations seeded on pinyon-juniper sites in Utah

(Monsen and Davis 1985). Hybrids with antelope

bitterbrush collected from Utah County, UT, also

offer excellent forage traits, including evergreen

growth, high productivity, dramatic regrowth, good

palatability, and an upright growth form. Davis

(1983a) listed five different central Utah accessions

of Stansbury cliffrose that have performed best in

artificial seedings in Utah nearly 30 years after

planting. Studies were based on continuous evaluations from 60 sites and include such factors as growth,

persistence, and vigor.

Stansbury cliffrose introgressions with desert bitterbrush acquired from Benton, CA, demonstrate a

wide range of adaptation and survival under harsh

circumstances. Selections from hybrids with desert

bitterbrush show promise for improved fire tolerance

and adaptation to arid situations.
560 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Crataegus douglasii _____________

Douglas or River Hawthorn

Douglas or river hawthorn is a compact, deciduous,

deep rooted shrub or small tree growing 3.3 to 13

(26) ft (1 to 4 [8] m) tall (fig. 10). The bark is smooth

and reddish brown, becoming reddish bronze in winter. Stems are armed with stout, usually straight

thorns 0.4 to 0.8 inch (1 to 2 cm) long. The petiolate

leaves are 1.2 to 2.4 inches (3 to 6 cm) long, usually less

than half as wide, and lanceolate to elliptic ovate. Leaf

edges are serrate to doubly serrate. Leaves turn bright

red in fall. Flowers are perfect, regular, and epigynous, developing in corymbose cymes that terminate

short lateral spurs. The clusters of fragrant white

blossoms sometimes appear to cover the crown of the

plant. Fruits are lustrous black, glabrous pomes about

0.25 inch (0.6 cm) in diameter that resemble small

apples. They are fleshy, dry, and mealy, and contain

light yellow pulp with one to five seeds. Flowering

occurs in May and June; fruits ripen from July to

September (Brinkman 1974e; Hitchcock and others

1961; Welsh and others 1987). Fruits are dispersed by

gravity and animals.

Ecological Relationships and Distribution

There are approximately 300 species of Crataegus

distributed throughout the Northern Hemisphere. In

North America the genus is most abundant east of the

Rocky Mountains. However, approximately 8 to 10

species occur in the Rocky Mountain area (Preston

1968). Hawthorn taxonomy is extremely complex; the

genus consists of numerous species exhibiting great

variation. Hybridization is common, particularly in

the central and eastern United States. Characteristic

Figure 10—Douglas hawthorn is an excellent

wildlife species that often occurs in or near

riparian areas.

of the subfamily Pomoideae, their chromosome number is n = 17 (McArthur and Sanderson 1985).

Douglas hawthorn, the most common species in our

area, is distributed from Alaska east to Alberta and

the Dakotas and south to California and northern

Wyoming. It also occurs in Michigan and Ontario

(Hitchcock and others 1961). In sagebrush and ponderosa pine communities it is largely confined to riparian

zones where it may form dense thickets. It is well

adapted to deep, moist, organic soils, but may be found

on soils ranging from silty to rocky clay loams. Although it commonly grows on well-drained sites, Douglas hawthorn is capable of withstanding periods of

soil saturation (Stark 1966). The species is an alternate host for cedar apple rust and may be damaged by

leaf spot in drier locations (Krebill 1972; Stark 1966).

Plant Culture

Ripe fruits are hand collected. Freshly collected

fruits must be kept cool and not allowed to mold prior

to extraction. Seed can be separated from fleshy material with a Dybvig cleaner or blender and water followed by air drying and fanning. A variable percentage of the seed is usually unsound but cannot be

removed by flotation. Some of these may be separated

on a gravity table. There are about 15,050 seed per lb

(33,179 per kg) at 100 percent purity (see chapter 24).

Treatment with concentrated sulfuric acid has been

used to release dormancy. The appropriate length of

acid treatment varies from 0.5 to 3 hours depending on

the seedlot (King 1947; McKeever 1938). Effectiveness of acid scarification has not been consistent; acid

can penetrate the seed coats and damage the embryos

of fresh seed. Embryo dormancy is released by a 3 to

5 month wet prechilling (King 1947; McKeever 1938;

SEAM 1976). Because of the variability in dormancy

among seedlots, combinations of standard pretreatments may be only partially effective in its release.

Tetrazolium or excised embryo tests are often used as

indicators of seed quality because current germination tests are time consuming and not highly reliable.

Results of direct seedings have been erratic. Spot

plantings in desirable locations conserve seed and

provide developing seedlings with the best opportunity for establishment.

Douglas hawthorn is most reliably established from

planting stock. Container and bareroot seedlings are

easily propagated, handled, and planted with good

survival on adapted sites. In the bareroot nursery,

seed should be planted to provide a seedling density of

25 per ft2 (278 per m2

). Untreated seed should be

planted in early fall, but wet prechilled seed must be

spring planted (Hartmann and others 1990; Van Dersal

1938). Container seedlings of uniform size are most

economically produced from germinants (Landis and

Simonich 1984). Rooting of hardwood or softwood
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Chapter 22 Rosaceous Shrubs

cuttings is not recommended (Marchant and Sherlock

1984). Root cuttings have been used to propagate

cultivated ornamental Crataegus species (Wyman

1971). Seedlings may be planted in mixtures with

other shrubs, but should be protected from competition with perennial grasses and weedy species.

Uses and Management

Douglas hawthorn is a valuable shrub deserving

more frequent inclusion in revegetation projects. It

has been successfully used in conservation and wildlife projects to furnish soil protection and food and

cover for wildlife. It is well suited to mixed plantings

with other shrub species. The dense thickets provide

hiding and thermal cover food for birds, small mammals, bear, big game, and livestock. They add valuable

structural diversity on grassland sites. Winter mule

deer use was evaluated as moderate to heavy in

Washington (Brown and Martinsen 1959). Douglas

hawthorn receives moderate summer use by mule

deer (Kufeld and others 1973). Use by white-tailed

deer and livestock is variable (Van Dersal 1938).

Douglas hawthorn is rated as a valuable honey plant

and fruits are heavily used by birds.

Fruits of Douglas hawthorn are high in sugar, but

low in fats and protein. They were eaten fresh or dried

and used in pemmican by Native Americans. They are

sometimes used for jams and jellies (Craighead and

others 1963). Where adapted, Douglas hawthorn is an

excellent candidate for landscape plantings in parks

and recreation areas.

Natural reproduction from seed is often prevented

by heavy use and trampling in riparian areas where

game and livestock tend to congregate. Seedlings

develop slowly and should be protected from browsing

and competition with weeds and grasses during the

first two seasons. Young seedlings may be girdled by

rodents. Established plants are not easily damaged by

excessive browsing.

Varieties and Ecotypes

Although Douglas hawthorn exhibits many good

attributes, selection trials have been limited. Appropriate growth forms of Douglas hawthorn may be

selected for shelterbelt and windbreak plantings. The

deep rooted, thicket-forming plants are useful for

stabilizing mine spoil banks, riparian sites, and other

disturbances. They are resistant to beaver damage

(Marchant and Sherlock 1984). A number of hawthorn

species and varieties are native to riparian zones in

the Intermountain region and may be selected for the

revegetation of adapted sites (Hitchcock and others

1961). Columbia hawthorn is a small tree or shrub

with red fruit and thorns 1.5 to 3 inches (4 to 7 cm) long.

It is distributed from British Columbia and Alberta

south to Oregon, Idaho, and Montana. It grows well in

heavy clay soils over basalt. Yellow hawthorn is found

from central Utah east to Pennsylvania and southeastern Canada.

Fallugia paradoxa _______________

Apache Plume

Apache plume, the only species of its genus, is a

much-branched shrub 2 to 7 ft (0.6 to 2.2 m) tall

(fig. 11). Branches are white to straw colored with

flaking or scaly bark (Harrington 1964). Leaves are

described as often evergreen in the southern portion of

its range (Blauer and others 1975; Deitschman and

others 1974b; USDA Forest Service 1937) and persistent or deciduous in cooler areas (Munz and Keck

1959; Welsh and others 1987). Leaves are borne in

fascicles along the branches and are 1.2 to 1.8 inches

(3 to 5 cm) long. They are cuneate with revolute

margins and pinnately divided into three to seven

linear lobes (Welsh and others 1987). Leaves are

lepidote on both surfaces, light green above and rusty

below (Harrington 1964; Munz and Keck 1959; Welsh

and others 1987). The large showy, perigynous flowers, 1.2 to 1.8 inches (3 to 4 cm) broad, develop singly

or in few-flowered cymes. A typical flower has five

broad sepals alternating with five bracts and five

large white petals. Numerous stamens are inserted in

three series on the margin of the hypanthium. There

are 20 to 30 pistils. The hairy achenes are 0.25 inch (6

mm) long with plumose persistent styles that elongate to 1 to 2 inches (2 to 5 cm) at maturity (Deitschman

and others 1974b; Young and Young 1986). The distinct plumose styles resemble an Indian war bonnet,

hence the name “Apache plume.”

Figure 11—Apache plume is a resprouting relative of bitterbrush that often grows in dry washes.
562 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Flowers are bisexual or rarely staminate (McMinn

1951). Blauer and others (1975) found monoecious as

well as dioecious populations. At a study site near

Richfield, UT, Blauer and others (1975) encountered

shrubs with mostly perfect flowers. However, some

plants had well-developed stamens but rudimentary

and nonfunctional pistils. Other plants bore flowers

with well-developed pistils, but rudimentary and nonfunctional stamens. Plants grown from seed collected

at this location exhibited similar differences when

grown at a study location near Boise, ID (Shaw and

Monsen 1983b). McArthur (n.d.) considers many

populations to be functionally dioecious.

Ecological Relationships and Distribution

Apache plume is found on a wide variety of sites from

southeastern California to southwestern Colorado

and south into Mexico and western Texas, largely

outside the Great Basin (Deitschman and others

1974b). It is more abundant in the southern portion of

its range. Apache plume grows in a variety of plant

communities from the lower desert brush types up

through the pinyon-juniper woodlands to open ponderosa pine communities at elevations from 4,000 to

8,500 ft (1,200 to 2,550 m) (Dietschman and others

1974b; Monsen and Davis 1985). It is commonly found

in washes and ephemeral waterways, but also occurs

on rocky or gravelly slopes and alluvial fans. Plants

tend to be widely spaced, even when in pure stands.

Apache plume superficially resembles Stansbury

cliffrose; both have lobed leaves and clusters of long

plumed fruits. Unlike Stansbury cliffrose, Apache

plume produces functionally dioecious plants (Blauer

and others 1975; McArthur and others 1983b), and it

does not form symbiotic nitrogen fixing root nodules

with the actinomycete Frankia spp. (Klemmendson

1979; Nelson 1983; Righetti and others 1983). The

species occur in broadly overlapping areas (Benson

1957; Blauer and others 1975), and a few suspected

natural hybrids have been reported (Blauer and

others 1975). Some seeds have been produced by

artificial crosses, but no progeny have been produced

(Blauer and others 1975). Chromosome number for

Apache plume is n = 7 compared to n = 9 for Stansbury

cliffrose.

Plant Culture

Flower buds are formed on elongated peduncles

produced from vegetative growth of the current season. Buds may appear on terminal or lateral leaders or

from new root sprouts (Rehder 1940). Flowering begins as early as April but may extend into August

(Rydberg 1954; Van Dersal 1938). Plants flower from

April to June in California (McMinn 1951), and as

late as October in Arizona (Kearney and Peebles

1960). Plants established in southwestern Idaho flowered erratically from May until early October (Shaw

and Monsen 1983b). Time from the appearance of

floral buds to fruit ripening was 67 days in 1979 and 86

days in 1980.

Pollination is insect dependent; the flowers attract a

variety of colorful bees, ants, hornets, and beetles

(Blauer and others 1975). Good seed (achene) crops

occur at 1 to 3 year intervals (Deitschman and others

1974b). Because flowers of Apache plume appear in

late spring, seed crops are usually not damaged by

spring frosts. Determining the appropriate date to

harvest seed is difficult; seeds ripen unevenly over a

period of several months. Most flowers develop early

in the season, producing up to 90 percent of the annual

seed crop. As seeds ripen they are detached and dispersed by the wind. Consequently, the period for

collecting the maximum amount of seed may be rather

short, perhaps only 1 to 2 weeks in duration. Later

blooming is associated with summer rains.

Achenes turn from green to reddish as they ripen

and plumes sometimes change from reddish to white.

They are harvested by dislodging the fruits from the

bush. The entire fruit, including the achene and plume

is collected. Fruits comprise only 15 to 20 percent of

the collected material by weight. Unless the plumes

are removed with a barley debearder, the collected

material remains in a thick entangled mass that

cannot be seeded. There are approximately 546,000

seeds (achenes) per lb (1,203,712 per kg) at 100 percent purity. For seed purchases, acceptable purity is 80

percent and germination is 75 percent (see chapter 24).

Apache plume has been somewhat difficult to establish from direct seeding. A better understanding of the

species’ seed germination requirements and seedbed

ecology would facilitate its use. Seeds are nondormant

and germinate quite readily. Plummer and others

(1969) reported 60 and 70 percent germination after

60 days at temperatures of 32 and 38 ∞F (0 to 3 ∞C).

Deitschman and others (1974b) found seeds sown in

nursery beds germinated within 4 to 10 days.

Although the plumed seeds are wind dispersed and

adapted to shallow or surface seeding, broadcast seeding without seed coverage has not been a satisfactory

means of planting. Apache plume is generally included in shrub seed mixtures that are planted separately from grasses. The seeds are easily sown with

conventional drills if the styles are removed. Seed

should be planted 0.25 to 0.5 inch (6 to 12 mm) deep in

a firm seedbed. Fall seeding or seeding prior to the

season of most dependable soil water is recommended.

Rodents often gather and bury or eat seeds that have

fallen from shrubs as well as sown seed. Seeding in late

fall or winter when rodents are less active reduces

these losses.

Seedlings emerge and grow rapidly under favorable

conditions. They are reasonably drought tolerant,

especially those that persist through the first summer.
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Chapter 22 Rosaceous Shrubs

The small plants are not able to persist amid herbaceous competition.

Bareroot and container reared seedlings are easily

started and grow rapidly with irrigation, often flowering during the first growing season. When grown

under field or nursery conditions in northern climates,

the seedlings are mostly deciduous. Once the leaves

have fallen, the plants are dormant and can be lifted

and transplanted. The brittle stems must be handled

carefully to prevent damage. In addition, the root

system usually consists of a main root with few side

branches. Any damage to the roots can be detrimental

to field survival. Greenhouse-maintained plants tend

to remain evergreen. These must be adequately hardened or overwintered in a shade house and planted as

dormant stock or field survival will be reduced.

Uses and Management

Apache plume occurs throughout the dry, hot portions of the pinyon-juniper, big sagebrush, and southern desert shrublands. It is a useful species for these

difficult-to-plant areas and can be planted on southern

aspects and shallow soils. To date, Apache plume has

most frequently been planted on drier portions of

pinyon-juniper communities in central and southern

Utah. Seed of the more northerly-occurring ecotypes

is harvested for these projects. Once established

Apache plume also does well on sites within its range

occupied by green ephedra, Stansbury cliffrose, and

black sagebrush. It can be planted on benchland sites

where spiny hopsage, shadscale, blackbrush, low rabbitbrush, desert bitterbrush, littleleaf mountain mahogany, and singleleaf ash may occur. It is also adapted

to fourwing saltbush, winterfat, and roundleaf

buffaloberry sites. The species has been established

from seed or planting stock on big sagebrush and

antelope bitterbrush/bunchgrass ranges in central

and northern Utah, southwestern Idaho (Shaw and

Monsen 1983b), and northeastern Oregon (Edgerton

and others 1983), all outside its native range.

Apache plume does not form thick stands. When

seeded with other shrubs it quickly thins out to occupy

only specific microsites. This characteristic has also

been observed when it is planted on mine wastes and

road disturbances. Apache plume has been slow to

spread from seed, particularly when planted outside

its natural range. Few seedlings are noted, even following years of heavy seed production. On coarse

soils, vegetative spread occurs through root proliferation and root sprouting. Sprouts are more numerous

during years with good precipitation or following

physical damage to the plant. Individual sprouts may

attain heights of 1 to 2 ft (30 to 70 cm) in one season

(Monsen n.d.).

Although Apache plume’s palatability to cattle and

sheep is generally considered low, it is rated as good on

some ranges. It does not receive heavy summer browsing

by livestock on most sites. However, because of its fine

twigs and evergreen leaves, it is a fairly valuable winter

forage, particularly in the southeastern portion of its

range (Ferguson 1983; Monsen and Davis 1985; USDA

Forest Service 1937). The bushy habit provides good

cover for birds and mammals (Blauer and others 1975).

Ferguson and Frischknecht (1985) considered Apache

plume a secondary species for use on mine disturbances in northeastern Utah. It is not nodulated by the

nitrogen-fixing endophyte Frankia (Nelson 1983;

Nelson and Schuttler 1984). Because of its ability to

spread vegetatively, Apache plume can be used to

provide ground cover and soil protection in dry regions

where infrequent storms can cause considerable erosion (Thornburg 1982). It is capable of recovering by

vigorous crown sprouting. On native sites it spreads

naturally onto disturbed areas.

Apache plume provides diversity for landscape

plantings, particularly for dry, low maintenance situations. The white branches and the flowers and plumes

are showy and quite unusual. Under cultivation the

plant can be over irrigated and will not withstand

even partial shade. Increased sprouting and a thicker,

bushy habit can be obtained for landscape or erosion

control plantings by trimming or grazing the planting.

Apache plume has been cultivated as far north as

Massachusetts (Deitschman and others 1974b).

The Tewa Indians of New Mexico used bundles of the

smaller branches of Apache plume as brooms and the

larger straight branches as arrow shafts (USDA Forest Service 1937).

Varieties and Ecotypes

Ecotypic differences have been observed among

Apache plume collections currently under test. Shrubs

of different populations vary in size and growth habit.

Some populations appear better adapted to more northerly climates than others. Breeding system expression, palatability, and rootsprouting following fire also

vary. There are at present no released varieties of the

species.

Holodiscus discolor _____________

Creambush Oceanspray

Creambush oceanspray, also known as arrowwood,

hardhack, rockspirea, and mountain spray, is a highly

variable, but usually erect multiple-stemmed, deciduous shrub (fig. 12). Mature plants range from 2.5 to

10 (20) ft (1 to 3 [6] m) in height, with large tree-like

forms growing in coastal areas. Young twigs are fine

hairy; bark of the older stems is gray to deep grayish

red. Leaf buds are covered by two to three purplishbrown bud scales. The alternate leaves are 1 to 4

inches (2.5 to 10 cm) long and ovate to ovate lanceolate

with lobed to doubly toothed edges. Upper surfaces are
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Chapter 22 Rosaceous Shrubs

hirsute, while lower surfaces are pilose lanate to

lanate and sometimes glandular. The common name

creambush oceanspray is derived from the appearance of the diffuse, showy panicles of tiny, perfect,

perigynous, white to pink flowers. Panicles are 4 to 7

(12) inches (10 to 17 [30] cm) long and overtop the

shrub. Flowers are about 0.2 inch (5 mm) in diameter,

regular, and 5-merous; petals exceed the sepals. Dried

flower parts and panicle branches turn brown and

persist well into the winter. Fruits are light yellow

hirsute achenes approximately 0.1 inch (2 mm) long.

In communities where it exists, creambush

oceanspray is one of the first shrub species to leaf out

in spring (Orme and Leege 1980). Floral buds develop

in late spring and flowering occurs from late June to

late August, depending on location. Fruits mature in

late summer and may persist into fall. Seeds are wind

dispersed (Hitchcock and others 1961; Orme and Leege

1980; Stickney 1974a; USDA Forest Service 1937).

Ecological Relationships and Distribution

The genus Holodiscus existed in the Madro-Tertiary

geoflora of western North America. It is now disjunct across the tropics, and occurs in western North

America and Columbia. Creambush oceanspray is

distributed from British Columbia east to western

Montana and south along the coast to southern

California. It grows in northeastern Oregon, Idaho,

and northeastern Nevada (Hitchcock and others 1961;

Munz and Keck 1959; Stark 1966; USDA Forest Service 1937). Creambush oceanspray grows on a wide

variety of sites at elevations ranging from sea level to

7,000 ft (2,100 m). It is adapted to rich, deep soils of

coastal forests, riparian areas, and canyon bottoms. It

may also be found growing on dry, shallow soils or

rocky talus slopes and along streams in drier areas

(Patterson and others 1985). Creambush oceanspray

is associated with drier conifer types of the interior

and is a climax species in a number of forested communities (Cholewa and Johnson 1983; Cooper and

others 1987; Halverson and others 1986; Steele and

others 1981). An understory shrub layer of mallowleaf

ninebark and creambush oceanspray is common in

ponderosa pine and Douglas-fir forests in eastern

Washington, northeastern Oregon, northern Idaho,

and western Montana (Daubenmire and Daubenmire

1968; Steele and others 1981).

Creambush oceanspray occurs as a climax in open

stands of conifers; it survives fires and other disturbances by resprouting from buds on the root crowns to

become a member of early seral brushlands (Mueggler

1966; Stickney 1986). Mixed seral brushfields provide

valuable forage and cover for deer, elk, and other

wildlife. Regeneration from seed following burns is

often limited and seedlings develop slowly (Ferguson

1983; Wright and others 1979; Young 1983). However,

regeneration from seed may be important following

extremely hot fires (Stickney 1974a). Owens (1982)

found weight and length of current annual leader

growth increased directly with degree of canopy mortality. However, regrowth was slightly lower for complete canopy mortality than for the 51 to 99 percent

mortality class. Prescribed burning at 10 to 15 year

intervals has been recommended to rejuvenate brush

fields that have grown beyond the reach of browsing

animals (Orme and Leege 1980).

Plant Culture

Collection of creambush oceanspray achenes is tedious; supplies are rare and costly. Fruits are “combed”

from the branches by hand in late fall and conditioned

by air drying and screening out larger material with a

fanning mill. Achenes of oceanspray are among the

smallest of shrub fruits. Only a small percentage of the

seeds are normally sound; these are identified by

examining wet seed through a dissecting microscope

for the presence of an embryo (King 1947). Seeds

require wet prechilling at 34 to 36 ∞F (1 to 2 ∞C) for

about 120 days to break dormancy (Marchant and

Sherlock 1984; Young and Young 1986).

Seeding may be accomplished by broadcasting fresh

achenes over a rough seedbed in fall and allowing

them to be covered by natural soil sloughing. Achenes

may be mixed with seeds of other shrub species, but

they should not be sown with grasses or forbs. Spots to

be planted should be carefully selected to make best

use of the seed. Seeding results have been erratic.

Creambush oceanspray is readily propagated from

bareroot or container stock. Achenes should be fall

sown in the nursery or artificially prechilled and

spring sown. Cleaned achenes can be planted at

Figure 12—Creambush oceanspray in full bloom.
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Chapter 22 Rosaceous Shrubs

reasonably uniform spacings. They can be surface

sown and covered by dragging a lightweight chain over

the seedbed. Seedlings develop rather slowly and may

be lifted as 1-0 or 2-0 stock depending on size specifications and growing conditions. Container seedlings

are propagated by planting germinants or about 10

wet prechilled seeds in each container. Plants can be

grown from cuttings, but techniques have not been

refined (Everett and others 1978a). Marchant and

Sherlock (1984) achieved 19 percent rooting success

with softwood cuttings harvested in late June and

propagated under a greenhouse mist system and 12

percent rooting success with hardwood cuttings overwintered in a cold frame.

Uses and Management

Creambush oceanspray is generally not considered

a palatable shrub, particularly when alternative browse

species are available (Daubenmire and Daubenmire

1968; USDA Forest Service 1937). It is taken by deer

and elk; peak use occurs in fall and winter (Stickney

1974a). Use varies geographically depending on ecotype

and associated species. Creambush oceanspray receives year round use by elk and deer in eastern

Washington, where it forms an important component

of the available browse (USDA Forest Service 1937).

Garrison (1953) recommended that browsing be limited to 50 to 60 percent of current annual growth on

these sites. In the northern Rocky Mountains, use by

big game is limited, but the species can be important

on some winter ranges (Leege 1969; Stickney 1974a).

Use increases tremendously the first winter following

fires, but decreases the second winter (Leege 1969).

Palatability of creambush oceanspray is rated fair for

cattle and poor to fair for sheep (USDA Forest Service

1937). The species tends to increase on heavily grazed

sites (Hall 1973; USDA Forest Service 1937). The

quantity of the plant consumed in summer and fall in

these areas may be considerable due to its abundance

and accessibility.

Creambush oceanspray is a useful erosion control

plant. The species is adapted to steep, dry, rocky areas

with shallow, sandy to clay loam soils (Stark 1966).

Seedlings quickly develop a spreading, highly

branched fibrous root system. Their low palatability

and their ability to resprout following top damage

contribute to their value in stabilizing critical sites

such as roadcuts and fills (Hungerford 1984; Stark

1966). In some areas, creambush oceanspray is

grown as a horticultural species for its dark green

foliage, extremely long flowering period, and orange to

dull red fall coloration (Sutton and Johnson 1974). It

is one of the first shrubs to leaf out in the spring. The

small, dry fruits were eaten by Native Americans.

Seedlings establish readily from seed or transplant

stock, but grow at only a moderate rate. Consequently,

they require protection from grass and forb competition

for one or two growing seasons. Excessive browsing is

rarely a threat to developing seedlings, but they may

be girdled by rodents or trampled by big game or

livestock.

Varieties and Ecotypes

To date, selection work with Holodiscus species has

been limited. The erosion control and ornamental

potential of both creambush oceanspray and rockspirea

deserve further investigation.

Holodiscus dumosus ____________

Rockspirea

Rockspirea, bush oceanspray, or rock spray spiraea is

somewhat similar in appearance to creambush oceanspray, but has a lower and more compact growth habit

(fig. 13). The rounded plants branch from the base and

range from 3 to 12 ft (1 to 4 m) in height with crown

diameters of 5 to 10 ft (1.5 to 3 m) (Hitchcock and

others 1961; Sutton and Johnson 1974; Welsh and

others 1987). Inflorescences and leaves are smaller.

Leaves are decurrent along the petioles; blades are

shallowly or coarsely toothed, sometimes with a few

secondary mucronulate serrulations. Roots are densely

fibrous and spreading. Flowering occurs unevenly all

summer (Mozingo 1987). The species has n = 18

chromosomes (McArthur and Sanderson 1985).

Rockspirea is distributed from northeastern

California across central and southern Oregon and

Idaho to Wyoming and south to Chihuahua, Mexico

(Hitchcock and others 1961) at elevations from 2,500

to 12,000 ft (762 to 3,660 m) (Dittberner and Olsen

1983; Harrington 1964; Hopkins and Kovalchek 1983).

It occurs in a wide array of plant communities including

Figure 13—Rockspirea has smaller leaves and a

more compact growth habit than creambush

oceanspray; it often occurs on cliffs and rock

outcrops.
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Chapter 22 Rosaceous Shrubs

big sagebrush, pinyon, curlleaf mountain mahogany,

chaparral, aspen-lodgepole pine, spruce-fir, Douglasfir, white fir, and ponderosa pine forests (Sutton and

Johnson 1974; USDA Forest Service 1937). Within its

range it commonly occurs on dry to moderately dry

sandy or gravelly soils, rocky ridges, talus slopes, and

basalt outcrops (Alexander and others 1974; Hitchcock

and others 1961; Welsh and others 1987). It is frequently a codominant in climax communities, but has

been reported to occur as a pioneer species following

disturbances in a white fir/Rocky Mountain maple/

rockspirea habitat type, where it persisted as the site

recovered (Alexander 1985).

Cultural and management practices recommended

for creambush oceanspray may be generally applied

to rockspirea. King (1947) determined that although

there are about 5,400,000 achenes per lb (12,000,000

per kg), only about 7 percent were sound. These required wet prechilling at 41 ∞F (5 ∞C) for 18 weeks to

release dormancy. Heat generated by wildfires may

also stimulate germination.

The palatability and forage value of rockspirea generally receive low ratings (USDA Forest Service 1937).

Kufeld and others (1973) reported its use by mule deer

was moderate in fall and light during the remainder of

the year. It is used by birds and rabbits to some extent

(Sutton and Johnson 1974; Van Dersal 1938). Based

on results of clipping studies in Oregon, Garrison

(1953) recommended a maximum of 50 to 60 percent

use for sustained production. Present primarily on

summer ranges where other species receive preferential use, excessive grazing of rockspirea rarely occurs,

and, at least on some western juniper/big sagebrush

habitat types of eastern Oregon, the species tends to

increase when other species receive excessive use

(Ferguson 1983; Hopkins and Kovalchik 1983).

Rockspirea thrives on dry slopes and unstable hillsides. It has high potential for use in revegetating

disturbed sites (Johnston 1987; Stark 1966).

Seeding practices and propagation of nursery stock

are as described for oceanspray. Plants are occasionally planted as ornamentals because of their long

flowering period, attractive foliage, and compact

growth habit (Stark 1966; Sutton and Johnson 1974).

Use of the species to date has been seriously limited by

a lack of commercially available seed.

Management of the species is also as described for

creambush oceanspray. Little is known of the plant’s

regenerative ability following use, fire, or other

physical damage (McMurray 1987a). Sprouting capabilities have not been well documented. Some regeneration might occur from wind-dispersed seed or from

soil seed banks.

Varieties and Ecotypes

There are no releases.

Peraphyllum ramosissimum ______

Indian apple

Indian apple is a deciduous rounded shrub growing

to 6 ft (2 m) in height from numerous, gray-barked

basal stems (fig. 14). Fascicles of simple, nearly sessile

oblanceolate leaves 0.6 to 0.8 inch (15 to 20 mm) long,

and 0.2 to 0.4 inch (5 to 9 mm) wide are produced at the

ends of short lateral spurs of the season. Leaves are

glabrous above with a minute pubescence below. Perfect, 5-merous, epigynous flowers appear with the

leaves, singly or in clusters of two to five. Leaves may

be dropped in midsummer, possibly in response to

water or temperature stress. Flowers are regular,

perfect, and fragrant with spreading pink to rose

petals. The style and the 15 or more stamens inserted

on the edge of the hypanthium are exserted at anthesis. Fruit is a yellowish, bitter tasting, apple-like

pome, 0.5 inch (12 mm) in diameter containing four

seeds. In Utah, Indian apple flowers from April to

July. Fruits ripen and are gravity or animal dispersed

in July or August (Hitchcock and others 1961; Welsh

and others 1987).

Ecological Relationships and Distribution

Peraphyllum is a monotypic genus of the subfamily

Pomoideae distributed from southern and eastern

Oregon along the east side of the Sierra Nevadas

through central California and east to western Colorado

and northwestern New Mexico at elevations ranging

from 3,000 to 9,000 ft (900 to 2,700 m) (Dayton 1931;

Ferguson 1983). It grows from the sagebrush/grass,

pinyon-juniper, mountain brush, and oak/sagebrush

zones to the lower fringes of ponderosa pine forests

Figure 14—Indian apple is an intricately branched

shrub with potential for use in low maintenance

landscaping.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 567

Chapter 22 Rosaceous Shrubs

(Hitchcock and others 1961; Welsh 1982). Chromosome

number is n = 17 (McArthur and Sanderson 1985).

Historically, Indian apple appeared in the Conifer

Woodland Element of central Nevada’s Upper Miocene floras in an area where the Arcto-and MadroTertiary floras overlapped (Axelrod 1950).

Plant Culture

Seed is harvested by hand picking or beating the

fruits into containers. Fresh or dry fruits are macerated in a Dybvig or blender with water. Pulp that

cannot be separated by flotation may be removed

from the seed by fanning after the pulp is dried.

Approximately 6.5 to 10.3 lb (2.9 to 4.6 kg) of cleaned

seed may be obtained from 100 lb (45 kg) of fruit

(Swingle 1939). There are approximately 23,000 seeds

per lb (50,706 per kg) at 100 percent purity. Acceptable

purity is 95 percent and germination is 50 percent for

seed purchases (see chapter 24).

Seed requires a wet prechilling period of approximately 45 days at 38 ∞F (3 ∞C) to relieve dormancy

(Belcher 1985; SEAM 1976; Smith 1974b). Germination of seedlots tested by the Coeur d’Alene Forest

Service Nursery ranged from 60 to 80 percent (SEAM

1976). Seed testing techniques have been described by

Belcher (1985). Seeds remain viable for up to 5 years

if stored in a cool, dry, well ventilated container (Smith

1974).

The existence of distinct ecotypes, possibly restricted

to specific soil types, makes selection of appropriate

seed sources important. Seed should be fall planted on

medium textured, well-drained soils in adapted areas

(Stark 1966). Seeding may be accomplished by hand

planting or drilling with other shrubs, but separately

from grass and forb species. Little seed is available

commercially; production of bareroot or container planting stock may maximize the number of seedlings

obtained (Ferguson 1983).

Uses and Management

The forage value of Indian apple varies with accession and associated plant species. It is grazed by deer,

particularly in fall. The fruits are used by birds and

rodents. Indian apple can be incorporated into windbreak

and conservation plantings on arid lands. The thick

branching habit of the shrub offers cover for birds and

other small animals in winter, even after the leaves

have fallen. It also has potential for landscaping in

arid areas, along roadways, and in recreation areas

(Monsen and Davis 1985). It increases diversity on

adapted sites. Indian apple is highly persistent once

established, leading Blauer and others (1975) to suggest that it be used as a rootstock for related, but less

tenacious species.

Initial survival of Indian apple plantings is often

quite low. Seedlings grow slowly and may require

protection for up to 3 years following planting. Indian

apple may be entirely defoliated by grasshoppers when

their populations are high. However, plants can recover even after successive defoliations.

Varieties and Ecotypes

Indian apple accessions have received little testing.

When selecting ecotypes for revegetation, emphasis

should be placed on plant size, fruit production, growth

rate, and palatability.

Physocarpus malvaceous ________

Mallowleaf Ninebark

Mallowleaf ninebark is an open, multiple-stemmed

shrub 3 to 8 ft (1 to 2.5 m) tall (fig. 15) that spreads

vegetatively from long rhizomes (Hitchcock and others

1961; Welsh and other 1987). Roots are shallow and

spreading. Bark is gray to red brown, shredding in

thin layers. Leaves are alternate and currant- or

mallow-like with three to five palmate lobes. Leaves

are doubly crenate, nearly glabrous above, stellate

pubescent to glabous below, and about 0.75 to 3.2

inches (2.0 to 8.0 cm) long. Clusters of stellate white

flowers develop on the terminal corymbs. Flowers are

regular and perigynous. Fruits are paired, densely

stellate, flattened follicles with ascending beaks that

dehisce along two sutures. Each follicle contains 1 to

several tiny, hard, shiny, pyramiform seed each

about 0.1 inch (2 mm) long (Gill and Pogge 1974a).

Mallowleaf ninebark flowers in May or June and sets

seed in August or September. Some fruits persist on

the plant through the winter.

Figure 15—Mallowleaf ninebark grows rapidly;

its dense root systems stabilize soil on unstable

disturbances.
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Chapter 22 Rosaceous Shrubs

Like other shrubby members of the subfamily

Spiraeoideae native to western North America, the

chromosome number for mallowleaf ninebark is n = 9

(McArthur and Sanderson 1985). Species with n = 7

and n = 9 occur among Asian taxa of the genus

(McArthur and others 1983b).

Ecological Relationships and Distribution

Mallowleaf ninebark occurs east of the Cascades

from British Columbia and Alberta south through

Nevada, Utah, and Wyoming. It grows in association

with mountain big sagebrush, aspen, and many conifer communities at elevations from 5,200 to 10,800 ft

(1,560 to 3,240 m) (Welsh 1982). It may be found on

dry open slopes, talus slopes, northern exposures, in

scattered to heavy timber, and on riparian sites (Davis

1952). Mallowleaf ninebark is adapted to deep, welldrained organic soils of medium texture and pH of

7.0 to 7.5 (Marchant and Sherlock 1984), but it will

grow on drier, sandy or rocky soils (Daubenmire and

Daubenmire 1968; Ferguson 1983). It is often abundant and highly productive.

Mallowleaf ninebark is frequently a member of

seral shrub communities that develop following logging or wildfires in conifer forests (Olsen and Nagle

1965; Owens 1982). The species is highly adapted to

fire; it resprouts from the root crown or horizontal

rhizomes. The more completely buried rhizomes have

the greatest resprouting capacity (Bradley 1984).

Owens (1982) and Merrill (1982) found that number of

plants and twig production increased on burned sites

relative to unburned sites. In addition, recovery is

enhanced by arrival of seed from undisturbed sites.

The light seed is dispersed over long distances by wind

and gravity (Young 1983).

Seral shrub communities provide forage and cover

for deer, elk, and other wildlife, but rapidly grow out of

reach of browsing animals. Overmature brushfields

have been rejuvenated by prescribed burning, cutting,

or topkilling tall shrubs with herbicides (Merrill 1982;

Mueggler 1966; Orme and Leege 1980). Control of

brush species to promote conifer regeneration has also

been accomplished with herbicides (Miller 1981; Miller

and Pope 1982a,b).

In addition to mallowleaf ninebark, several other

ninebark species occur within the Intermountain region. Dwarf ninebark occurs in pinyon-juniper and

related habitats of Idaho, Nevada, Colorado, and Utah.

Mountain ninebark is found on moist soils of aspen

and coniferous forests from Nevada east to Wyoming

and south to Arizona and New Mexico. Pacific ninebark

also grows on moist soils, but is restricted to California

and the area west of the Cascades in Oregon and

Washington. A disjunct population occurs in northern

Idaho (Ferguson 1983; Hitchcock and others 1961).

Plant Culture

Mallowleaf Ninebark seed are hand collected by

detaching the dried flower heads (Van Dersal 1938).

Seed is separated from debris using a barley debearder

and an air screen fanning machine. There are approximately 756,000 seed per lb (1,666,700 per kg).

Acceptable purity for seed purchases is 98 percent

and germination 40 percent (see chapter 24). Seed is

expensive and rarely available. Little is known of the

seed biology of this species. A high percentage of the

tiny seeds are normally not sound. Sound seeds are

dormant and require a 77-day prechilling at 40 to 43 ∞F

(4 to 6 ∞C) (Gill and Pogge 1974a; Marchant and

Sherlock 1984).

Direct seeding is accomplished by broadcasting

seed onto a rough seedbed in fall and allowing natural

soil sloughing to cover it. Best results are obtained

when seed is planted separately from other species in

selected spots. Seedlings can establish with some

shade and limited herbaceous competition.

For bareroot nursery plantings, mallowleaf ninebark

is fall seeded or artificially wet prechilled and spring

seeded. Seed should be covered lightly and mulched.

Swingle (1939) estimated that a pound of seed would

yield about 30,000 usable seedlings. Seedlings are

lifted as 1-0 to 2-0 stock depending on purchase specifications and growing conditions. Container stock may

be propagated from germinants or cuttings. Hardwood stem cuttings are easily propagated and frequently used. Container and bareroot stock develop

slowly, producing densely branched root systems. Both

container and bareroot seedlings establish readily

(Monsen n.d.).

Uses and Management

The value of mallowleaf ninebark as a forage plant

varies with season, location, class of game or livestock

being considered, and availability of other vegetation.

It is generally considered to be more valuable on

ranges where better forage plants are depleted or

unavailable, and on drier sites (Noste and Bushey

1987). Stands are rarely lost as a result of browsing.

The USDA Forest Service (1937) rated mallowleaf

ninebark as fair for sheep, fairly good for goats, and

poor for cattle; Van Dersal (1938) rated it as good

browse for sheep, goats, and cattle. Kufeld and others

(1973) reported it receives light spring use and moderate summer and fall use by mule deer. In cafeteria

style feeding trials with tame mule deer, Smith (1953)

found mallowleaf ninebark was one of the most highly

preferred native forages in the Logan, UT, area from

May 1 to June 20. Because of its deciduous habit, it is

of little value on winter ranges. New growth is more

heavily browsed by deer for up to 3 years following

burning (Noste and Bushey 1987).
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Chapter 22 Rosaceous Shrubs

Mallowleaf ninebark has potential value as an erosion control plant. It grows rapidly, producing a network of rhizomes and a densely branched fibrous root

system. It is a useful species for the restoration of

high elevation acid minespoils in Nevada (Butterfield

and Tueller 1980; Everett and others 1980). In spite of

its rapid growth rate, mallowleaf ninebark receives

little use in windbreak and farmstead plantings because of its bushy growth habit (Olsen and Nagle

1965). Mallowleaf ninebark has gained acceptance as

an ornamental in some areas. It is a hardy, easily

grown plant with attractive white flowers and dull to

bright reddish brown leaf coloration (Gill and Pogge

1974a; Sutton and Johnson 1974). Plants that become

too bushy can be pruned back.

Once established, seedlings of mallowleaf ninebark

grow rapidly compared to those of many other members of the Rosaceae. They compete successfully with

other shrub species, but require some protection from

grass competition for one or two growing seasons.

Seedlings are not browsed heavily if alternative forage is available, but they may suffer predation by

rodents and trampling by big game and livestock.

Potentilla fruticosa ______________

Shrubby Cinquefoil

Shrubby cinquefoil, also known as bush cinquefoil,

buckbrush, hardhack, ninebark, or yellow rose, is a

rather short lived, densely branched, round-topped to

low spreading shrub with spreading and fibrous

roots. It produces white to bright yellow flowers

throughout the summer, when adequate soil water is

available (fig. 16) (USDA Forest Service 1937). The

shreddy brown bark of the main stems peels off in long

strips. Leaves are pale to gray-green, leathery, 0.4 to

1.2 inch (1 to 3 cm) long, and pinnately compound

with five to seven narrow, often revolute, leaflets.

Leaves turn yellow in fall, but may be retained over

winter. Flowers are 5-merous, complete, and solitary

or in three to seven-flowered cymes. Each flower is

0.75 to 1 inch (2 to 2.5 cm) in diameter with a saucershaped hypanthium and five petals. There are 10 to 30

stamens and numerous pistils. Fruits are golden tan,

teardrop-shaped achenes crowded on a hairy receptacle. Individual achenes are 0.04 to 0.08 inch (1 to 2

mm) long with very thin seed coats covered with white

hairs. Fruits mature and are dispersed continuously

by wind and gravity throughout the flowering season

(Hitchcock and others 1961). However, in Alberta,

Canada, flowering apparently occurs in two flushes,

one in May and the other in August (Scotter 1975).

Fruits mature in mid to late summer and fall.

Ecological Relationships and Distribution

There are about 250 species of Potentilla occurring

in temperate to subarctic regions of the Northern

Hemisphere (Young and Young 1986). Shrubby cinquefoil is widely distributed in the Northern Hemisphere,

ranging east from Alaska’s north slope to Greenland

and Labrador and south to California, New Mexico,

Minnesota, and New Jersey. It also grows in Europe

and Asia (Hitchcock and Cronquist 1973; USDA Forest Service 1937). In common with several mesic

shrubby North American members of the subfamily

Rosoideae, the diploid chromosome number of shrubby

cinquefoil is 2n = 14 (McArthur and Sanderson 1985);

that of European plants is 2n = 28 (Elkington and

Woodell 1963). Hitchcock and others (1961) reported

hybridization occurs within this genus, but hybridization of shrubby cinquefoil with other species has not

been observed (Elkington and Woodell 1963).

Shrubby cinquefoil is associated with cool climates

and wet areas; it grows from plains and lower foothills to above timberline over an elevation range of

6,100 to 12,000 ft (1,850 to 3,600 m) (Belcher 1985). In

the western United States shrubby cinquefoil is a

common summer range plant in wet subalpine meadows. It also grows near springs and marshes and along

streambanks and floodplains in chaparral, sagebrush,

aspen/lodgepole pine, ponderosa pine, Douglas-fir/white

Figure 16—Shrubby cinquefoil is a common riparian species in wet meadows at mid to upper

elevations.
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Chapter 22 Rosaceous Shrubs

fir, and spruce/fir forests (Sutton and Johnson 1974;

Welsh 1982; Welsh and others 1987). It is most

abundant on areas with deep, wet, medium-textured

soils and can withstand some salinity (Davis 1952;

Stark 1966).

The response of shrubby cinquefoil to burning depends on the season, site conditions, and fire intensity,

and may vary among populations. Plants may be

relatively unaffected to heavily impacted by burning

(Fischer and Clayton 1983; Mueggler and Stewart

1980). Little damage is incurred by low intensity

spring burns that fail to kill the root crown (Wasser

1982), particularly on wet or rocky sites. Plants

regenerate rapidly by sprouting from root crowns,

rhizomes, and stem layering (Wasser 1982). Severe

burns occur when dense mature stands burn in late

summer, particularly under droughty conditions.

Some soil seed reserves may also be lost under such

circumstances with the result that recovery occurs

very slowly.

Plant Culture

Seed collection is difficult. Fruits are scattered over

the crown of the plant and ripen unevenly during the

growing season. Plants are not shade tolerant and

produce little seed when shaded (Scotter 1975). Little

seed is mature at any given time and hand collection

proceeds slowly. Consequently, seed is rarely available commercially (Ferguson 1983). Seed is cleaned

with a debearder and fanning mill. It may be stored in

a warehouse for 5 years. Seventy percent is considered

an acceptable level for both purity and germination

for seed purchases (see chapter 24). Seed quality is

highly variable, but often rather low (Elkington and

Woodell 1963; Wasser 1982).

Little information is available regarding germination requirements of shrubby cinquefoil seed. Vories

(1981) reported the species is difficult to grow from

seed. Belcher (1985) and SEAM (1976) found no germination pretreatment was required although a water

soak at 86 ∞F (30 ∞C) for 18 hours did enhance germination. McDonough (1969) applied an 11-minute soak

in sulfuric acid and obtained 73 to 87 percent

germination at 90/72, 72/63, and 63/53 ∞F (32/22, 22/

17, and 17/12 ∞C). Light is required during incubation

(Young and Young 1986). Tetrazolium and x-ray testing instructions were provided by Belcher (1985).

Direct seeding information is scarce, but seeding can

be successful on sites with good soil water. Babb (1959)

recommended planting as soon as seed ripens. Planting before mid-July or in late fall was recommended by

Wasser (1982). Seeding may be accomplished by drilling, broadcasting, or spot seeding on prepared sites

(Wasser 1982). Seeds are small and should be covered

lightly, about 0.25 inches (6 mm) deep or less (Wasser

1982).

Bareroot and container stock can be produced from

seed. Container stock started from germinants or

small transplants requires a 3 to 5 month cropping

season (Landis and Simonich 1984). Wildings are an

alternative source of planting stock.

Production of shrubby cinquefoil for the commercial

ornamental market generally involves vegetative

propagation of horticultural varieties. Plants are capable of natural root sprouting following disturbances

(Wasser 1982). Stem layering also occurs and may be

effective in spread and survival where inundation

occurs (Elkington and Woodell 1963). Vegetative propagation is accomplished by rooting softwood or

semihardwood stem cuttings in a greenhouse. Root

cuttings, layers, and crown divisions are also propagated (Doran 1957; Mahlstede and Haber 1957; Toogood

1980).

Uses and Management

Shrubby cinquefoil is an indicator of heavy grazing;

it may become more abundant when other species are

depleted. It is outcompeted by grasses and declines if

grazing is eliminated. Although its palatability is

often low due to its coarse, astringent foliage (Dayton

1931), game and livestock will use shrubby cinquefoil

when other species are not available (Wasser 1982).

Young plants are generally more palatable than mature shrubs (Wasser 1982). In general, it is more

palatable to sheep and goats than to cattle and horses

(Sampson and Jesperson 1963). Because of its higher

elevational distribution, it normally occurs on summer ranges, but where available in winter, it is sometimes used because of its semi-evergreen habit and

dominance in some areas (USDA Forest Service 1937).

Kufeld and others (1973) found mule deer make

light use of shrubby cinquefoil in winter and take only

trace amounts in spring and summer. Elk and whitetailed deer make some use of the plant during all

seasons. Bighorn sheep use it primarily in autumn

(Van Dyke and others 1983). Seeds are consumed by

birds and small mammals (Wasser 1982). Palatability

of shrubby cinquefoil varies geographically and

seasonally. It is generally more palatable in spring

and summer (Plummer and others 1968). Although it

is avoided by big game in Alberta, it is unusually

palatable in parts of Montana and Arizona. Livestock

also make greater use of it on scattered high altitude

summer ranges of the southwestern states, southwestern Montana, southeast Idaho, and Utah (Dayton

1931; Ferguson 1983; USDA Forest Service 1937).

Plants are moderately tolerant of browsing, but they

can be damaged by repeated grazing, particularly

during periods of drought when plants may become

hedged and stunted and eventually lost (USDA Forest

Service 1937; Wasser 1982).
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Chapter 22 Rosaceous Shrubs

Whether transplanted or grown from seed, shrubby

cinquefoil is a useful soil stabilization species for

disturbed sites with fairly wet soils. It is useful for

mine reclamation, roadway plantings, and game

range rehabilitation (Dittberner and Olsen 1983;

Fedkenheuer and others 1980; Ferguson and

Frischknecht 1985; Stark 1966; Wasser 1982). It is an

excellent species for ornamental plantings because of

its rapid growth rate, ease of cultivation, attractive

semi-evergreen leaves, colorful flowers, and long

flowering period.

Although shrubby cinquefoil seedlings have been

described as weak, they begin to grow rapidly once

established and are better able to survive herbaceous

competition than most other rose species. Plants are

cold, but not drought tolerant (Sutton and Johnson

1974). If planted in areas not heavily grazed by game

or livestock, they develop to maturity in about 5 years

(Wasser 1982).

Varieties and Ecotypes

Considerable natural variability has been observed

in this species. Numerous ornamental varieties of

shrubby cinquefoil are marketed commercially. However, the species has received little testing for wildland uses. Populations with a rapid growth rate,

spreading growth habit, and low palatability should

be of greatest value for soil stabilization and

streambank erosion control. Numerous showy flowers, a long flowering period, low palatability, and

evergreen leaves are desirable characteristics for populations used for landscaping in recreation areas.

Prunus americana _______________

American Plum

American or Pottawatami plum is an erect shrub or

rarely a small tree. In the Great Plains it grows

from 9 to 26 ft (3 to 8 m) in height (fig. 17) (Harrington

1964; Hitchcock and others 1961). In Utah plants are

thicket-forming shrubs up to 16 ft (5 m) tall (Welsh

and others 1987). Regeneration by rhizome sprouting is the main method of spread (McMurray 1987b;

Welsh and others 1987). Branches are thornlike at the

tip. Leaves are 1 to 2.8 inches (2.5 to 7 cm) long, elliptic

to ovate or lanceolate and sharply long attenuate.

They have pubescent, glandular petioles. Small umbels of white flowers are borne before the leaves on

spur branchlets or axillary buds formed the previous

season. Fruits are fleshy yellow to red drupes 0.8 inch

(2 cm) in diameter, each with a single large stone.

Some fruits may be hard and dry (Welsh and others

1987). Seed are dispersed by gravity and frugivorous

mammals.

Figure 17—In the Intermountain area, American

plum is a thicket-forming shrub that spreads from

rhizomes.

Plant Culture

American plum ranges from Manitoba through northeastern Canada and south to New Mexico and northwestern Florida (Grisez 1974; Hitchcock and Cronquist

1961). It occurs in the Rocky Mountain region and has

been introduced into portions of the Intermountain

area west of its native range. It will grow on deep soils

in mountain brush, pinyon-juniper, and sagebrush

grass habitat types receiving at least 16 inches (40 cm)

of annual precipitation (Plummer and others 1968;

Thornburg 1982). Within its native range it is often a

climax dominant or codominant, but it may be an early

seral species in some forested habitat types (Elkington

and Woodell 1963). It acts as a colonizer on unstable

sites along streams or rivers (Elkington and Woodell

1963; Mueggler and Stewart 1980). Where introduced

in the Intermountain region, it often grows in sandy to

loamy soils along ditch banks, drainage areas, and

around home or farm buildings where supplemental

water is available. Plants are capable of withstanding

some salinity.

American plum does not establish well from direct

seeding. However, it can be established from bareroot

or container stock transplanted singly or in rows or

clusters to meet planting objectives. Rootstocks are

utilized for propagation of domestic plum in northern

climates (Grisez 1974). American plum grows well in

mixtures with other woody species. Herbs with some

shade tolerance do well as understory species.

American plum will sprout when burned, trampled,

or otherwise damaged. Spread also occurs from seed

transported by animals. The species has good drought

and excellent cold tolerance.
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Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution

American plum flowers in April or May and fruits

ripen in September or October (Grisez 1974; Plummer

and others 1968). Fruits are harvested, conditioned,

and stored as described for common chokecherry.

Seed requires a 60 to 150 day wet prechilling to

relieve dormancy (Grisez 1974). There are about 810

seed per lb (1,786 per kg) at 100 percent purity.

Acceptable purity is 98 percent and germination 70

percent for seed purchases (see chapter 24).

Uses and Management

American plum has been planted for many years in

conservation plantings, windbreaks, and shelterbelts

in the Midwest and Western States. It provides excellent escape, nesting, thermal, and travel cover for big

game, livestock, and small mammals and birds. Leaves,

twigs, and fruit are utilized by these animals.

American plum can be an important component of

erosion control plantings and low maintenance landscape projects. It has excellent potential for use in

recreation areas, roadside rest areas, roadways, and

administrative sites. It is also used as an aesthetic

screen for wildlife in areas requiring thermal and

concealment cover.

Varieties and Ecotypes

A number of American plum cultivars have been

selected, primarily for their fruiting characteristics.

There are no selections specifically developed for wildland use. Available material varies considerably in its

range of adaptability and suckering habit. Stock

adapted to the planting site and suitable for the

planting goals should be acquired.

Prunus andersonii_______________

Desert Almond or Anderson Peachbrush

Mozingo (1987) described desert almond as “one of

the most beautiful and probably the most unappreciated shrubs in the western Great Basin.” Also known

as desert peach, Anderson almond, or Nevada wild

almond; desert almond generally occurs as low growing, densely shrubby, clonal plants. Each clone may

cover several acres. Individual stems are evidently

short lived, usually producing 6 to 8 annual rings

(Kay and others 1977f; Young and Evans 1973, 1974).

The plants have an average height of about 3 ft (1 m),

but range from 1.5 to 6.5 ft (0.5 to 2 m) in height

(fig. 18). They are intricately branched and densely

thorny; each branch ends in a sharp spine. Narrow

leaves develop in fascicles on short, lateral branches.

Small pink to light red flowers appear in April to

June and cover the shrub. The flowering period for

Figure 18—Desert almond is a clonal shrub

valuable for soil stabilization, cover, and as a

low water-use ornamental.

separate clones within an area may vary by up to a

month, ensuring that some flowers and fruits will

survive late frosts (Kay and others 1977e). Fruits

ripen in June or July. The drupes resemble small, dry,

fuzzy greenish-orange peaches and have a thin, rather

dry fleshy layer. The thick hard coat of the stone opens

easily along one suture (Mozingo 1987). Fruits and

seeds are dispersed by gravity and wildlife.

Ecological Relationships and Distribution

The native range of desert almond extends from

northeastern California southward along the eastern

slopes of the Sierra Nevada Mountains and eastward

across western and southern Nevada at elevations

from 3,450 to 7,400 ft (1,050 to 2,250 m) (Ferguson

1983; Munz and Keck 1959; Young and Evans 1973).

It is a fairly drought tolerant species, most commonly

found scattered through big sagebrush and pinyonjuniper communities on foothills, mesas, alluvial terraces, and in canyons. It is especially common on coarse

sandy or gravelly to rocky soil derived from decomposing granite (Dayton 1931; Smith and others 1978).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 573

Chapter 22 Rosaceous Shrubs

Plant Culture

Seedling establishment apparently occurs only

infrequently. Seedlings have been noted following

removal of woody species in sagebrush/grass and

pinyon-juniper sites (Young and Evans 1973, 1974);

they develop from soil seedbanks or from off-site seed

transported by mammals (McMurray 1987c). Plants

resprout vigorously following fire (Young and Evans

1978b). Clones may survive into later seral stages of the

community; maximum occurrence is on old burns that

have reached mid-successional stages (Koniak 1985).

Fruits are hand collected. Conditioning involves

macerating the fruits in a Dybvig or blender and

floating the pulp. To relieve dormancy, Kay and others

(1977f) recommended a 30 day wet prechilling; 60 days

were recommended by SEAM (1976). Seed should be

fall planted to provide for field prechilling. Container

and bareroot seedlings have been used successfully.

Planting stock can also be propagated from cuttings,

although the success rate may not be high. Everett and

others (1978a) recommended use of cuttings collected

during the period of seed maturation. Rooting required approximately 6 weeks under a greenhouse

mist system.

Uses and Management

Desert almond has good potential for use as a soil

stabilization species on disturbed sites due to its low,

dense growth form, clonal habit, and good establishment. Everett (1980) obtained second year survival of

77 percent and 25 percent for containerized desert

almond seedlings planted on south- and north-facing

roadcuts, respectively, near Reno, NV. Seedlings were

tenacious, but tended to grow slowly. Smith and others

(1978) obtained 3-year survival of 73 percent when

desert peachbrush seedlings grown in containers

1.5 ft long (0.5 m) were transplanted in holes drilled

into granitic bedrock near Mammoth Lake, California.

They recommended the species for granitic sites within

its range.

Desert almond foliage is somewhat palatable, accessible, and probably seasonally important. Livestock

and wildlife make some use of the species, particularly

in spring or when summer storms induce new leaf

production (Dayton 1931). The dense patches provide

good cover for birds and other small animals. Desert

almond has good potential as a low maintenance ornamental for roadsides, parks, and campgrounds because of its colorful flowers and long flowering period.

Varieties and Ecotypes

No improved varieties are available for revegetation

projects. Some variability among clones has been

noted, but comparative testing of different populations has been limited. Desert almond has been grown

as far north as Boise, ID, (Monsen n.d.).

Prunus besseyi _________________

Bessey Cherry

Bessey cherry or western sand cherry is a lowgrowing shrub with numerous spreading basal stems

1 to 5 ft (0.3 to 1.5 m) tall (fig. 19) (Vories 1981; Welsh

1982). Leaf blades are 0.6 to 2 inches (1.5 to 5.0 cm)

long and 0.2 to 0.7 inch (5 to 18 mm) wide, elliptic to

oblanceolate, and acute to cuspidate apically. Leaves,

sepals, petals, hypanthia, and fruits are glabrous.

White flowers appear with the leaves in clusters of two

to four. The black fruits are 0.5 to 0.7 inch (12 to 18

mm) in diameter. Flowering occurs in April or May;

fruit ripening and dispersal occur from July to September in Nebraska (Grisez 1974), and from August to

September in Utah (Plummer and others 1968).

Ecological Relationships and Distribution

Bessey cherry is native to the Great Plains and is

distributed from Manitoba south to Wyoming,

Colorado, and Kansas (Grisez 1974). The species is

commonly used as a dwarfing rootstock for other

Prunus species. It has been introduced into the

Intermountain region in wildlife and conservation

plantings and by dispersal from orchards. Bessey

cherry has been successfully established on sites

ranging from basin big sagebrush communities to

openings in ponderosa pine forests. Adkins (1980)

reported survival in conservation plantings in eastern

Washington was 28 percent in the 9 to 12 inch (23 to

30 cm) precipitation zone; survival ranged from 3 to

61 percent when plants were fully established. He

concluded that in low precipitation areas Bessey

cherry produced inadequate growth to provide good

cover and was unable to compete successfully with

cheatgrass and other annual and perennial weeds.

Figure 19—Native to the Great Plains, Bessey

cherry has been used in wildlife and conservation

plantings in the Intermountain region.
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Chapter 22 Rosaceous Shrubs

Plant Culture

Seed collection, conditioning, storage, and planting

practices are generally as described for common

chokecherry. Seed is often collected from stock maintained at a nursery or from conservation plantings.

Swingle (1939) found an average of 17 lb (7.6 kg) of

seed could be recovered from 100 lb (45 kg) of fruit

yielding 1,500 to 2,260 clean seed per lb (3,300 to

4,980 per kg). Babb (1959) obtained greatest germination following a 100 day wet prechilling at 41 ∞F

(5 ∞C) or 120 days at 33 ∞F (1 ∞C); he noted a high

degree of variability in requirements among seedlots. A 72 hour gibberellic acid soak (3.46 g Rootone F/

1000 g H2O) followed by wet prechilling for 20 days

at 41 ∞F (5 ∞C) was recommended by SEAM (1976).

Bessey cherry can be direct seeded. Seed should be

fall planted 0.5 to 1.0 inch (1.0 to 2.5 cm) deep on

seedbeds cleared of competition. The species can also

be established from container or bareroot stock (Monsen

1974). It can be propagated frombuds, grafts, suckers,

or root cuttings (Babb 1959). For windbreaks or conservation plantings, a 4 ft (1.2 m) spacing is recommended (Cook 1981).

Uses and Management

Monsen (1974) and Monsen and Christensen (1975)

recommended use of Bessey cherry to provide good

ground cover and soil stabilization on adapted sites in

Utah and Idaho. It has been used extensively in the

Midwest as a low growing erosion control and wildlife

plant; it provides fruits and cover for birds and other

small animals. Attractive flowers, fall leaf coloration,

and its low, spreading growth habit make Bessey

cherry a unique plant for landscaping.

Rodents and birds often collect planted seed and

emerging seedlings. Severe damage to planted Bessey

cherry seedlings has resulted from rodent girdling

(Brown and Martinsen 1959). Plants developing from

seed or planting stock should be protected from excessive grazing and competition with grasses and forbs.

Varieties and Ecotypes

Some selections have been developed for horticultural plantings, but no specific material has been

developed for wildland uses.

Prunus emarginata ______________

Bitter Cherry

Hitchcock and others (1961) recognized two varieties of bitter cherry: Prunus emarginatavar. mollis, an

arborescent variety growing to 42 ft (15 m) in height

and distributed west of the Cascade Mountains, and

var. emarginata, a more shrubby form with an open

growth habit and several crooked stems commonly

3 to 13 (26) ft (1 to 4 [8] m) tall that grows in and east

of the Cascade Mountains (Hitchcock and others 1961).

The deep reddish-purple bark on young twigs of both

varieties turns gray to dark brown on older branches

and stems. The entire, alternate leaves are 1.2 to 2

inches (3 to 5 cm) long, glabrous to pubescent, and

oblong to obovate with crenulate to serrate margins.

Stipules are caducous and attenuate to linear. Corymbs

of 3 to 12 white insect-pollinated flowers (fig. 20) are

produced from axillary buds on twigs of the previous

season. Flowers are complete, regular, perigynous, and

5-merous. Sepals are produced on a turbinate, deciduous, disk-lined hypanthium that varies from glabrous

to hairy. There are 12 to 20 exserted stamens and a

single pistil. Only one of the two ovules develops. The

fruit is a bitter tasting, red to black, one-seeded drupe

from 0.3 to 0.5 inch (8 to 12 mm) in diameter. Bitter

cherry flowers from April to June. Fruits ripen from

July to September depending on elevation and local

weather conditions (Grisez 1974; Plummer and others

1968). Fruits are dispersed in August or September.

Seed are often distributed by birds and mammals.

Figure 20—Bitter cherry flowers from April to June,

depending on elevation. Fruits are used by many

birds and mammals.
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Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution

Bitter cherry occurs from the Pacific Coast of British

Columbia east to northern Montana, and south to

southern California and southwestern New Mexico

(Ferguson 1983; Hitchcock and others 1961). Its

elevational range varies with latitude. In the north it

grows from sea level to 3,000 ft (915 m); in the southwest it occurs from 5,000 to 9,000 ft (1,525 to 2,745 m)

(USDA Forest Service 1937).

Bitter cherry is most frequently found in the upper

portion of the ponderosa pine belt. It is frequently

found on well drained, moderately fertile, rocky sites

such as ridge tops and southwest slopes in mountain

brush communities. It is capable of resprouting

following burning and often dominates other shrubs

in seral brushfields. Bitter cherry grows along watercourses in grasslands and sagebrush communities.

Plant Culture

To maximize the number of viable seed obtained,

fruits should not be harvested until they are bright red

and fully mature (Grisez 1974). Fruits are collected by

hand stripping or by placing tarps under trees to catch

them following natural dispersal. They may also be

harvested with a mechanical tree shaker. Swingle

(1939) reported that about 25 lb (11.4 kg) of seed can

be obtained from 100 lb (45.5 kg) of fruit yielding

1,800 to 5,471 seeds per lb (3,965 to 12,050 per kg).

A Dybvig or commercial blender may be used to

macerate the fruit. All pulp and juice should be cleaned

from the stones. Part of the pulp and some low quality

seeds can be removed from the mixture by flotation,

the rest by drying and fanning. Dried fruits are cleaned

by hammermilling at a low speed, followed by fanning.

Specific storage requirements have not been ascertained for bitter cherry (Grisez 1974); seeds are commonly stored as described for common chokecherry.

Bitter cherry seed fill is generally low. Filled seeds

are difficult to germinate. Treatments to increase

germination give inconsistent and generally poor results. Germination of treated seed is generally less

than 20 percent (Monsen and Davis 1985). Poor germination has discouraged use of this species in direct

seedings. The species can be intertransplanted using

bareroot or container seedlings on favorable sites.

Seedlings grow remarkably well on dry and exposed

sites with shallow soils where few other shrubs

establish. Wildings and root cuttings are alternative

methods of propagation.

Uses and Management

Bitter cherry is frequently planted to provide browse

for wildlife and livestock. Elk, deer, and moose eat

twigs and buds of plants growing on ridges and open

slopes of winter ranges. The species is ranked as

highly valuable for elk in summer and fall (Kufeld

1973). Mule deer use trace amounts in winter and

make moderate use of the plant during other seasons

(Kufeld and others 1973). Its palatability has been

ranked as poor to fair for cattle and fair to fairly good

for sheep. Major livestock use occurs primarily in late

summer or fall when other vegetation is no longer

available. In many areas dense thickets of bitter

cherry grow out of reach of grazing animals, but

provide good cover. Lower growing forms may be

heavily browsed. Fruits are consumed by birds and

mammals; rodents harvest the seed.

Bitter cherry may be planted on shallow soils of

disturbed sites within its natural range to control

erosion on roadways and other disturbances (Monsen

and Davis 1985). Butterfield and Tueller (1980) and

Everett and others (1980) concluded bitter cherry

held considerable promise on acid mine spoils in

Nevada. Establishment from seed or seedlings can be

hindered by trampling and grazing; the seedlings

develop slowly. Rodents consume seed and damage

seedlings. Swihart and Yahner (1983) found rodents

preferred Prunus species over the remaining 29 common windbreak and conservation species tested. For

cottontails, Prunus fell into a neutral to preferred

category. Seedlings develop slowly and must be protected from competition with more aggressive plants

for up to 3 years following establishment.

Bitter cherry spreads by root sprouts; new shoots

develop readily following burning. Leege and Hickey

(1971), working near the Lochsa River of northern

Idaho, found spring and fall burns increased bitter

cherry sprouting about equally; mortality was slightly

higher in fall than in spring. Spring burning was

recommended for decreasing height of shrubs in seral

brush fields and improving browse conditions for elk.

Adequate regrowth occurred during the summer to

provide fall forage.

Varieties and Ecotypes

There are no releases.

Prunus fasciculata ______________

Desert Peachbrush

Desert peachbrush is a low growing, intricately

and divaricately branched deciduous shrub with a

rounded growth form (fig. 21). Shoots may attain

heights of 5 ft (1.5 m) and diameters of 8 ft (2.4 m).

Plants have a deep taproot as well as a spreading

fibrous root system. They are not rhizomatous

(Harrington 1964; Welsh and others 1987).
576 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Branches are grayish white, ending in pointed spines.

The light green leaves are 0.1 to 0.2 inch (2.5 to 5 mm)

long and 0.04 to 0.2 inch (1 to 5 mm) wide, cuneate to

spatulate, and entire to few toothed at the apex.

Flowers are small with five cream colored petals. Fruit

is a pubescent greenish-orange drupe with a rather

dry fleshy layer.

Ecological Relationships and Distribution

Desert peachbrush occurs in central and southern

Nevada, southwestern California, and western Arizona

on valley floors and foothills at elevations from 2,050 to

5,800 ft (625 to 1,770 m) (Ferguson 1983; Welsh and

others 1987). Desert peachbrush occurs in association

with chaparral, basin big sagebrush, Wyoming big

sagebrush, sand sage, blackbrush, creosotebush,

fourwing saltbush, quailbush, and lower pinyon-juniper communities. It can be found on well drained to

heavy soils ranging from basic to neutral. Greatest

area of occurrence is on sandy to rocky loam soils in

depressions along drainage areas, intermittent

streams, and creeks. Within these areas plants occur

as individuals or more typically in small groups, usually with a rather sparse understory. Following fire,

desert peachbrush sprouts readily and profusely from

the root crown. Seed dispersal and seedling establishment may be heavily dependent on rodents that

gather and cache seed.

Plant Culture

Harvestable seed crops are produced in about 1 of

every 5 years. Fruits are gathered by hand stripping or

by beating them into a container. The pulp attached to

Figure 21—Desert peachbrush provides soil

stabilization and cover for wildlife. Because of its

unique branching habit, it has potential as a low

maintenance ornamental.

the stony endocarp is thin and dry and does need not be

extracted. Leaves and sticks are separated by screening. Fruits are then dried and stored or seeded. There

are about 4,500 seed per lb (9,900 per kg). Viability is

generally good and is retained for 5 to 10 years of

storage following harvest (Stevens and others 1981a).

Seed retained in storage for 20 years exhibited fair

viability. Acceptable germination for seed purchases

is 90 percent and purity 70 percent (see chapter 24).

Seed can be drilled or broadcast and covered. It

must be placed 0.5 to 1 inch (1.25 to 2.5 cm) deep in a

firm seedbed. Fall or winter seeding provides the wet

prechilling period required to relieve dormancy. A

strong, fast growing seedling is produced and good

establishment is common if soil water conditions are

favorable. This species can also be propagated and

outplanted quite easily as bareroot or container nursery stock (Monsen and Davis 1985). Considerable

numbers of seedlings sometimes establish naturally

from rodent caches.

Uses and Management

Desert peachbrush is generally seeded in a mix with

other shrubs to improve diversity. California quail

and Gambel quail, chukar, nongame birds, and small

mammals use it extensively for cover, nesting, and

forage. With the exception of pronghorn (Smith and

Beale 1980), livestock and big game make little use of

the species. Although the shrubs are sometimes infested with tent caterpillars, they persist in spite of

defoliation.

Desert peachbrush establishes and does well when

seeded on disturbed sites. Individual plants can cover

large areas. Branches contact the soil surface over

most of the area and provide considerable ground

cover and soil protection. Perennial forbs and grasses

emerge and develop beneath and adjacent to the crown.

Desert peachbrush can establish on roadcuts and fills,

in sand and gravel pits, and on mined areas. It also has

ornamental potential, especially for low maintenance

landscaping projects. The intricate branching pattern, grayish-white stems, and light green leaves give

the plant a unique appearance.

Seeded plants should be given 2 to 3 years to establish and attain reasonable size prior to grazing. Once

established, this species is very hardy and can withstand considerable use.

Desert peachbrush has performed well when planted

on big sagebrush sites. It can be interseeded or

intertransplanted in crested wheatgrass or native

perennial herb seedings. Growth of established seedlings is not suppressed by annual grasses. Because of

its ability to persist, even with wildfires and invasions

of annual weeds, increased study and use of desert

peachbrush to improve diversity of plantings on desert

shrublands within its range are warranted.
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Chapter 22 Rosaceous Shrubs

Varieties and Ecotypes

There are no subspecies or released varieties of

desert peachbrush. Little selection work has been

attempted, but the species is being tested as a drought

tolerant rootstock for other Prunus species (Welsh and

others 1987).

Prunus spinosa and Prunus

tomentosa _____________________

Blackthorn and Nanking Cherry

A number of introduced Prunus species from eastern

Europe and Asia are adapted to mountain brush,

upper pinyon-juniper, and big sagebrush communities of the Intermountain region. Blackthorn and

Nanking cherry, both Asian species, have been successfully planted in the Intermountain West. Blackthorn grows to 13 ft (4 m) and Nanking cherry to 10 ft

(3 m) in height (Grisez 1974). Flowers of both species

are generally numerous and very showy. Blackthorn

produces a fleshy, dark purple, glabrous fruit; fruits

of Nanking cherry are semifleshy, light purple, and

pubescent.

Both species have been used in shelterbelt, windbreak, and conservation plantings. They have high

potential for use in and around recreational areas,

administration sites, low maintenance landscaping

projects, and on disturbed sites such as roadcuts and

fills or mine disturbances.

Both species transplant very well. However, they

are fairly difficult to establish from direct seeding.

Fall planting is required as the seed require wet

prechilling. Good diversity can be provided by including these species in mixtures with other shrubs. Grasses

and forbs grow well in association with both species,

but seedlings should be protected from herbaceous

competition. They have excellent resistance to insects

and grasshoppers, good cold tolerance, and are fairly

drought tolerant. Livestock and big game make some

use of twigs and leaves; fruits are taken by birds and

rodents. Birds use these species for cover, nesting, and

brood rearing.

Seed and bareroot or container stock of both species

may be obtained from commercial nurseries in the

Midwest and Intermountain regions.

Prunus virginiana _______________

Chokecherry

Chokecherry is a thicket-forming shrub or small

tree with several deep roots and a well-developed,

fibrous root system (fig. 22). Plants develop rhizomes

that are 0.4 to 0.6 inch (1 to 1.5 cm) in diameter and

extend laterally about 3 ft (1 m) from the stems. New

stems sprout from the rhizomes and the root crown.

Leaves are alternate and elliptic, about 0.1 to 4 inches

(2.5 to 10 cm) long and finely serrate along the edges

(Welsh and others 1987). Purplish-red glands are

usually found near the tip of the petiole or along the

leaf base. The leaves are bright green above and paler

beneath. They turn bright red in early autumn.

Elongate racemes of fragrant white flowers develop

from April to June. Racemes are about 3 to 6 inches

(8 to 15 cm) long. The insect-pollinated flowers are

perfect; five petals spread from a gland-lined hypanthium. Only one of the two ovules normally develops. The fruits are red to purplish-black cherrylike

drupes about 0.4 inch (1 cm) in diameter; each contains a single seed with a stony endocarp (Grisez 1974;

Welsh and others 1987). Seeds ripen from July to

September. Fruit production is greater during years

of average or better rainfall and on plants growing in

full sunlight.

Hitchcock and others (1961) recognized three varieties of chokecherry. Common chokecherry is a large

shrub or small tree growing to 50 ft (15 m) in height

with bright to deep red fruits. Black chokecherry is a

small to medium sized shrub or small tree 13 (20) ft (4

[6] m) tall with dark, nearly black fruits and thick

leaves. Western chokecherry ranges from 7 to 13

(20) ft (2 to 4 [6] m) tall with dark, nearly black fruits

and thick leaves.

Ecological Relationships and Distribution

Chokecherry occurs primarily in central and eastern United States. It ranges from Saskatchewan to

Newfoundland and south to Kansas and North Carolina (Hitchcock and others 1961; USDA Forest Service

1937). Black chokecherry is distributed from eastern

British Columbia to the Dakotas and southward

east of the Cascades from northern California to

New Mexico. Western chokecherry occurs primarily

Figure 22—Chokecherry provides palatable and

nutritious browse that is used by many wildlife

species.
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Chapter 22 Rosaceous Shrubs

along the west coast from British Columbia south to

California (Hitchcock and others 1961; Welsh and

others 1987).

Chokecherry is distributed from low elevation foothills into mountainous areas at elevations from sea

level to 9,000 ft (2,750 m) on sites with annual precipitation of 12 to 30 inches (305 to 760 mm) (Thornburg

1982; USDA Forest Service 1937). It is often associated with sagebrush, mountain brush, pinyon-juniper, and dry conifer types. It grows on moderately

acidic to basic or saline soils with silty to sandy

textures that are moderately deep, fertile, and usually

well drained. Scattered plants or small clumps may be

found on talus slopes and other rather dry sites; larger

thickets develop along streams and around springs,

ditches, or roadsides in semi-arid areas. In higher

elevation communities, common chokecherry tends to

occur on more open sites and along riparian areas. It

is not adapted to high water tables or prolonged

flooding (Wasser 1982).

Plant Culture

Chokecherry spreads by seed. It also spreads vegetatively from buds produced on rhizomes and the root

crown. However, it is not commonly propagated by

vegetative means. Leafy softwood cuttings can be rooted,

but hardwood cuttings are not recommended (Hartmann

and others 1990; Marchant and Sherlock 1984).

The drupes fall soon after reaching maturity (Grisez

1974). Birds and mammals harvest and spread the

seed. Germinability may be increased after seeds

have passed through their digestive tracts (Grisez

1974). Fruits are collected either by hand picking or

beating them into a canvas or into a container. Seed

harvested too early may have low germinability. If

harvested too late, many seeds may already have

fallen and others will split while being cleaned, leaving the embryo vulnerable to mechanical damage

(Grisez 1974; Marchant and Sherlock 1984). Harvested fruits should not be exposed to heat as mold

may develop, and viability may be reduced. Fruits are

cleaned by macerating the fleshy material with a

Dybvig cleaner and flushing with water to remove the

flesh from the seed. This is followed by drying and

further cleaning with a clipper cleaner. Fermentation

may facilitate cleaning, but may adversely affect

germination (Grisez 1974). There are about 4,100

cleaned seed per lb (9,020 per kg) (see chapter 24).

For seed purchases, acceptable purity is 90 percent

and germination 70 percent.

Seed may be sown immediately after cleaning

without further drying. Short-term storage for a few

months requires only air drying. Optimal long-term

storage conditions have not been determined, but

warm, humid conditions result in rapid deterioration

of seed quality. On the other hand, excessive drying

can also be harmful. Belcher (1985) and Marchant and

Sherlock (1984) reported that air dried seed stored in

sealed containers at 37 to 41 ∞F (3 to 5 ∞C) will remain

viable for 3 to 5 years.

Chokecherry seeds generally have fairly high viability, but germination of untreated seed is generally

quite low. About one-half of the viable seed will germinate in 69 days (Wasser 1982). Western chokecherry

generally germinates more readily than black

chokecherry (Monsen and Davis 1985). Wet prechilling

can increase both the rate and percent of germination.

Seed is wet prechilled at 36 to 41 ∞F (20 to 23 ∞C) for 120

to 180 days (Wasser 1982). Thoroughly soaking the

seed prior to wet prechilling may improve effectiveness of the treatment. Wet prechilling is adequate if

the stones have cracked, but the radicles have not yet

emerged. The rate of radicle emergence is normally

rather low, but can be further reduced, if necessary, by

lowering the prechilling temperature once the stones

have broken open (Grisez 1974; Wasser 1982). Fracturing the endocarp, boiling, freezing, and various

chemical treatments have provided inconsistent results and are sometimes detrimental (Grisez 1974).

The Association of Official Seed Analysts recommended

a 60 to 90 day wet prechill period at 37 to 41 ∞F (3 to

5 ∞C) for testing germination (Belcher 1985). They

also provided tetrazolium testing and radiographic

procedures.

Nonprechilled seed must be fall seeded, but artificially prechilled seed may be planted in spring. Seed

should be planted 0.5 to 1 inch (1.25 to 2.5 cm) deep in

a firm seedbed. Seed can be drill seeded or metered

through a seed dribbler mounted on a crawler tractor.

The former method has provided excellent results

(Stevens 1992). This species can be seeded individually or as a component of a mixture of other shrubs and

nonaggressive forbs and grasses. Seedlings establish

most successfully when planted on sites that receive

at least 15 inches (38 cm) of annual precipitation

(Monsen and Davis 1985). They have often been planted

on drier sites with poor success. Seedlings of black

and western chokecherry are not highly competitive,

but become more so after 2 to 3 years (Monsen and

Davis 1985; Plummer and others 1968; Thornburg

1982). Once the root system becomes well established,

plants can exist with other woody species or herbaceous vegetation and can withstand considerable

grazing.

Bareroot stock of all chokecherry species may be

grown in the nursery. However, seed and new seedlings are often gathered by birds and small rodents.

One growing season usually provides stock of adequate size for transplanting (Shaw 1984). Although

initial growth of germinants is very rapid, the rate

sometimes drops off considerably during the growing season; a 2 year cropping period is often necessary
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 579

Chapter 22 Rosaceous Shrubs

(Marchant and Sherlock 1984). Bareroot seedlings

have a strong taproot system. Rhizome and new shoot

development are initiated early in the season, often

prior to lifting. New growth can easily be damaged

during lifting and planting, thus reducing the success

of establishment (Monsen n.d.). Seed, germinants, or

small transplants can be used to propagate container

stock. Seedling development requires a 3 to 5 month

cropping period (Landis and Simonich 1984).

Uses and Management

Chokecherry has been extensively used to improve

wildlife habitat. Plants typically form relatively open

thickets that allow livestock and big game access to

abundant amounts of nutritious and palatable browse.

Despite its deciduous nature, chokecherry maintains

relatively high nutrient levels in late fall and winter

(Dietz 1972). Chokecherry provides fairly palatable

browse for big game and is widely utilized by elk,

moose, and bighorn sheep (Plummer and others 1968);

it is also the preferred deer browse on many winter

ranges throughout the Intermountain west (Dietz

1972; Kufeld 1972; Kufeld and others 1973).

Chokecherry receives its greatest use in spring and

fall, but elk and deer use it to some extent year around.

It is moderately palatable to all classes of livestock,

although it seems to be more heavily browsed by sheep

than cattle (USDA Forest Service 1937; Wasser 1982).

Fruits are taken by birds, bear, and many small

mammals (Aune and Stivers 1985; Blauer and others

1975; Gullion 1964; Noste and Bushey 1987; Wasser

1982). Plants are productive and withstand moderate

use. However, repeated heavy browsing results in

highlining, hedging, and lower availability of forage.

New sprouts may be killed by trampling or browsing.

Grazing does not affect the next year’s fruit production; flowers are borne on twigs of the season.

Chokecherry may be toxic if consumed in large

amounts; leaves contain the glucoside amygdalin

(Noste and Bushey 1987). Young shoots contain the

greatest concentrations. Most problems occur in spring

and early summer. Following fall frosts, the plants

become essentially harmless. Livestock are notlikely

to consume fatal amounts of chokecherry except in

areas of animal concentration.

Chokecherry is extremely valuable for watershed

and streambank stabilization, restoration of mine and

roadway disturbances, landscape plantings in recreation areas, and increasing diversity in conservation

and windbreak plantings (Ferguson 1983; Plummer

and others 1968; Shaw and Cooper 1973; Wasser

1982).

All chokecherries are edible, but have an extremely

sharp and biting flavor if eaten before fully ripe. The

fruits make excellent wine, jellies, syrup, honey, flavorings, and perfume oil. Native Americans used the

dried fruits to make pemmican cakes for winter use

(Welsh and others 1987). Tea was prepared from the

stems and bark (Craighead and others 1963).

All species of chokecherry are well adapted to fire,

particularly if burned when dormant or when soil

moisture is high (Anderson and Bailey 1980; Tisdale

and Hironaka 1981; Young 1983). Although top growth

is easily killed, vigorous sprouting from rhizomes or

the root crown provides for rapid recovery following

the initial decline (Leege and Hickey 1971; Noste and

Bushey 1987). After the first postburn season, numbers of stems and coverage by the species can exceed

preburn levels for a number of years. Although establishment from seed is quite rare on undisturbed sites,

the species may be dependent on seedbanks for postfire

regeneration, at least in riparian areas or higher

elevation communities (Kramer 1984; Wright and

others 1979; Young 1983).

Varieties and Ecotypes

There are no releases.

Purshia glandulosa ______________

Desert Bitterbrush

Desert bitterbrush is an evergreen shrub resembling Stansbury cliffrose and frequently mistaken for

it. Welch and others (1987) described desert bitterbrush as ranging from 5 to 9 ft (1.5 to 2.7 m) in height

(fig. 23), generally upright, but sometimes low and

Figure 23—Desert bitterbrush is a stable hybrid

derivative of Stansbury cliffrose and antelope

bitterbrush.
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Chapter 22 Rosaceous Shrubs

spreading. Branchlets are prominently glandular. The

glandular leaves are 0.1 to 0.4 inch (3 to 10 mm) long

and up to 0.2 inch (5 mm) wide, cuneate, glabrous

above, and slightly tomentose beneath. Petals are

0.2 to 0.3 inch (5 to 8 mm) long and creamy white to

yellowish. Achenes are oblique and 0.8 inch (2 cm)

long, including the elongate style. Glandular evergreen leaves and smaller, elongate achenes differentiate desert bitterbrush from antelope bitterbrush

(fig. 24).

Ecological Relationships and Distribution

Desert bitterbrush is restricted to southern

California, southern Nevada, southern Utah, and

northern Arizona (Koehler and Smith 1981; McArthur

and others 1983b; Nord 1965; Stutz and Thomas

1964). Koehler and Smith (1981) and Stutz and

Thomas (1964) considered it to be a hybrid derivative

of Stansbury cliffrose and antelope bitterbrush parentage that arose from hybridization and subsequent

introgression between these two species (McArthur

and others 1983b). This hybridization is a relatively

recent development and populations remain genetically distinct. Welch and others (1987) transferred

cliffrose, Cowania, to the genus Purshia because hybrids of cliffrose, antelope bitterbrush, and desert

bitterbrush occur where the distribution of these species overlap.

In southern Utah, desert bitterbrush grows

principally in the blackbrush type, and in pinyonjuniper extensions into this type, at elevations from

3,460 to 4,470 ft (1,055 to 1,363 m) (Welsh 1987). It is

more common in southern Nevada and California

than in Utah. It often occupies sites where cool and

warm desert species interface and is often the dominant or codominant shrub when present.

Figure 24—Desert bitterbrush, (left) is

distinguished from antelope bitterbrush

(right) by its glandular, evergreen leaves.

Plant Culture

Seed production, harvesting, conditioning, and

storage are similar for desert and antelope bitterbrush (Deitschman and others 1974c; Giunta and

others 1978). When ecotypes of both species are grown

at common garden sites in Idaho and Utah, desert

bitterbrush is normally smaller in stature than the

erect antelope bitterbrush growth form. It produces

less seed than antelope bitterbrush and good seed

crops occur less frequently. There are approximately

21,000 seed per lb (46,200 per kg). Seed retain viability

for up to 25 years when stored under warehouse

conditions (Stevens and Jorgensen 1994). For seed

purchases recommended purity is 95 percent and

germination 90 percent (see chapter 24).

A high percentage of desert bitterbrush achenes

fail to mature normally. Commercially harvestable

seed crops are produced once in every 2 to 6 years at

native collection sites. Consequently, it is necessary

to harvest adequate quantities of achenes in good

years to fill anticipated needs (Plummer and others

1968). Desert bitterbrush seed orchards have been

established at various locations in Utah and Idaho in

an attempt to develop more consistent seed production centers.

Germination requirements of desert and antelope

bitterbrush are similar. Both require short periods of

wet prechilling to release dormancy. Desert bitterbrush seedlings are more difficult to establish than

those of antelope bitterbrush. They grow more slowly

and attain maturity later when grown together in

common gardens. Seedlings also grow more slowly

under nursery conditions with regular irrigation.

Because of its slow growth, desert bitterbrush is

less competitive with annual weeds than antelope

bitterbrush.

Direct comparison of establishment attributes between the two species of bitterbrush may create a

somewhat distorted picture. It is adapted to more

xeric sites and when planted in such areas can perform well. It usually grows in areas receiving less

than 14 inches (36 cm) of annual precipitation.

Artificial plantings on these sites are often unsuccessful; the precipitation received may be too low or

erratic to permit establishment. However, desert bitterbrush seedlings are relatively hardy and can survive adverse weather conditions if they once establish.

Rodents quickly gather planted desert bitterbrush

seed. Consequently, planting in late fall or winter is

advised to reduce seed predation. Seeding directly

into a weedy understory should be avoided. Seedlings normally require 1 to 3 years to fully establish.

Both bareroot and container grown seedlings establish well and can be used to provide more rapid

establishment.
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Chapter 22 Rosaceous Shrubs

Uses and Management

To date, most plantings of desert bitterbrush have

been made on upland pinyon-juniper sites where

antelope bitterbrush is native (Davis 1983a). In general, antelope bitterbrush ecotypes respond better

than desert bitterbrush at most of these planting

sites. However, desert bitterbrush plants have established and persisted on antelope bitterbrush sites in

central and northern Utah, southern Idaho, and southcentral Idaho. Desert bitterbrush plantings have also

been successfully established on pinyon-juniper/

blackbrush sites in southern Utah (Davis 1983a).

Desert bitterbrush exhibits important vegetative traits

such as production of semi-evergreen leaves, ability to

resprout following fire, and good drought tolerance.

Desert bitterbrush demonstrates adaptability to disturbances and can be used to seed infertile mine spoils

and roadways in adapted areas. Welch and others

(1983a) reported that it reduces soil erosion and provides habitat for game and non-game animals. Roots

may form symbiotic nitrogen-fixing relationships

with the endophyte Frankia (Nelson 1983). Once established, desert bitterbrush grows well with herbaceous understory plants and can be managed to provide seasonal herbage, particularly winter forage for

big game and livestock. Welch and others (1983a)

found desert bitterbrush plants in a southwestern

Idaho common garden retained a significantly greater

quantity of leaves in winter than collections of antelope bitterbrush and cliffrose. Crude protein content of

the desert bitterbrush accessions exceeded that of the

other two species. Palatability may be somewhat lower

than that of antelope bitterbrush. However, where

desert bitterbrush occurs, variety, quality, and quantity of other browse species are often low. As a result,

considerable use is made of the species, particularly by

deer and cattle. Its growth habit provides much needed

shade and cover for small game and birds. Seeds are

collected, eaten, and cached by rodents, birds, and

reptiles. Insect populations associated with desert

bitterbrush are often high, especially during flowering and seed set.

Varieties and Ecotypes

No selections of desert bitterbrush have been developed for release (Davis 1983a). Ecotypes vary in growth

form from decumbent to upright. Populations or

ecotypes having good drought tolerance (Davis 1983a),

fire tolerance, winter leafiness (Welch and others

1983a), and nitrogen-fixation capabilities (Righetti

and others 1983) have been identified. Because desert

bitterbrush, antelope bitterbrush, and Stansbury

cliffrose hybridize freely, it may be possible to combine desired attributes of these three species.

Purshia tridentata _______________

Antelope Bitterbrush

Antelope bitterbrush is an intricately branched,

deciduous shrub varying in growth habit from low

decumbent spreading forms to upright arborescent

plants over 13 ft (4 m) in height (fig. 25) (Blauer and

others 1975; Welsh and others 1987). Leaves are

alternate, simple, pinnatified or apically three-toothed

and usually glandular. Flowers are borne on short

spurs of previous year’s growth. They are numerous,

white to yellow, and about 0.3 inch (8 mm) in diameter (Harrington 1964; Munz and Keck 1959; Welsh

and others 1987). The fruit is a cartilaginous achene

0.2 to 0.5 inch (6 to 12 mm) long (Young and Evans

1983) with a persistent tapering style and a black

pyriform seed (Blauer and others 1975). Plants are

reported to be long lived. Nord (1965) found the average age of decumbent plants growing above 8,000 ft

(2,440 m) in northern California was 52 years; at

Figure 25—(A) Mature antelope bitterbrush plant

and (B) fruit.

A

B
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Chapter 22 Rosaceous Shrubs

other locations the average age exceeded 150 years

(Ferguson 1983a; Nord 1959).

Populations of antelope bitterbrush are extremely

variable. Giunta and others (1978) described ecotypes

with differing growth forms, fire tolerance, rates and

periods of growth, seed production, drought resistance, nutritional qualities, heat tolerance, cold hardiness, climatic adaptation, resistance to disease or

insect damage, palatability to game or livestock, evergreen habit, vegetative reproduction, and shade tolerance. Winward and Findley (1983) recognized similar

variations in Oregon and California populations, but

also noted marked differences in leaf size, shape, color,

presence or absence of glands, flowering, and seed set.

Many differences observed among populations are

genetically influenced (Klemmedson and Ferguson

1969; Nord 1959a; Plummer and others 1968; Wagle

and Vlamis 1961). Ecotypes maintain their native

growth habits and morphological characteristic when

planted over a wide range of sites. However, climatic

and edaphic conditions alter growth response to some

extent (Davis 1983a; Edgerton and others 1983; Nord

1965; Shaw and Monsen 1983a).

Ecological Relationships and Distribution

Cowania is derived from the neotropical Tertiary

flora that emerged in the Sierra Madre region of

Mexico during the Tertiary epoch (Axelrod 1958).

McArthur and others (1983b) postulated that Purshia

is an early derivative of Cowania that evolved in isolation. Chromosome number for both genera is n = 9

(Alderfer 1977; McArthur and others 1983b; Sanderson

1969). Present contacts between Purshiaand Cowania

are considered very recent; their partially separated

flowering periods form a weak isolation barrier. Natural hybrids are found throughout most of Utah and

Nevada where Cowania and Purshia overlap (Stutz

and Thomas 1964). The hybrids backcross with the

parental stock, producing genetic introgression in

both genera. Introgression is considered to be the

major process contributing to genetic differentiation

of ecotypes (McArthur and others 1983b; Stebbins

1959; Stutz and Thomas 1964; Thomas 1957). Desert

bitterbrush is a southern adaptive derivative resulting from this process. In addition, cliffrose characteristics are present in most populations of antelope

bitterbrush that occur beyond the northern margins of

cliffrose distribution (Stutz and Thomas 1964).

Antelope bitterbrush occurs from the Cascade Mountains eastward through the Rocky Mountains and from

British Columbia south to Arizona and New Mexico

(Ferguson 1983; Hitchcock and others 1961; Koehler

and Smith 1981; McArthur and others 1983b; Stutz

and Thomas 1964). It grows in big sagebrush, pinyonjuniper, and mountain brush communities and

ponderosa pine forests. At lower elevations, in the

southern portion of its range, it extends into blackbrush

communities. It may also be encountered on high

mountain exposures intermixed with aspen, limber

pine, and Douglas-fir. It usually exists at elevations

ranging from 200 ft (60 m) in the Pacific Northwest to

11,500 ft (3,510 m) in the Sierra Nevada Mountains in

areas receiving 12 to 25 inches (30 to 64 cm) of annual

precipitation (Nord 1965; Stanton 1959; Tew 1983).

Antelope bitterbrush is often a component of mixed

shrub communities (Blaisdell 1953; Ferguson and

Medin 1983; Holmgren and Basile 1956), and is

frequently a dominant in the northwestern part of its

range (Franklin and Dyrness 1973; Mueggler and

Stewart 1980; Nord 1965). Prostrate and semi-erect

growth forms of antelope bitterbrush occur most

commonly as understory plants in forested communities or at high elevations (Nord 1965). Some lowgrowing types are also encountered at low elevations

and under dry conditions, but the upright, tree-like

forms usually dominate on more xeric sites (Nord

1965).

Some ecotypes are adapted to specific soil conditions. Prostrate growth forms occur on shallow, coarse

textured soils in central Idaho, while tall, erect forms

grow on deep productive soils where basin big

sagebrush may also occur (Tew 1983a). Plummer and

others (1968) reported that ecotypes from neutral or

slightly acidic soils in Idaho are not adapted to calcareous soils of central Utah. Davis (1983a) found that

only 24 of 259 accessions planted on pinyon-juniper

sites in Utah scored performance index values that

would recommend their use in these areas.

Antelope bitterbrush ecotypes vary in their response

to burning. Adams (1980) suggested that antelope

bitterbrush evolved as a fire climax. Edgerton (1983)

reported rapid establishment of new seedlings following fire in ponderosa pine/antelope bitterbrush

communities in the southern Cascade Mountains.

Reestablishment from seed occurs erratically in response to climatic conditions and may be limited by

grazing, rodent activity, presence of annual weeds,

and other factors. Emergence from rodent caches is

common in many areas.

Resprouting capabilities vary with ecotype. Some

ecotypes in forested communities recover well by

resprouting after burning (Martin 1983; Rice 1983;

Sherman and Chilcote 1972; Zlatnik 1999). Driver and

others (1980) found 100 percent resprouting of plants

burned in a ponderosa pine/grass/antelope bitterbrush

habitat type. Antelope bitterbrush plants recovered

well from spring burning in ponderosa pine-antelope

bitterbrush-pinegrass habitats, but little resprouting

followed fall burning when soil moisture was low

(Driver 1983).
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Chapter 22 Rosaceous Shrubs

Billings (1952), working in the western Great Basin,

and Hormay (1943) in California found that antelope

bitterbrush rarely resprouted following fire and suggested burning would permanently eradicate the species. Blaisdell (1953) found that not all plants growing

on the Upper Snake River Plain in Idaho resprouted

even after a light burn. Murray (1983) reexamined

sites studied by Blaisdell and reported no antelope

bitterbrush recovery 43 years after burning. Bunting

(1985) and Bunting and others (1987) concluded that

the columnar ecotypes growing in central Idaho do

not recover from burning. At Wallsburg, UT, 80 to

90 percent of the antelope bitterbrush plants burned

in a wildfire resprouted the first year following the

fire. By the end of the third year mortality exceeded

90 percent. This pattern is commonly observed following wildfires in antelope bitterbrush stands in Idaho

and Utah.

Plant Culture

Antelope bitterbrush was perhaps the first native

shrub used for wildlife habitat and rangeland restoration in the Intermountain region. Since the early

1940s antelope bitterbrush has been seeded in many

areas in the western United States. Holmgren and

Basile (1956) reported that of approximately 50 species tested in Idaho, antelope bitterbrush was the

easiest to establish from direct seeding.

Nord (1965) reported that young plants growing

on native sites usually begin producing seed in about

10 years. By contrast, plants grown in cultivated

nurseries, or in native stands managed for seed production, often begin producing seed when 2 to 4 years

of age. Latitude, elevation, season, and site conditions

influence vegetative phenology and the date of seed

maturation (Nord 1965).

Leaf growth of antelope bitterbrush precedes the

appearance of floral buds. At a common garden near

Boise, ID, leaf growth of ecotypes from a wide geographic range was initiated between March 17 and 23

(Shaw and Monsen 1983b). Leaves reached mature

size by about mid April when flower buds were beginning to develop. Leader growth of the different

ecotypes was initiated during the first week of May,

coinciding with anthesis. Most stem elongation occurred

in June and early July. Stem growth rate diminished

in July. Little elongation occurred in late summer or

fall. All five ecotypes examined responded similarly.

Climatic conditions favorable to good seed production antedate the crop by at least 1 year; flower buds

develop primarily on second year wood (Nord 1965).

Plants that receive favorable moisture produce good

leader growth and increased root reserves that support seed production the following year (McCarty and

Price 1942) (fig. 25b).

Shaw and Monsen (1983b) reported that accessions

of antelope bitterbrush growing in a common garden

in southern Idaho set and matured seed in 42 to 48

days in 1979 and 57 to 60 days in 1980. Blaisdell (1958)

found the period of flowering and seed maturation was

slightly longer (66 days) at sites near Dubois, ID. Seed

maturation dates can be predicted with reasonable

accuracy. Nord (1965) developed an equation using

elevation and latitude to predict mean dates of seed

maturation for stands in California. Actual harvesting

dates generally fell within 6 to 10 days of predicted

dates.

Accurate determination of seed ripening dates is

critical for commercial seed harvesters as seeds shatter quickly when mature. Achene development

progresses through a number of recognizable stages.

In southern Idaho developing achenes reach nearly

mature size by mid June (Shaw and Monsen 1983b).

During this period of rapid growth, the endosperm

remains milky. About 10 to 16 days prior to maturation, the endosperm changes from white to dark red

(blood stage). Achenes dry rapidly during the last 3 to

5 days of this period.

Achenes can wither and abort at any stage of development. Late frosts, poor or inadequate soil water, and

insect damage are principal reasons for crop failure.

Many failures occur during the “blood stage” just prior

to achene maturation. Insects may destroy a large

proportion of the seed crop (Basile and Ferguson 1964;

Basile and others 1964; Ferguson 1967; Ferguson and

others 1963; Furniss 1983; Furniss and Barr 1975).

Shaw and Monsen (1983b) reported 50 to 60 percent of

all flowers that initially appeared failed to develop a

mature fruit in a southern Idaho seed orchard as a

result of insect damage.

Prospective seed collection sites must be inspected

frequently, particularly when seeds are in the blood

stage to determine when and where harvestable crops

will be available. Harvesting is accomplished by striking branches with a paddle to dislodge seeds into

canvas or metal trays (Giunta and others 1978; Nord

1967). Most native sites produce 150 to 200 lb of seed

per acre (168 to 224 kg per ha) although production

may approach 500 lb per acre (560 kg per ha) (Nord

1965).

Few seed nurseries have been established on agricultural land. Cultivated plantings should be established on or near native sites. Plantings should be

located in areas protected from high wind; flowers

are cross pollinated by insects and seed shatter

easily (Blauer and others 1975). In southern Idaho 6 to

10 year old plants of an upright ecotype planted at 6 ft

(1.8 m) spacings produced more seed per acre than

plants planted at 6 by 12 or 12 by 12 ft (1.8 by 3.7 or

3.7 by 3.7 m) spacings. However, as plants attained

mature stature, those at 6 ft (1.8 m) spacings became

so overgrown that seed harvesting became impractical. Depending on growth habit, shrubs should be
584 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

spaced 12 to 15 ft (3.7 to 4.6 m) apart to permit good

seed production and facilitate site maintenance and

seed harvesting.

Maintaining a stand of native herbs as an understory is a practical approach for controlling weeds and

supporting predatory insects. Field cultivated plants

appear to benefit from cultural treatments, but response to fertilizer, water, and pesticides is not known.

Native stands of antelope bitterbrush can also be

managed for seed production. Plants respond favorably to pruning, thinning, weed control, and protection from grazing.

Annual collections of antelope bitterbrush from western rangelands have sometimes exceeded 40,000 lb

(18,182 kg). Considerable seed is harvested annually

from most Intermountain and Northwestern States.

Demand varies considerably from year to year. Areas

in south-central Idaho and northern California tend

to produce more consistent seed crops than most sites

within the Intermountain region. More seed is usually

harvested from areas in central Idaho and the Pacific

Northwest than is used in those regions. However,

seed collected from central Utah and eastern Nevada

is generally not sufficient to meet regional demands.

Likewise, the quantity of seed harvested from arid

regions and decumbent growth forms growing in

ponderosa pine and aspen communities is generally

inadequate.

Seed are easily hand harvested. Effective mechanical harvesters have not been developed (Nord and

others 1967). Air-dried collections are cleaned by

first screening to remove sticks and debris. The

husks are then detached with a barley debearder,

Dybvig, or other scarifier. Remaining material is again

screened to separate seed from the detached husks

(Giunta and others 1978). At 100 percent purity there

are approximately 15,000 seed per lb (33,000 per kg).

Seed are reasonably hard and are not easily broken,

fractured, or damaged when cleaned or planted. Recommended purity for seed purchases is 95 percent and

germination 90 percent (see chapter 24).

Cleaned seed can be stored under warehouse conditions for extended periods without loss of viability.

Afterripening improves germination. Stevens and others (1981a) reported a significant increase (79 to 86

percent) in germination between the second and third

year of storage. Seed germination remained unchanged

after 20 years of warehouse storage, but decreased to

24 percent after 25 years. Antelope bitterbrush seed

can usually be stored without suffering extensive

insect damage, but storage areas must be kept rodent

free.

Antelope bitterbrush seed are large, smooth, and

fairly heavy. Consequently, they are easily carried

through most seeders. When planted in mixtures, antelope bitterbrush can be used to carry other light-weight

seed through seeding mechanisms. Seeds require a 2

week wet prechilling to release dormancy, thus they

should be fall or winter seeded.

Recommended seeding rates for antelope bitterbrush vary considerably. The shrub may be drill seeded,

interseeded in rows at various interspacings, or hand

planted in clearings or selected spots. Hubbard (1964)

recommended drilling at a rate of 3 lb per acre (1.4 kg

per ha). Extensive pinyon-juniper chainings have been

interseeded using a Hansen seed dribbler. Rows are

usually spaced 8 to 12 ft (2.4 to 3.7 m) apart when

mechanical interseeders are used (Stevens and others

1981b). Between 5 and 20 seeds are usually dispensed

per linear ft (16 to 66 per m) in each interseeded row.

At a rate of 20 seeds per ft (66 per m), approximately

87,000 seeds are planted per acre (214,974 per ha), if

rows are spaced 10 ft (3 m) apart. This equates to 3 to

5 lb per acre (3.4 to 5.6 kg per ha) depending on the seed

lot planted.

Hand seeding has been used extensively to plant

steep or inaccessible slopes. Herbaceous vegetation

must be cleared to enhance shrub seedling survival.

Clearings amid annual or perennial vegetation

should be between 2.5 to 3.3 ft (0.75 to 1 m) square

to effectively reduce competition. Hand seeding antelope bitterbrush and other shrub seed into depressions created by uprooting pinyon-juniper trees following anchor chaining is also a successful planting

technique.

Rodents actively gather planted seed (Evans and

others 1983; Young and Evans 1978a,b), and forage on

small seedlings. Rodent damage can be extremely

serious and limit planting success. This problem can

be reduced by planting late in fall after rodents hibernate. Coating seeds with a combination of Arasan and

Endrin eliminates nearly all rodent predation (Medin

and Ferguson 1980), but the toxic effects of Endrin on

other animals have eliminated its use. Everett and

Stevens (1981) found thiourea treatments reduced

deer mouse predation. However, the compound is

now classified as a carcinogen. No other chemical has

been developed that effectively diminishes rodent

predation.

Evans and others (1983) and Reichman and Oberstein

(1977) reported that size and depth of seed caches

influenced detection by rodents. Rodents locate seed

primarily by olfactory cues (Howard and Cole 1967;

Johnson and Jorgensen 1981), thus edaphic conditions can affect seed discovery (Johnson and Jorgensen

1981). Rodents were better able to locate antelope

bitterbrush seeds when two or more seeds were planted

together. Rodents failed to locate individual seeds

placed 0.4 inch (1 cm) deep, but removed 75 percent of

the seeds if two seeds were placed together at this

depth. Predation increased to 98 percent when more

than 10 seeds were planted together. Evans and others
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Chapter 22 Rosaceous Shrubs

(1983) found that planting depth affected seed detection. Rodents consumed all seeds placed on the soil

surface or planted at a depth of 0.4 inch (1 cm), but

only about 50 percent of seeds planted between 0.8

and 1.6 inch (2 and 4 cm) deep.

Available cover also strongly influences seed predation. Evans and others (1983) reported significantly

less rodent predation of seed caches in burned areas

lacking plant cover than in unburned sites with considerable vegetative cover. Similar responses have

been noted in seedings on mine and roadway disturbances. Rodents apparently cache antelope bitterbrush seeds in small openings, but are not likely to

venture onto exposed sites for extended foraging.

Planting success of range and wildlife seedings in

different plant communities also indicates considerable differences in rodent foraging habits (Basile and

Holmgren 1957; Casebeer 1954; Holmgren and Basile

1959; Nord 1965). Success has generally been better

in pinyon-juniper communities cleared of trees by

anchor chaining than on burned antelope bitterbrush/

bunchgrass ranges infested with cheatgrass.

Booth (1980) and Evans and others (1983) found

pretreating antelope bitterbrush seeds with a fungicide reduced damping-off problems and ultimately

increased the number of successfully established seedlings. Cutworms and wireworms can also destroy

newly developing seedlings of antelope bitterbrush

(Hubbard 1956), but outbreaks are usually limited to

isolated situations.

Compared to most shrub and many herb species,

antelope bitterbrush seedlings establish quite successfully even under harsh circumstances. Consequently, the shrub can be used to revegetate roadways, mine disturbances, and other infertile sites. If

protected from browsing, young plants grow rapidly

and provide considerable forage and ground cover

within 3 to 5 years.

Antelope bitterbrush is often interseeded with a

combination of grasses and broadleaf herbs. However,

Monsen and Shaw (1983c) reported that intermediate

wheatgrass and crested wheatgrass planted as an

understory can significantly reduce or minimize natural seedling recruitment of this shrub. It is essential

that the shrub be seeded with less competitive herbs.

Interseeding antelope bitterbrush onto sites dominated by cheatgrass is not recommended unless remedial treatments are employed to reduce or replace the

cheatgrass understory.

Bareroot or container seedlings are frequently used

to circumvent seed predation by rodents and speed

establishment. Dormant or carefully hardened seedlings should be planted before native antelope bitterbrush shrubs at or near the planting site break

dormancy (Carpenter 1983). Establishment success is

usually quite high. Antelope bitterbrush seedlings

and young plants are often browsed heavily. Project

seedings or plantings should be large enough to dissipate animal browsing or establishment may be adversely affected.

Uses and Management

The broad genetic base of antelope bitterbrush and

its wide geographical distribution (Stutz and Thomas

1964; Thomas 1957), provide considerable opportunity to promote the use of natural and artificial populations and hybrids with specific traits. Few other

shrub species are so diverse.

Antelope bitterbrush is widely regarded as an important browse plant for big game and domestic livestock. It has been planted extensively in big game

habitat improvement projects throughout the Intermountain and Pacific Northwest regions to provide a

forage base in areas where game and livestock use

may be seasonally heavy. Numerous seeding and

transplanting projects have been conducted to improve sites where shrub populations have been depleted or eliminated. Antelope bitterbrush grows

well with a wide number of native understory species

and other shrubs and is often planted in mixtures to

reestablish plant diversity. It forms important associations with various conifers, mountain brush species, and bunchgrasses. However, in certain areas

this plant exists with few other species and may be

the dominant plant encountered. In either situation,

reestablishing the shrub improves wildlife habitat,

watershed resources, and community and structural

diversity.

A wide array of ecotypes exists within the broad

geographic range of antelope bitterbrush. Numerous

ecotypes have been propagated and planted at various

study sites in the western United States. Some collections exhibit adaptation to a rather broad range of site

conditions; others require rather specific edaphic and

climatic conditions. An upright growth form from

Sanpete County, UT, for example, performed much

better than any other source when seeded on a broad

array of Utah sites (Davis 1983). A natural hybrid

of antelope bitterbrush and Stansbury cliffrose displayed better survival, persistence, and herbage yield

than most other antelope bitterbrush ecotypes when

seeded on a number of Utah sites.

Planting failures sometimes result when antelope

bitterbrush ecotypes are planted on similar parent

materials but at elevations about 2,000 ft (610 m)

higher than the native stands. In addition, ecotypes

have failed to persist if sources from strongly basic soils

are planted on acidic sites, or vice versa. Selective

grazing by livestock, big game, rodents, and insects has

also eliminated specific ecotypes when they were planted

in small test plots (Edgerton and others 1983; Giunta
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Chapter 22 Rosaceous Shrubs

and others 1978). If nonfire-tolerant ecotypes are planted

on sites subjected to burning, entire stands can be

eliminated in a single fire. Land managers should be

very selective when considering the use of seed collections obtained from areas differing from the prepared

planting site in edaphic or climatic conditions.

Antelope bitterbrush is particularly well adapted to

harsh sites and frequently acts as a pioneer species.

Consequently, it is frequently seeded on mines, roadways, and similar sites. Decumbent and semi-erect

growth forms can be used to control erosion; they

provide a dense, spreading ground cover capable of

spreading by stem layering.

Plants may form symbiotic nitrogen-fixing relationships with the endophyte Frankia (Nelson 1983).

Antelope bitterbrush is more frequently nodulated

than other rosaceous nitrogen-fixing species, but the

degree of nodulation varies considerably among

antelope bitterbrush populations. The significance of

nitrogen fixation in arid land rosaceous species is

poorly understood.

Fire-tolerant ecotypes have been utilized to plant

areas subjected to frequent burning. Most ecotypes

planted for this purpose retain their resprouting characteristics when planted on adapted sites. Consequently, selection programs have been conducted to

identify populations that are capable of resprouting.

Antelope bitterbrush possesses many attributes that

contribute to its widespread use. Seed are large, easy

to collect, process, store, and plant with conventional

equipment. Sufficient seed crops are produced with

frequent regularity to provide satisfactory amounts of

seed for most project plantings. In addition, nursery

and greenhouse-grown transplants are easily produced and establish well when outplanted. Commercial growers produce good quality stock, and with

careful planning, land managers can normally contract production of adapted ecotypes.

The broad array of available growth forms provides

a useful assemblage of material for planting disturbances. With proper selection, adapted ecotypes can

be used to provide windbreaks, serve as conservation

and wildlife plantings, or be used for more formal

maintenance landscape purposes. Different growth

forms can be used to enhance recreational sites, summer homes, and parkways.

Because antelope bitterbrush is a major wildlife

species and establishes quite well, some grasslands

and shrublands have been converted to antelope bitterbrush by artificial seedings. Some plantings have

failed when the shrub was not well suited to the

problem area. Attempts to convert wildland sites to

antelope bitterbrush stands should be carefully

evaluated. Unless adapted ecotypes of antelope bitterbrush exist, more suitable species should be selected

for planting.

Varieties and Ecotypes

Numerous populations of antelope bitterbrush with

distinctive attributes have been recognized and are

commercially harvested and sold (table 2). Ecotypes

from central Utah are widely used for big game habitat

improvement, but are best adapted to calcareous soils.

Decumbent growth forms commonly found as an understory in lodgepole pine or ponderosa pine forests are

used in revegetation of forested sites. Upright ecotypes

from southern Oregon and central Utah are used to

stabilize sandy soils and restrict human activity.

‘Lassen’ antelope bitterbrush, the only release to

date, is a robust, upright ecotype that produces and

maintains moderate quantities of overwintering

leaves that provide a source of protein for fall and

winter grazing (Shaw and Monsen 1986; Welch and

others 1983a). This selection is native to Lassen County,

CA, where it grows on coarse textured, granitic soils.

It has done well throughout the Pacific Northwest and

the Intermountain regions when planted on sandy,

slightly acidic or neutral soils in areas receiving 14 to

18 inches (36 to 46 cm) of annual rainfall. Seedlings

are vigorous and grow rapidly.

Rosa woodsii ultramontana _______

Woods Rose

Hitchcock and others (1961) and Welsh and others

(1987) described Woods rose as a deciduous shrub with

a growth habit ranging from nearly prostrate to upright and height from 3.3 to 10 ft (1.0 to 3.0 m). Its root

system is dense and fibrous. Thick briar patches of

arching branches are formed by heavy suckering from

spreading rhizome systems. Branches are armed with

straight to curved infrastipular spines. They may

produce internodal prickles, but are occasionally unarmed. The alternate leaves are odd pinnate with five

to nine ovate to obovate or elliptic leaflets. The leaves

are 0.6 to 5.1 inches (1.5 to 13 cm) long and 0.2 to 1 inch

(0.4 to 2.5 cm) wide, singly to doubly serrate, and

pubescent to glandular or rarely stipitate glandular.

The fragrant flowers are solitary or in few-flowered

corymbs or corymbiform cymes that terminate lateral

branches of the season (fig. 26). Each flower has five

persistent erect to spreading sepals and five white to

deep rose petals. Numerous stamens are inserted on a

disk lining the throat of the calyx. Carpels are numerous. The “hips” are globose to ellipsoidal and 0.2 to

0.5 inch (6 to 12 mm) in diameter. They turn a bright

orange red at maturity. Each contains 15 to 30 straw

colored to white achenes 0.1 to 0.2 inch (3 to 5 mm) long

with long stiff hairs along one side.

Woods rose blooms from late spring to summer

depending on water availability. Fruits ripen in late

fall and remain on the shrub through winter (Blauer

and others 1975; Hitchcock and others 1961).
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Chapter 22 Rosaceous Shrubs

Table 2—Principal ecotypes of antelope bitterbrush used in wildland restoration projects.

Mean annual Growth Areas of Availability

Ecotype/origin Elevation precipitation form Attributes adaptability Seeda Transplantsb Referencesc

feet inches

Boise Front 2,800 17 Erect Highly productive, Neutral to acidic 4 3 2,3,4

Ada Co., ID good seedling soils, south-central

vigor, excellent Idaho

cover

Bryce Canyon 7,000 16 Spreading to Leafy, highly Colorado River 2 1 3,4

Garfield Co., UT semi-erect available forage, drainage, but

resilient to heavy widely adapted

browsing, excellent

ground cover

Columbia Basin, 12-16 Spreading Excellent drought Columbia River, 2,3

eastern Washington tolerance, highly western and

available forage, central Idaho

good ground cover

Elko 5,200 14 Semi-erect Excellent drought Northern Nevada, 3 2 3,4

Elko Co., NV tolerance, persistent western Utah,

responds well follow- big sagebrush

ing fires, widely and pinyon-juniper

adapted sites

‘Lassen’ 4,200 14 Erect Good winter Mid-elevations, 4 2 5

Lassen Co., CA leaf retention, northern California

seedling vigor, eastern Oregon,

palatability southern Idaho

Maybell 6,000 14 Spreading to Stem layering, Specifically 3 2 3

Moffat Co., CO semi-erect adapted to sandy adapted to

soils, good sandy soils

ground cover

Panguitch Lake 8,250 20 Decumbent Cold tolerant, Spruce-fir,aspen, 2 1

Garfield Co., UT excellent summer and mountain brush

forage, resists types, exposed sites,

browsing, excel- windblown exposures

lent ground cover

Pinto 6,000 14 Upright Drought tolerant, Adapted 2 1

Washington Co., UT productive, good throughout the

forage, excellent pinyon-juniper

wildlife habitat, type within the

consistently good Great Basin

seed crop

Sanpete 5,000 14 Erect, upright Excellent seedling Most widely adapted 4 2 1,3,4

Sanpete Co., UT vigor, highly ecotype tested throughproductive, good out the pinyon-juniper

seed producer, type within the Interpersistent mountain area

Shoshone 4,200 10 Decumbent Drought tolerant, Southern Idaho, 1 1

Lincoln Co., ID spreader, adapted basalt soils

to rocky lava

substrates

Snowville 5,100 15 Semi-erect Cover and forage Adapted to big 3 2

Box Elder Co., UT for big game and sagebrush and

upland birds,excel- pinyon-juniper

lent seed producer communities

Starvation 8,124 18 Decumbent, Highly palatable, Adapted to 2 2 3,4

Canyon spreading excellent summer high elevations,

Wasatch Co., UT forage, resistant aspen openings

to heavy grazing

Salt Lake 5,525 17 Erect, upright Good seed Well-drained 3 2 3,4

Salt Lake Co., UT producer, soils, mountain

forage available brush and upper

elevations of

pinyon-juniper

communities

a

Seed availability: (1) Erratic producer - limited amounts available; (2) Fair to good producer - limited amounts collected; (3) Fair to good producer - moderate amounts collected most years;

(4) Fair to good producer - large collection sites, seed usually available; (5) Good producer - seed normally available. b

Transplant availability: (1) Produced only on request; (2) Produced mainly on request; (3) Produced mainly on request, some speculative rearing; (4) Produced for speculative sales. c

References: (1) Davis (1983a); (2) Edgerton and others (1983); (3) McArthur and others (1983b); (4) Plummer and others (1968); (5) Shaw and Monsen (1986).
588 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Ecological Relationships and Distribution

Rosa is a difficult genus taxonomically. It occurs

throughout the Northern Hemisphere, and exhibits a

high degree of intraspecific variability. There are

approximately 115 species of Rosa in temperate North

America (Young and Young 1986). About 10 of these

occur in the Rocky Mountains and most will hybridize

(Hitchcock and others 1961). Chromosome number for

this group is n = 7. The genus Rosa was represented

in the western American element of the Arcto-Tertiary Flora (Axelrod 1950).

The native range of Woods rose extends from British

Columbia to southern California and east to

Saskatchewan, Minnesota, Missouri, and Texas. There

are two varieties. The eastern variety, Rosa woodsii

var. woodsii occurs on plains and prairies as far west

as eastern Montana. Plants are about 3.3 ft (1 m) tall

with crowded leaves. The leaflets are 0.4 to 0.8 inch

(1 to 2 cm) long and gland tipped. The cordilleran

variety, Rosa woodsii var. ultramontana, covers the

western portion of the species range. It is taller than

var. woodsii, often reaching heights of 3.3 to 6.6

(9.9) ft (1 to 2 [3] m). Its leaves are also larger, up to

2 inches (5 cm) long by 1 inch (2.5 cm) wide with coarse

teeth that are not gland tipped. It exhibits tremendous

Figure 26—Woods rose flowers in late spring or

early summer. Fruits ripen in fall and contain

about 15 to 30 achenes.

variation in height, spininess, vegetative spread, and

drought tolerance. This variety occurs from plains to

alpine areas along marshes, lake shores, and in other

riparian communities of the sagebrush, juniper,

mountain brush, aspen, and spruce-fir zones. Plants

may be found in well drained, wet to dry, loamy to

sandy soils on all aspects, and in moderate shade to

full sunlight. Woods rose grows at elevations from

2,800 to 11,000 ft (850 to 3,355 m) (Welsh and others

1987), generally in areas receiving 16 to 20 inches

(41 to 51 cm) of annual precipitation (Monsen and

Davis 1985; Wasser 1982).

Woods rose invades disturbed areas along roadways, eroded drainages, fence rows, irrigation canals,

and abandoned fields. It is intolerant of poor drainage,

high water tables, and prolonged flooding (Wasser

1982). Ecotypes vary in drought tolerance; flowering

is more adversely affected by drought than is rhizome

growth.

Woods rose resprouts following fire. However, intense fires or repeated burns can damage or kill the

shallow root crown. Annual growth may increase following light to moderate burns (Wasser 1982).

Plant Culture

Woods rose begins flowering when plants are 2 to

4 years old. Plants growing in full sunlight produce

more flowers than those growing in shade. Good seed

crops are produced at about 2 year intervals (Gill and

Pogge 1974b). The hips of Woods rose are harvested by

hand as soon as they ripen and before the flesh begins

to soften. Achenes are extracted by macerating the

hips in water in a blender or Dybvig and floating off

the pulp and empty achenes. Achenes should be dried

thoroughly prior to storage. Acceptable viability is retained for 2 to 4 years by sealing the dry achenes in

containers at 34 to 38 ∞F (1 to 3 ∞C) (Gill and Pogge

1974b). There are about 45,300 seed per lb (99,868 per

kg) at 100 percent purity. Acceptable germination for

seed purchases is 95 percent and purity 70 percent

(see chapter 24).

Dormancy of Woods rose seed is caused by inhibitors

in the seed coverings and mechanical restriction of the

pericarp wall (Jackson and Blundell 1963). In nature,

consumption and digestion of seeds by animals disperse the seed and alleviates dormancy (Morris and

others 1962; Shaw 1974). Overall seed viability is

decreased by digestion, but dormancy of unharmed

seed is reduced. A wet prechilling at 34 to 41 ∞F (1 to

4 ∞C) for 30 to 365 days is recommended to relieve

dormancy of planted seeds (Gill and Pogge 1974b).

Warm pretreatment for 60 days at room temperature

preceding the wet prechilling may also be helpful.

Hot water and acid treatments have been used prior to

wet prechilling with mixed results (Gill and Pogge

1974b).
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Chapter 22 Rosaceous Shrubs

In the field, seed may be broadcast or drilled at a rate

of about 0.5 to 1.0 lb per acre (0.6 to 1.1 kg per ha). It

should be covered with 0.25 to 0.75 inch (0.6 to 1.9 cm)

of soil. Seed can be mixed with other shrub seed, but

it should be seeded separately from grasses; seedlings

are only poorly to moderately competitive with herbs.

Seed may also be spot planted in areas cleared of

competition. Due to variability in seed dormancy among

seed collections, and the extremely long wet prechilling

requirements, results of seedings have been quite

erratic.

Transplanting, although more expensive than direct seeding, is often recommended, at least for small

planting sites. Bareroot stock is usually lifted after

one growing season. Seedlings produce an extensive

root system during the first growing season in the

nursery. They may initiate new rhizome growth prior

to lifting. Rhizome growth is initiated early in spring.

New rhizomes are easily damaged during lifting and

outplanting. Large bareroot seedlings with shoots 2 to

3 ft (0.6 to 0.9 m) tall are grown at some nurseries.

These are useful on unstable or dry sites; the large root

system provides anchoring and a greater water absorption area, and the long shoots are not easily buried

by soil sloughing. Augers or other special planting

tools may be required to properly plant these seedlings. Wildings may be dug when dormant and kept in

cold storage for later transplanting. They should be

handled and planted like bare root stock.

Containerized seedlings may be grown as tublings

or as larger plants. Seedlings may be started from

seed, small transplants, or cuttings (Landis and

Simonich 1984). They require at least a 3 to 5 month

cropping time.

Hardwood cuttings collected when dormant may be

stored at 34 ∞F (4 ∞C) and rooted in the nursery in areas

with mild climates or under a greenhouse mist system

in other areas. Softwood cuttings of partially mature

wood root quickly (10 to 14 days) under a mist system.

Rooted cuttings may be transplanted in fall, overwintered in a shadehouse or cold frame, or transferred

to a nursery bed (Gill and Pogge 1974b). Everett and

others (1978a) recommended the use of hardwood

rather than softwood cuttings. They used a mist system and 0.8 percent indole-3-butyric acid to induce

rooting in a greenhouse. Layers or suckers may also be

propagated in the nursery or greenhouse and field

planted when adequate root systems have developed

(Hartmann and Kester 1990).

Uses and Management

Because of its wide geographic distribution and

revegetation values, Woods rose is a candidate for use

in a wide variety of planting situations. High survival

rates of transplanted seedlings make Woods rose a

desirable and reliable addition to many projects.

Livestock, game animals, and rodent populations may

require control during the period of establishment.

Rodents consume planted seed and young seedlings

sometimes suffer serious debarking and girdling

(Spencer 1958; Wasser 1982). Plants are sometimes

attacked by leaf spots, rusts, downy mildews, blights,

and cankers. Periodic close use, mowing, or burning

under safe conditions may improve the appearance

and stimulate production of plants along roadways or

in recreation areas.

Woods rose is an extremely useful plant on disturbed sites such as roadcuts and fills, gullies, riparian areas, and mining disturbances where soil stabilization is essential (Monsen 1974, 1975; Plummer

and others 1968). At the Leviathan Mine in western

Nevada, Woods rose and mountain snowberry provided the highest third year survival and cover of the

species tested (Everett and others 1980). Suckers were

produced the third year. Plants survived in spite of a

low soil pH (4.1) and low nitrogen. Liming and other

amendments were recommended. Although natural

seedlings may invade disturbances, they are easily

buried by soil sloughing. However, seedlings do develop quite rapidly if established from rooted cuttings

or nursery stock. The fibrous, rapidly developing root

system and rhizomes provide soil stabilization. Rhizomatous spread has been noted by the third year

after transplanting, augmenting the stabilization process. In addition, the brushy habit, thorniness, and

low palatability of some ecotypes permit development

of impenetrable thickets that reduce livestock and

wildlife trampling on unstable sites. Many ecotypes,

however, are grazed rather heavily. Planting them in

a mixture with other shrubs is advised to take advantage of differing growth habits and microsite requirements and maximize the diversity obtained.

Woods rose obtained the highest assessment of suitability of 18 shrubs tested on forest roads in northwestern Montana (Hungerford 1984). The assessment

was based on growth, spread, vigor, survival, flowering, and regeneration. Overall survival after 4 years

was 62 percent with similar survival on south, west,

and east aspects. Plants spread by root sprouting and

were indistinguishable after 4 years. The plant was

recommended for use in this area to hold soil against

wind and water erosion in open to light shade and in

dry to moist soil. Everett (1980) found Woods rose and

antelope bitterbrush were among the best species

tested for revegetating roadcuts in Nevada. After 3

years, Woods rose survival remained high and the

species was among the best for both north and south

slopes when ranked by the amount of cover produced.

The species also exhibited the highest survival of

species tested on forest roadcuts at 14 locations on two

forests in eastern Washington (Tiedemann and others

1976). Overall survival of Woods rose was 56 percent
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Chapter 22 Rosaceous Shrubs

on the Colville National Forest and 63 percent on the

Okanogan National Forest. Plants were generally of

rather low vigor and on these sites may have required

fertilization. Successful planting required stems tall

enough that they were not buried by raveling.

Woods rose is used by big game during most seasons;

it generally receives heaviest use in early spring and

fall (Blauer and others 1975; Wasser 1982). Plants are

moderately grazing tolerant, particularly on moist

soils. Palatability of leaves and stems varies with

ecotype, density of prickles, and availability of other

forage. Wood rose is rated as valuable in winter and

highly valuable in summer and fall for elk (Kufeld

1973). Mule deer make light use of it in winter and fall

(Kufeld and others 1973). In a cafeteria style study,

Smith (1953) found Woods rose was one of the most

preferred shrubs for mule deer throughout the summer (May 1 to September 30) in northern Utah.

Woods rose provides food and excellent nesting,

escape, and thermal cover for birds and small mammals during all seasons (Blauer and others 1975;

Thornburg 1982; Young and Young 1986). Although

not highly preferred, the hips remain on the shrubs

well into the winter and provide an excellent energy

source for many species when the ground is covered

with snow (Mace and Bissell 1986; Monsen and Davis

1985; Plummer and others 1968; Welch and Andrus

1977).

Nutritive value of Woods rose varies considerably by

season. In the Black Hills of South Dakota, protein

value varied from 4.9 percent in summer to 12.0

percent in spring for stems and from 5.7 percent in fall

to 16.4 percent in spring for leaves (Dietz 1972). Rose

hips have high digestibility and are moderately high

in crude protein in winter; they provide food for many

birds and rodents.

Rose hips have been used as preserves and as an

ingredient in herbal tea and candy. They are high in

vitamin C. American Indians used the bark, roots, and

stems to produce a dressing for wounds, sores, and

burns. Roots were used for treating diarrhea (Craighead

and others 1963). Woods rose has high ornamental

value for recreation areas, parks, and campgrounds.

The dense growth habit can also be used to direct

pedestrian traffic and provide low maintenance

shrubbery. The attractive fruits, flowers, and fall

coloration also contribute to its landscape value.

Varieties and Ecotypes

No populations of Woods rose have been released for

commercial production. Tremendous variability occurs

in growth habit and many other characteristics. Some

accessions have a wide range of adaptability.

Several additional Rosa species are used in wildlife

plantings. Rosa nutkana is distributed from Alaska

south to northern California and the Blue Mountain

region of Oregon. It also extends south into Colorado

and Utah at moderate elevations. Rosa acicularis, a

circumpolar species, extends south from Idaho and

Montana to northern New Mexico. Rosa gymnocarpa

occurs from sea level to 6,000 ft (1,830 m) from British

Columbia to northwestern Montana, south to the

Sierra Nevada Mountains of California and the Blue

and Ochoco Mountains of eastern Oregon (Hitchcock

and others 1961). It is a larger shrub with a more erect

and less thicket-like growth habit than Woods rose.

Uses and culture of these species are generally as

described for Woods rose.

Rubus leucodermis ______________

Black Raspberry, Blackcap

Black raspberry or blackcap is a sprawling, deciduous shrub with arching biennial canes ranging from

3 to 10 ft (1 to 3 m) in length (Hitchcock and others

1961). Prickles of the leaves, petioles, and stems are

retrorsely curved. The pinnate leaves are petiolate

with linear, caducous stipules, and three to five

doubly serrate leaflets. Leaflets are green and nearly

glabrous above and white tomentose beneath. They

are about 2.4 to 5.6 inches (6 to 14 cm) long; the

terminal leaflet is 1 to 3 inches (3 to 7.5 cm) long.

Perfect, 5-merous white flowers are produced in

racemes in groups of two to seven. Stamens and pistils

are numerous. Fruits consist of aggregates of black

drupelets on a convex receptacle. Each drupelet contains a hard seed. Flower and fruit development occur

unevenly through the summer. Seed is dispersed by

birds and mammals.

Ecological Relationships and Distribution

Rubus is distributed worldwide, but is most common

in temperate regions of the Northern Hemisphere.

There are several hundred species. Many taxa are

interfertile; most are polymorphic, and a few are

apomictic. Rubus belongs to the subfamily Rosoideae

and has chromosome number n = 7 (McArthur and

Sanderson 1985).

Black raspberry is distributed from British Columbia

east to Montana and south to California and New

Mexico (Davis 1952). Although commonly a plant of

moist environments such as riparian zones and forest

openings, black raspberry also exists on dry, open

slopes of mountain brush communities and on talus

slopes or roadbanks. It spreads by layering and rhizome sprouting as well as from seed and may be one

of only a few species growing on some rocky sites.

Uses and Management

Ferguson (1983) considered black raspberry the most

likely of the Rubus species to be used for revegetation
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Chapter 22 Rosaceous Shrubs

purposes. It is easily propagated vegetatively and

transplants readily. On roadcuts, drainageways, or

other disturbed sites it grows rapidly, spreads vegetatively, and provides soil stabilization as well as dense

cover and food for birds and other small animals. Black

raspberry grows rapidly and is used only lightly as a

summer forage by big game and livestock, thus its

establishment is normally not hindered by excessive

use.

Plant Culture

Seed dormancy and cultural practices for black raspberry are similar to those described for thimbleberry.

Time requirements for wet prechilling and scarification have not been determined. Fruits are collected

when they are black and detach easily from the receptacle. Use of planting stock rather than seed on disturbed sites may provide better and more rapid success; these sites offer poor conditions for seed

germination and early seedling establishment. Planting stock may be grown from seed. Root cuttings,

suckers, layers, stem cuttings, and leaf-bud cuttings

are easily started (Hartmann and Kester 1990). Root

cuttings may be treated as wildings, particularly on

moist sites. Seedlings grow fairly rapidly, but are

sensitive to dense competition and shading.

Varieties and Ecotypes

There are no releases.

Rubus parviflorus _______________

Thimbleberry

Thimbleberry is a low growing, rhizomatous, thicketforming shrub. It is unarmed, deciduous, and 2 to 7 ft

(0.5 to 2.0 m) tall (Hitchcock and others 1961). Bark of

the main stems is gray to brown and shreddy. Stems,

petioles, and leaves are unarmed and stipitate glandular. The large leaves are long petioled with membranous lanceolate stipules. They are alternate, simple,

palmately lobed, and veined from a cordate base.

Leaves are 2 to 6 inches (5 to 15 cm) long and 0.6 to 2

inches (1.5 to 5 cm) wide. The showy, white flowers are

perfect and 5-merous, in groups of two to seven in flattopped cymes or terminal corymbs (fig. 27). Stamens

and pistils are numerous. Fruits are approximately

0.5 to 1 inch (1.2 to 2.5 cm) wide aggregates of coherent

red drupelets that are thinly fleshy and almost dry at

maturity. Fruit aggregates form over a conical receptacle, thus the common name “thimbleberry.” Each

drupelet contains a single hard beige to pink seed

about 0.1 inch (2 mm) long. Flowering and fruit ripening continue through the summer. Seeds are dispersed

by birds and mammals.

Figure 27—Thimbleberry spreads by seed or

resprouting in natural forest openings and disturbed

areas.

Ecological Relationships and Distribution

Thimbleberry is distributed from Alaska east to

Ontario and south to Chihuahua, Mexico. It grows in

all western States from elevations near sea level on

the Pacific Coast to near 10,000 ft (3,050 m) in Colorado

(Harrington 1964; Sutton and Johnson 1974; USDA

Forest Service 1937). Thimbleberry forms dense

patches in natural forest openings or disturbed areas,

but it is also fairly common in areas shaded by forest

canopies, brushfields, or woody riparian plants, particularly in the southern portion of its range. It is a

particularly common understory species in the humid

forests of the Pacific Northwest.

Thimbleberry grows on moist, sandy loam soils rich

in organic matter, but may also be found on rocky talus

slopes, road banks, and creek bottoms. It regenerates

rapidly from rhizome and root crown sprouting following burning or logging (Cholewa and Johnson 1983;

Leege and Hickey 1971; Mueggler 1966; Noste and

Bushey 1987). Buried thimbleberry seeds have been

reported in a number of forest seed bank studies
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Chapter 22 Rosaceous Shrubs

(Kramer 1984). Although Kramer (1984) found low

densities and low viability of thimbleberry seeds in

soil banks for three forest habitat types in central

Idaho, he concluded that the rhizomatous nature of

the species would allow even isolated seedlings to

spread over significant areas.

Plant Culture

Good seed crops are produced almost every year but

fruits are difficult to collect in large quantities as they

are scattered over the crown of the plant and ripen

unevenly. Ripe fruits are bright red and detach easily

from the receptacle. They are dispersed by gravity and

by animals that consume them.

Fruits are hand collected. Seeds are extracted by

macerating the fruits in water in a Dybvig or blender

and floating off the pulp. Remaining debris and light

seed are removed by drying and fanning. Dried seeds

may be stored at 41 ∞F (5 ∞C) for several years (Brinkman

1974h).

Dormant embryos and an impermeable endocarp

prevent rapid seed germination. In nature, dormancy

is overcome by warm pretreatments and wet prechilling

occurring between early summer seed fall and spring

germination, although seeds are capable of remaining

viable in the soil for more than 1 year if germination

conditions are not met (Brinkman 1974h). A 2-hour

soak in sulphuric acid or a 7-day soak in 1 percent

sodium hypochlorite are commonly used germination

pretreatments. However, results of acid scarification

have been erratic (Marchant and Sherlock 1984).

Swingle (1939) and Babb (1959) recommended planting the seed immediately after it is harvested. Summer seeding in the nursery provides warm pretreatments and wet prechilling that relieve both blocks to

germination for many Rubus species (Babb 1959;

Brinkman 1974h). A 90-day warm pretreatment at 68

to 86 ∞F (20 to 30 ∞C), followed by a 90 day wet

prechilling at 41 ∞F (5 ∞C) in the laboratory, prepares

seed for spring sowing.

Thimbleberry is easily established from seed-grown

bareroot or container stock. Plants are also easily

propagated from layers or suckers. Sprigging, the collection and planting of dormant rhizomes, provides a

means of quickly obtaining large quantities of plants.

Uses and Management

Rubus was ranked first among 18 genera of shrubs

in terms of the number of wildlife species using them

(Martin and others 1951; Robinette 1972). The drupelets are sought by birds and many other animals;

leaves are used to varying degrees in summer. The

dense sprawling plants provide cover for small animals during all seasons. Kufeld and others (1973)

reported that mule deer take trace amounts of

thimbleberry in fall and moderate amounts in summer. Thimbleberry is ranked as highly valuable and

valuable for elk in summer and fall, respectively

(Kufeld 1973). Palatability for livestock varies with

geographic location and plant association. In general, it is considered worthless to poor for cattle and

fair for sheep, although it may be heavily used if other

vegetation is scarce or unpalatable (USDA Forest

Service 1937).

Primary values of thimbleberrry in revegetation

plantings are to provide soil stabilization and thickets

for wildlife food and cover on disturbed forest sites,

sand dune areas, and streambanks. Seedlings are

not likely to be browsed heavily unless other more

palatable vegetation is unavailable. They are susceptible to white fly and spider mite infestations

(Marchant and Sherlock 1984).

Thimbleberry is a common plant of seral shrub

communities; it resprouts following logging or burning, and establishes in natural forest gaps. Burned

plants resprout from root crowns and rhizomes. Intense fires might destroy these structures, but permit

germination of seed banked in the soil (Noste and

Bushey 1987). Gratkowski (1978), Leege and Hickey

(1971), and Miller and Kidd (1983), discussed the

response of thimbleberry to burning and herbicide

treatments, practices used to kill shrubs and permit

conifer regeneration or rejuvenate overgrown shrub

fields to improve browse availability for wildlife.

Varieties and Ecotypes

None.

Sorbus scopulina _______________

Greene’s Mountain Ash

Greene’s mountain ash is an attractive semi-erect

shrub, branching from the base and forming thickets

by suckering or layering (fig. 28). Bark of younger

stems is smooth and yellowish gray becoming grayish

red on older stems. Young growth is sparingly to

densely grayish strigillose pilose. Winter buds are

glutinous and white hairy to glabrous. The alternate

deciduous leaves are glossy dark green above and pale

and glabrous beneath, turning a showy red orange in

autumn. They are pinnately compound with 7 to 13

serrate leaflets each, short cuneate to acute at the

base, and acuminate at the tip. Stipules are green,

membranous, and caducous to persistent. The inflorescence is a 70 to 200 flowered flat-topped corymb.

The creamy-white flowers are perfect, 5-merous, and

regular (Preston 1968). Hypanthium lobes are triangular, persistent, and white hairy externally. There

are 15 to 20 stamens and two to five carpels and

styles. Fruits are fleshy scarlet to orange pomes that

dry to a purplish color. Each contains four seeds
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Chapter 22 Rosaceous Shrubs

Figure 28—Greene’s mountain ash fruits persist

into winter and are consumed by birds and other

wildlife species.

(Harris and Stein 1974; Hitchcock and others 1961;

Welsh and others 1987). The shrubs flower from May

to July and fruits mature from August through October. Many fruits remain on the plant through winter.

Seeds are spread primarily by birds.

Ecological Relationships and Distribution

Approximately 50 species of Sorbus are distributed

through temperate and subarctic areas of the Northern Hemisphere (Hitchcock and others 1961). The

chromosome number n = 17 is typical of woody

Pomoideae genera in the western United States

(McArthur and Sanderson 1985). Greene’s mountain

ash is distributed from Alaska east to Alberta and

south to California, New Mexico, and the Dakotas

(Welsh 1982). It grows in openings in conifer forests

from foothills to subalpine or alpine sites at 4,000 to

9,000 ft (1,200 to 2,700 m) (Davis 1952; Stark 1966;

Welsh 1982). Greene’s mountain ash grows in sun or

partial shade on well-drained, neutral to acidic soils

forming dense patches or intermixed with other shrub

species (Van Dersal 1938). It is common in mountain

brush communities of forest openings, riparian areas,

and on disturbed sites.

Plant Culture

Fruit should be gathered immediately when ripe to

prevent losses to birds. Seeds are collected by hand or

shaken onto a canvas. Maceration of fruits in water in

a Dybvig or commercial blender separates seeds from

the remainder of the fruit. The pulp contains germination inhibitors and is removed by flotation and by

fanning and screening the dried seed and trash

(Hartmann and Kester 1990; Heit 1968). Seed quality

is often low. Seed should be dried to a 6 to 8 percent

water content and stored dry in sealed containers at 34

to 38 ∞F (1 to 3 ∞C) (Harris and Stein 1974). Properly

stored seeds remain viable for 2 to 8 years.

Seed requires wet prechilling for germination. A 72-

hour soak in gibberellic acid (3.46 g Rootone-F/1000cc/

distilled H2O) followed by a 90-day wet prechilling at

41 ∞F (51 ∞C) was recommended by SEAM (1976) and

Belcher (1985). Seed can also be given a 115-day warm

pretreatment at 77 ∞F (25 ∞C) followed by a 75-day wet

prechilling at 41 ∞F (5 ∞C), although exact requirements vary widely between and within seedlots. Laboratory germination tests require extended incubation

periods. Excised embryo tests and tetrazolium chloride tests are frequently used alternatives; both can be

completed rapidly (see Belcher 1985). Results of both

tests are generally higher than laboratory germination test results (Harris and Stein 1974).

Greene’s mountain ash may be established by sowing untreated seed in late summer to provide warm

and cold wet pretreatment periods. Laboratory

prechilled seed may be spring planted. Seed should be

covered lightly. Even when wet prechilled, seed lots

often germinate in the field over a 2 to 3 year period.

Seed should be planted in areas where it will be

protected from competing vegetation.

Production of planting stock may be necessary if

seed supplies are limited or more rapid establishment

is desired. Container and bareroot stock are usually

grown from seed. Bareroot stock develops slowly and

may require two growing seasons before lifting. Propagation by cuttings and layering is difficult. Wildings

may be used as alternative sources of planting material if only small quantities of plants are needed.

Cook (1981) recommended use of Greene’s mountain

ash on adapted sites below 6,500 ft (2,000 m) in

Wyoming, and suggested that it be planted at 4 ft

(1.2 m) spacings. Young plants develop rather slowly

and require protection from competition with grasses

and forbs. Seedlings are tenacious and evidently not

very vulnerable to insect and disease problems. They

are taken by deer (USDA Forest Service 1948).

Uses and Management

The persistent fruits of Greene’s mountain ash are

taken by bears and birds in the fall and into the winter.

Moose often browse the shrubs heavily in winter.

Twigs are eaten by game and livestock. Dayton (1931)

reported that Greene’s mountain ash produces good

sheep feed. It provides good cover and soil stabilization

on steep sites within its range as it spreads to form

dense thickets. As an ornamental, it offers attractive

foliage, flowers, and fruits for mass plantings in full

sunlight or partial shade. The foliage and berries are

brightly colored in fall, and clusters of berries remain

on the plant through the winter (Sutton and Johnson

1974).
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Chapter 22 Rosaceous Shrubs

Miller (1981) and Miller and Kidd (1983) examined

the response of Greene’s mountain ash on seral shrub

fields in northern Idaho to various herbicide treatments. Lyon (1966) found that plants were eliminated

by burning.

Varieties and Ecotypes

Various selections are used for horticultural

plantings, but none have been developed for wildlife or

wildland uses.

Sorbus sitchensis _______________

Sitka Mountain Ash

Sitka mountain ash grows from 3 to 13 ft (1 to 4 m)

tall with a rounded crown. It may be distinguished

readily from Greene’s mountain ash; leaves consist of

7 to 11 leaflets, each semitruncate to rounded at the

tip and usually at least one-third as broad as long. The

winter buds, young growth and inflorescences are

rufous hairy; the calyx is glabrous externally. Stipules

are persistent and the fruits are red with a slight

bluish cast (Hitchcock and others 1961; Preston 1968).

They are dispersed by birds, but some persist on the

tree through winter.

Sitka mountain ash is distributed from Alaska and

the Yukon south to the Cascades in northern California and east to British Columbia and northwestern

Montana at elevations from 2,000 to 10,000 ft (610 to

3,050 m) (Hitchcock and other 1961). Although it

frequently grows in wet soils along streams, it may

also be found in forest openings created by natural

forest gaps, logging, burning, or other disturbances.

Sitka mountain ash produces abundant seed crops.

Harvesting, cleaning, storage, germination, and planting requirements are similar to those described for

Greene’s mountain ash. Seeds sown late or without

adequate wet prechilling do not germinate. Approximately 1 to 5 lb (0.45 to 2.25 kg) of seed may be

extracted from 100 lb (45 kg) of fruits (Harris and

Stein 1974).

Marchant and Sherlock (1984) considered the species a useful high altitude shrub for rocky sites. Management and use of the species are as described for

Greene’s mountain ash. It is frequently cultivated as

an ornamental.

Spiraea betulifolia _______________

Bridal Wreath

Bridal wreath spiraea, birchleaf spirea, or white

spiraea is a low, glabrous, deciduous shrub or subshrub ranging from 0.5 to 2 ft (0.2 to 0.6 m) in height

Figure 29—Bridal wreath spiraea produces flat

corymbs of white flowers.

(Hitchcock and others 1961). Plants occur singly or in

dense patches interconnected by rhizomes. Stems are

slender and yellowish brown to brownish with few

branches. The simple, alternate, oval to obovate leaves

range from 1 to 2.8 inches (2 to 7 cm) in length; the

largest leaves occur near the tips of vegetative and

flowering branches. The leaves immediately subtending the inflorescence tend to be slightly smaller. Leaves

are short petiolate, exstipulate, strongly veined, and

coarsely or doubly serrate to shallowly lobulate along

the upper one-half to two-thirds of the leaf. Upper leaf

surfaces are smooth and shiny; lower surfaces are

pale.

Inflorescenses are dense, flat-topped terminal

corymbs 1 to 3.2 inches (3 to 8 cm) in diameter (fig. 29).

The tiny, white to pinkish 5-merous flowers are perfect, perigynous, and regular. Stamens are numerous

and exceed the petals. Fruits consist of five light

brown, dehiscent, several-seeded follicles. Flowering

occurs from June to early August; fruits ripen from

mid July to early September. Fruits normally dehisce

in October and seeds are shed over time as winter
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Chapter 22 Rosaceous Shrubs

storms dislodge them from the follicles (Hitchcock and

others 1961; Stickney 1974c; USDA Forest Service

1937).

Ecological Relationships and Distribution

The genus Spiraea includes about 70 species distributed through temperate and cooler areas of the

Northern Hemisphere. About 10 species are found

in the western United States, concentrated in midelevations of mountainous areas. They are particularly abundant in the Douglas-fir region of the western

United States. The genus is a member of the subfamily Spiraeoideae. Chromosome number of the

North American species is n = 9. Asian species commonly have chromosome numbers of n = 9 or n = 7

(McArthur and Sanderson 1985).

Bridal wreath spiraea occurs from sea level to

11,000 ft (3,300 m) and is distributed from British

Columbia to north-central Oregon and east to

Saskatchewan and South Dakota. It grows on a wide

range of sites from riparian areas to open or wooded

valleys, hills, and rocky slopes. It is particularly abundant on logged, burned, or otherwise disturbed mesic

forest sites at intermediate elevations. It is the dominant understory for a number of forest habitat types

and community types in the Rocky Mountains where

ponderosa pine, lodgepole pine, grand fir, subalpine

fir, and aspen are the dominant overstory species.

Common shrub associates include roses, huckleberries, willows, and snowberries.

Plant Culture

Bridal wreath spiraea grows in full sun or shaded

areas, although most seed is produced on shrubs

growing in full sunlight. Cleaning involves rubbing

the follicles to release the seeds and sieving to separate the seeds from the trash. The tiny seeds are

nondormant. Germination reportedly occurs following 120 days of wet prechilling at 32 to 34 ∞F (0 to 1 ∞C)

(Stickney 1974b); this mechanism permits emergence

shortly after snowmelt. Bridal wreath spiraea may be

planted by spot seeding disturbed sites in fall or

spring. Seed may be mixed with other shrub seeds, but

should be planted separately from grasses and forbs.

Propagating and planting bareroot or container

stock provides for better utilization of the normally

limited seed supplies and more rapid development of

cover on disturbed sites than does direct seeding.

Bareroot stock may be propagated from seed. Hand

seeding may be required in the nursery; it is difficult

to clean the seed to a high level of purity and the light

seed does not flow well through a drill. Seed should be

covered very lightly and mulched. Bareroot stock may

also be obtained by lining out hardwood cuttings in the

spring. Container stock is grown from germinants or

from hardwood or softwood cuttings (Babb 1959).

Hartmann and Kester (1990) recommended the use of

leafy softwood cuttings taken in summer. Cuttings

are treated with a root-promoting substance and

rooted under high humidity. Container and 1-0 or 2-0

bareroot seedlings develop rapidly and are easily

handled and transplanted.

Uses and Management

Although it produces abundant available forage,

bridal wreath spiraea has been rated as worthless for

cattle and poor to fair for sheep (USDA 1937). The

dried leaves persist on the plant in fall and may receive

moderate use after leaves have fallen from other

species. Kufeld and others (1973) reported bridal

wreath spiraea receives light winter use and heavy

summer and fall use by mule deer. Because of its low

palatability, livestock and wildlife rarely damage established stands of bridal wreath spiraea. Consequently, it may become abundant in heavily grazed

sites (USDA Forest Service 1937), spreading vegetatively and from seed.

Spiraeas, particularly those species that spread

rhizomatously, are valuable plants for soil stabilization along waterways. Adapted species or varieties

may be selected for specific areas and conditions.

Bridal wreath spiraea has also been used to stabilize

roadways, burns, and logged areas. It develops rapidly, producing a spreading root system and rhizomes.

The value of this and other spiraea species for site

stabilization is increased by its typically low palatability to game and livestock.

A largely unexploited potential of native spiraea is

their use as landscape plants. They are easily established and maintained and produce attractive foliage

and showy flowers, although flowering is normally

restricted to a short period.

Plants developing from seed or transplants mature

rapidly. Those growing in full sunlight may flower and

produce seed during the first or usually the second

growing season. Although seedlings are reasonably

vigorous, their growth may be restricted by competition with weeds or seeded grasses. Young seedlings

may be trampled by game or girdled by rodents.

Most horizontal rhizomes and fibrous roots are located beneath the mineral soil surface. After fires or

other injury, rhizomes sprout vigorously just behind

the damaged tissue. Roots beneath the surface of

mineral soil also resprout (Bradley 1984; McLean

1967; Fischer and Clayton 1983). Merrill (1982) reported white spiraea achieved a greater volume and

biomass on burned sites relative to unburned sites

1 year after burning. This difference was maintained

over a 4 year study. Rate of recovery was related to

moisture conditions.

Varieties and Ecotypes

None.
596 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 22 Rosaceous Shrubs

Spiraea densiflora _______________

Subalpine Spiraea

Subalpine spiraea occurs from British Columbia

south to northern California, Idaho, and Wyoming at

elevations ranging from 2,000 to 11,000 ft (600 to

3,300 m) (Hitchcock and others 1961). It grows along

lakes and waterways and on wooded to open rocky

slopes, frequently in association with lodgepole pine

and western white pine. It is a low, highly branched

shrub 1.6 to 3.3 ft (0.5 to 1 m) tall with strong root

stocks (fig. 30). Flowers develop in small flat topped

to rounded pink to rose corymbs 0.8 to 1.6 inches (2 to

4 cm) wide. Young stems, inflorescences, and lower

surfaces of leaves are thickly puberulent. Most material in the Intermountain region belongs to var.

splendens.

Spiraea douglasii _______________

Douglas Spiraea

A relatively tall species, Douglas spiraea grows from

3 to 8 ft (1 to 2.4 m) in height with erect stems

branching primarily below the center. The species

spreads by rhizomes, often forming thickets in openings along lake and stream margins, bogs, and swamps.

It grows on boggy or peaty ground and wet heavy soils

subject to annual flooding (Marchant and Sherlock

1984). Stems are pubescent to glabrous. The oblongelliptic leaves are serrate above the center. They are

1.6 to 4 inches (4 to 10 cm) in length, dark green

above, and woody beneath. The inflorescence is a

pyramidal rose to pink panicle 1.6 to 11.8 inches (4 to

30 cm) long with numerous tiny flowers (fig. 31). Dry

flower parts are retained as brown tufts overtopping

Figure 31—Elongate panicles of pink to rose flowers characterize Douglas spiraea.

Figure 30—Subalpine spiraea inflorescences

are thickly puberulent, flat to rounded corymbs

of pink to rose flowers.

the shrub in winter. Fruits are dry, glabrous, beaked

follicles. Seeds are green to brown and less than 0.1

inch (2 mm) long.

Ecological Relationships and Distribution

Douglas spiraea is distributed from Alaska south to

northern California and Montana, occurring primarily in coastal areas (Davis 1952; USDA Forest Service

1937).

Plant Culture

Douglas spiraea is easily propagated from cuttings.

Marchant and Sherlock (1984) obtained 98 percent

rooting of softwood cuttings under mist in June and 98

percent rooting of hardwood cuttings in fall. They

recommended lining out hardwood cuttings in spring

and lifting them the same fall for outplanting. Stock

could also be spring lifted and outplanted. Sprigs or

chunks of rhizomes can be dug when dormant and

spring planted in areas with moist soils.

Uses and Management

Because of their large size, rhizomatous habit, and

ease of propagation, both varieties of Douglas spiraea

are prime candidates for revegetation of riparian areas, where adapted. They also have potential for use as

ornamentals.

Varieties and Ecotypes

Spiraea douglasii var. menziesii is separated from

var. roseata by its tomentulose inflorescence. It occurs

from southern Alaska to northwestern Oregon and

northern Idaho. It is one of the most palatable spiraeas

and is rated as fair to good fall forage for cattle and

sheep (USDA Forest Service 1937). The variety roseata

occurs primarily in central Idaho (Hitchcock and others 1961). It is a smaller and less palatable shrub.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 597

Stephen B. Monsen

Richard Stevens

Nancy L. Shaw

23

Chapter

Introduction _____________________________________

Numerous genera and species of shrubs occur throughout the

Intermountain region in addition to those included in the

Asteraceae, Chenopodiaceae, and Rosaceae families. Although

shrubs are widespread throughout this region and dominate

many areas, species richness is low compared to the shrub flora of

the Pacific United States, Chile, western Australia, and South

Africa (Stebbins 1975). Generally, evolution proceeds most rapidly

when populations are isolated from one another and exposed to

different environmental conditions (Dobzhansky 1970; Stebbins

1950); this is the case in the Intermountain region. However,

fewer numbers of species within this region are, in part, due to

the relatively recent advance and retreat of continental seas,

drastic environmental changes, and instability that result in a

high rate of extinction (Stebbins 1975).

Most species included in this chapter have been used in revegetation programs. Many other shrubs are recognized as important,

but information related to their propagation, seed production,

germination, or establishment by direct seeding is not available.

Shrubs of Other

Families
598 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Chapter Contents by Species

Acer glabrum (Rocky Mountain maple)............................................................................................. 600

Rhus aromatica (Skunkbush sumac)................................................................................................. 608

Rhus glabra (Rocky Mountain smooth sumac) ................................................................................. 611

Mahonia aquifolium (Shining barberry) ........................................................................................... 613

Mahonia fremontii (Fremont barberry)............................................................................................. 614

Mahonia repens (Oregon grape, creeping barberry, mountain holly, creeping mahonia) .............. 614

Alnus incana (Thinleaf alder) ............................................................................................................ 616

Betula glandulosa (Bog birch)............................................................................................................ 618

Betula occidentalis (Water birch) ...................................................................................................... 620

Betula papyrifera (Paper birch) ......................................................................................................... 622

Lonicera ciliosa (Orange honeysuckle) .............................................................................................. 623

Lonicera involucrata (Bearberry honeysuckle) ................................................................................. 623

Lonicera utahensis (Utah honeysuckle) ............................................................................................ 624

Sambucus cerulea (Blue elderberry).................................................................................................. 624

Sambucus racemosa (Red elderberry) ............................................................................................... 625

Symphoricarpos albus (Common or white snowberry) ..................................................................... 627

Symphoricarpos longiflorus (Desert snowberry) .............................................................................. 627

Symphoricarpos occidentalis (Western snowberry, wolfberry) ........................................................ 628

Symphoricarpos oreophilus (Mountain snowberry) .......................................................................... 629

Cornus stolonifera (Redosier dogwood).............................................................................................. 630

Cupressus arizonica (Arizona cypress) .............................................................................................. 632

Juniperus horizontalis (Creeping juniper) ........................................................................................ 633

Juniperus osteosperma (Utah juniper) .............................................................................................. 635

Juniperus scopulorum (Rocky Mountain juniper) ............................................................................ 638

Elaeagnus commutata (Silverberry) .................................................................................................. 640

Shepherdia argenta (Silver buffaloberry).......................................................................................... 641

Shepherdia canadensis (Russet buffaloberry, soapberry) ................................................................ 642

Shepherdia rotundifolia (Roundleaf buffaloberry) ........................................................................... 643

Ephedra nevadensis (Nevada ephedra) ............................................................................................. 644

Ephedra viridis (Green ephedra) ....................................................................................................... 645

Arctostaphylos uva-ursi (Bearberry manzanita) ............................................................................... 646

Quercus gambelii (Gambel oak) ......................................................................................................... 648

Fraxinus anomala (Singleleaf ash).................................................................................................... 650

Eriogonum heracleoides (Wyeth or whorled eriogonum, buckwheat).............................................. 652

Eriogonum umbellatum (Sulfur eriogonum, buckwheat) ................................................................. 653

Eriogonum wrightii (Wright eriogonum, buckwheat) ...................................................................... 654

Clematis ligusticifolia (Western virginsbower)................................................................................. 654

Ceanothus species ............................................................................................................................... 656

Ceanothus cuneatus (Wedgeleaf ceanothus) ..................................................................................... 656

Ceanothus greggii (Desert ceanothus) ............................................................................................... 657

Ceanothus integerrimus (Deerbrush ceanothus)............................................................................... 658

Ceanothus lemmonii (Lemmon ceanothus) ....................................................................................... 659

Ceanothus prostratus (Prostrate ceanothus)..................................................................................... 659

Ceanothus martinii (Martin ceanothus) ............................................................................................ 661

Ceanothus sanguineus (Redstem ceanothus) .................................................................................... 662

Ceanothus velutinus (Snowbrush ceanothus) ................................................................................... 665

Salix species (Willow) ......................................................................................................................... 668

Salix bebbiana (Bebb willow, beaked willow) ................................................................................... 670

Salix boothii (Booth willow) ............................................................................................................... 672
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 599

Chapter 23 Shrubs of Other Families

Salix drummondiana (Drummond willow, beautiful willow, blue willow) ..................................... 673

Salix exigua (Coyote willow, dusky willow, narrowleaf willow, sandbar willow) ........................... 675

Salix geyeriana (Geyer willow) .......................................................................................................... 676

Salix glauca (Grayleaf willow, glaucous willow) .............................................................................. 677

Salix lasiandra var. caudata (Whiplash willow) .............................................................................. 679

Salix lasiolepis (Arroyo willow) ......................................................................................................... 680

Salix lutea (Yellow willow, shining willow) ...................................................................................... 681

Salix planifolia (Plainleaf willow) ..................................................................................................... 683

Salix scouleriana (Scouler willow, fire willow, black willow, mountain willow, nuttall willow) ... 684

Salix wolfii (Wolf willow) ................................................................................................................... 686

Philadelphus lewisii (Lewis mockorange) ......................................................................................... 687

Ribes aureum (Golden currant) ......................................................................................................... 689

Ribes cereum (Wax currant) ............................................................................................................... 691

Ribes viscosissimum (Sticky currant) ................................................................................................ 693

Penstemon fruticosus (Bush penstemon) ........................................................................................... 694

Celtis reticulata (Netleaf hackberry) ................................................................................................. 696

Consequently, they are not widely used in large-scale

seedings where uniform stands are expected. A number of important species could be more widely utilized

in artificial revegetation and increased through management if more information related to their culture

was available.

Sixty-seven species representing 20 families are

discussed in this chapter. These species occur over a

wide range of conditions and habitats. Certain species

are common and widespread; others are less abundant

and often restricted to localized conditions. However,

all are important. In combination, these shrubs offer a

diverse range of material for planting and management. Eriogonum and Ceanothus are examples of two

widespread genera discussed in this chapter. Approximately 130 species of Eriogonum, including shrubs

and broadleaf herbs, occur within the Intermountain

region (Welsh and others 1987). Of these, fewer than

10 have been evaluated or used for range or wildland

plantings (Miles and Meikle 1984; Plummer and others 1968; Sampson and Jespersen 1963). By contrast,

about two dozen species of Ceanothus are reported in

this same region and slightly more than half have been

studied for wildlife, watershed, or horticultural uses.

In most instances only one or two species of any genus

discussed in this chapter have been studied as potential revegetation species. Seed or planting stock of

most species is often relatively scarce.

Shrubs discussed in this chapter occupy rather specific sites. Mountain snowberry, Rocky Mountain

maple, and Gambel oak are confined to specific

elevations and climatic conditions, yet occupy rather

large acreages, often as the dominant species. Other

shrubs, including green ephedra and singleleaf ash,

are common throughout the southern portion of the

Intermountain area but do not extend into the colder

northern section. By contrast, red elderberry and dwarf

snowberry occupy only mountainous situations.

A characteristic of many shrubs discussed in this

chapter is their close association or relationship to soil

or site conditions. Rather extensive areas of willow

occur through many vegetative communities but are

most often confined to moist soils. The relationship of

blue elderberry, bush penstemon, and other shrubs

that are also adapted to specific soil conditions is less

well understood.

Introduced grasses have not been found to be as

specifically adapted as many of these shrubs. This is

an important consideration; some shrub-dominated

associations have been planted to introduced species,

mostly grasses. Attempts have been made to substitute other species for native shrubs. Many substitute

plants do not provide the forage or cover values expected or simply are not adapted for long-term persistence on some sites. Thus, it is important to learn

restoration and management practices required to

perpetuate endemic shrub associations.
600 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Some shrubs are particularly important during

various stages of succession. Species of redstem

ceanothus, snowbrush ceanothus, and red elderberry

recover well following fires and assume dominance

after coniferous forests have burned. These shrubs

provide important habitat, but unless they are able to

perpetuate during cyclic fire events, game and livestock habitat may eventually diminish as conifers

gain dominance.

Many shrubs occur as components of associated

forest and herbaceous communities. Considerable fluctuation in species composition often occurs as a result

of fires, and from extremes in climatic events, grazing,

and disease. The opportunity to reestablish plants on

these disturbances is, in part, dependent upon use of

ecotypes with appropriate adaptive features. In an

attempt to enhance the shrub component where desirable species have been lost, planting in some situations, has been very successful. Characteristics that

ensure the survival of shrubs under adverse conditions are important in selecting plants to rehabilitate

mine disturbances and other harsh sites.

Some shrubs are often the only or dominant species

to occur on harsh and disturbed sites. Soils throughout

many areas of the Intermountain region are infertile

and do not support large numbers of species. It is

unlikely that successional changes will result in the

replacement of many shrubs with other, equally

adapted species in such areas. Certain species of

buckwheat and penstemon are able to populate harsh,

exposed sites. Many of these sites are important

midwintering areas where high quality forage is essential for big game animals. In addition, these exposed sites are often invaded by weeds that may enter

and, subsequently, spread to adjoining locations. Without the presence of adapted species, weedy plants can

invade open and seemingly harsh situations.

Many shrubs are important forage plants. Some are

browsed readily throughout the year; others are selectively grazed during certain seasons. For example,

Gambel oak, Rocky Mountain maple, redstem

ceanothus, and Martin ceanothus are often heavily

browsed and can contribute a considerable amount of

forage throughout the entire year. Green ephedra,

Utah juniper, and Oregon grape usually are grazed

during specific seasons.

Woody plants provide not only forage, but are equally

important as cover and concealment for wildlife. Utah

juniper, Gambel oak, skunkbush sumac, Rocky Mountain maple, and willows are very important for this

purpose. Many shrubs discussed in this chapter are

also useful for soil stabilization, watershed protection,

and ground cover. Deerbrush ceanothus, Scouler willow, mountain snowberry, thinleaf alder, and golden

currant are examples of species useful for conservation plantings. The native shrub communities provide

aesthetic and cultural resources that are unique to the

Western United States.

Important characteristics of a number of shrubs

are listed in table 1. Seeding recommendations for

major vegetative types and conditions are discussed

in chapter 17. Shrubs adapted to these situations are

included in the seeding recommendations. Seed characteristics are found in chapters 24 through 27.

Family Aceraceae _______________

Acer glabrum

Rocky Mountain maple

Description—Rocky Mountain maple is a longlived, shade-tolerant, deciduous shrub or small tree

native to North America (Van Dersal 1938). Plants

vary in height from 6 to 20 ft (1.8 to 6.1 m) (Harrington

1964), and also demonstrate considerable variation in

stature, leaf form, and fruit shape (Kartesz and Kartesz

1980). Some plants reach 20 to 30 ft (6.1 to 9.1 m) in

height with multiple trunks up to 1 ft (0.3 m) in

diameter, but they occur most often as shrubby plants

with multiple stems less than 15 ft (4.6 m) tall

(Haeussler and Coates 1986; Hitchcock and Cronquist

1973). Leaves are simple, opposite (Alexander and

others 1987), palmately three- to five-lobed or three

foliate, and mainly 0.8 to 3.2 inches (2 to 8 cm) wide

(Welsh and others 1987) (fig. 1). The herbage is glabrous.

Figure 1—Leaves of Rocky Mountain maple

are opposite and palmately five-lobed.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 601

Chapter 23 Shrubs of Other Families Table 1—Species characteristics of selected Intermountain shrubs.

Vegetative typesa

 to which

the species is adapted

Ease of cleaning seed

Ease of seeding

Ease of transplanting

Germination

Initial establishment

Final establishment

Persistence

Natural spread

Growth rate

Herbage yield

Availability of current growth

Palatability, early spring growth

Palatability, summer growth

Edible foliage retained fall and winter

Grazing tolerance

Seed production

Soil stability

Compatibility with other species

Shade tolerance

Flooding tolerance

Disease and insect resistance

Alder, thinleaf 3b 25445444543424355545 A,WM,PP,MB,R

Apache plume 1 33423432343445244234 MB,JP,BS,WS,BB

Ash, singleleaf 4 32223522342445244124 MB,JP,BS,WS

Barberry, creeping 3 44223442223354234434 A,PP,MB,JP,MS,BS

Bearberry 3 44213432242244254424 A,PP

Birch, bog 3 2 3 3 3 4 4 4 4 4 3 3 4 3 4 2 4 4 4 4 3 SA,A,PP,WM,R

Birch, paper 3 25444445432424344443 A,PP,WM,MB,R

Birch, water 3 25444445422424344443 A,PP,WM,MB,R

Bitterbrush, antelope 4 55545444443445545424 PP,MB,JP,MS,BS,BB

Bitterbrush, desert 4 53543333443345434224 MB,JP,BS,BB

Blackcap 3 25224554442324154543 A,WM,PP,MB

Blackbrush 3 52411431333435133125 BS,WS,BB

Buckthorn, cascara 3 54333434443434344444 A,PP,MB,MS

Buffaloberry, roundleaf 3 42223322333344143124 MB,JP,BS,WS,BB

Buffaloberry, russet 3 44224433343444243534 SA,A,PP

Buffaloberry, silver 3 44334544443434344434 WM,MB,JP,BS,WS,BS,IS,R

Ceanothus, deerbrush 5 55444445553435354334 A,PP,MB

Ceanothus, Fendler 5 53323333433445243214 A,PP,MB,PJ,MS

Ceanothus, Martin 2 44335543433435254424 A,MB,PJ,MS,BS

Ceanothus, prostrate 3 54433542223445253424 A,PP,MB,MS

Ceanothus, redstem 3 55445555553435344434 A,PP,MB,R

Ceanothus, snowbrush 3 55445555553445354434 A,PP,MB,MS

Ceanothus, wedgeleaf 3 55434444553445354324 A,PP,MB,MS

Cherry, Bessey 3 45444414344424444433 A,WM,PP,MB,JP, MS,BS

Cherry, bitter 3 54224533445435354324 A,PP,MB,JP,MS

Chokecherry, black 3 3 4 3 2 4 5 4 4 5 4 5 4 3 5 4 5 4 4 3 3 SA,A,PP,MB,JP,R,MS

Cinquefoil, bush 4 34224533344434244444 SA,A,WM,R

Cliffrose, Stansbury 2 43433443443455435124 PP,MB,JP,MS,BB

Currant, golden 3 55444544455524344443 WM,PP,MB,JP,MS,R,BS,R

Currant, sticky 3 34323323444434334434 SA,A,PP,MB

(con.)
602 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families Currant, wax 3 44224433344434234334 A,PP,MB,JP,BS Cypress, Arizona 2 43334413333544333234 MB,JP,MS,BS Dogwood, redosier 3 45334445444434345444 A,PP,WM,MB,JP,R Elaeagnus, autumn 3 55334534444434344434 PP,MB,MS,WM Elderberry, blue 3 4 4 1 2 5 5 3 5 5 4 5 5 2 5 3 4 5 3 3 5 A,PP,MB,JP,MS,BS,R Elder, box 3 25542234224414433244 A,MB,JP,MS Elderberry, red 3 44125535545525344445 SA,A,PP,MB Ephedra, green 5 52543422344555334115 MB,JP,MS,BS,WS,SS,BB Ephedra, Nevada 5 52543422344455234115 MB,JP,BS,WS,SS,BB Eriogonum, sulfur 3 5 3 4 4 5 3 5 4 3 3 4 4 4 4 3 3 3 3 2 3 PP,MB,JP,MS,BS,WS Eriogonum, Wyeth 3 53445444344444334424 SA,A,PP,MB,JP,MS,BS,WS Forestiera, New Mexico 2 44324524232534434224 MB,JP,BS,WS Greasewood, black 3 43414523333525334155 BS,WS,SS,BG,IS Hackberry, netleaf 3 44224532433434244334 WM,PP,MB,MS,R Hawthorn, Douglas 3 34124552342334234343 WM,PP,MB,MS,R Honeylocust 3 45334544332422333322 PP,MB,JP,MS Honeysuckle, bearberry 3 35214533445544244434 SA,A,WM,MB Honeysuckle, Utah 3 45424534444424244435 A,PP Honeysuckle, orange 3 45324544443344253324 A,PP,MB Hopsage, spineless 3 44442433334535343135 JP,BS,WS,SS,BB Hopsage, spiny 3 42412512225115233135 JP,BS,WS,SS,BB Indian apple 3 53333533445425343323 PP,MB,JP,MS,WS Juniper, creeping 5c 53114522242255242324 A,PP Juniper, Rocky Mountain 5c 53214452343255332334 A,MB,JP,MS,BS Juniper, Utah 5c 51115552232355214224 MB,JP,MS,BS,WS,SS,BB

Vegetative typesa

 to which

the species is adapted

Ease of cleaning seed

Ease of seeding

Ease of transplanting

Germination

Initial establishment

Final establishment

Persistence

Natural spread

Growth rate

Herbage yield

Availability of current growth

Palatability, early spring growth

Palatability, summer growth

Edible foliage retained fall and winter

Grazing tolerance

Seed production

Soil stability

Compatibility with other species

Shade tolerance

Flooding tolerance

Disease and insect resistance

Table 1 (Con.)

(con.)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 603

Chapter 23 Shrubs of Other Families

Vegetative typesa

 to which

the species is adapted

Ease of cleaning seed

Ease of seeding

Ease of transplanting

Germination

Initial establishment

Final establishment

Persistence

Natural spread

Growth rate

Herbage yield

Availability of current growth

Palatability, early spring growth

Palatability, summer growth

Edible foliage retained fall and winter

Grazing tolerance

Seed production

Soil stability

Compatibility with other species

Shade tolerance

Flooding tolerance

Disease and insect resistance

Table 1 (Con.)

Kochia, gray-molly 3 43324532224545244134 WS,SS,BG

Kochia, forage 3 44424543444545444224 MB,PJ,MS,BS,WS, SS,BB, C,AW

Locust, black 3 34344544332433333323 PP,MB,JP,MS

Maple, bigtooth 3 33422333445423444434 A,MB,JP

Maple, Rocky Mountain 3 22212333234423335533 A,PP,MB

Mockorange, Lewis 1 2 3 4 3 3 4 4 4 4 4 3 2 4 4 3 3 2 2 3 2 A,PP,WM,MB,MS

mountain ash, Greene’s 3 43323442333234322233 A,WM,PP,MB

 Mountain lover 2 3 2 2 2 2 4 3 3 4 3 4 2 3 4 2 2 4 2 4 4 SA,A,PP,WM,MB

Mountain mahogany,

 curlleaf 2 43433433333554333313 PP,MB,JP

Mountain mahogany,

 littleleaf 2 43432433343444342213 PP,MB,JP

Mountain mahogany, true 2 43443444344424343323A,PP,MB,JP,MS

Ninebark, mallowleaf 3 4 4 4 3 3 4 4 3 4 4 4 2 4 4 2 3 4 3 3 4 A,WM,PP,MB,MS

Oak, Gambel 5 21432423441525353434 A,MB,JP

Oceanspray, creambush 4 44334433342255233343 A,WM,PP,MB,MS,R

Peachbrush, desert 5 44544443335414333133 MB,JP,BS,WS,SS,BB

Penstemon, bush 4 55444433544433323431 A,PP,MB,MS

Plum, American 3 23324443334414343344WM,MB,R

Rabbitbrush, Douglas 4d 24334453333434334224 A,PP,MB,JP,MS, BS,WS,SS

Rabbitbrush, dwarf 4d 25444553234445354114MB,JP,MS,BS,WS,BB

Rabbitbrush, low 4d 24334544344453442344 A,MB,JP,MS,BS,WS, SS,BG,BB,IS

Rabbitbrush, Parry 4d 25434443534445344224 SA,A,MB,SS

Rabbitbrush, rubber 4d 24444455553355454223 SA,A,PP,MB,JP, MS,BS,WS,SS,

BG,IS

Rabbitbrush, small 4d 25444554333445343115 JP,MS,BS,WS,SS

Rabbitbrush, spreading 4d 25445444445344353234 BS,WS,SS,BG,BB

Rockspirea 4 4 3 2 3 4 4 2 3 4 4 2 3 5 4 2 4 3 2 4 3 SA,A,WM,PP,MS

Rose, Woods 3 45234534434425244444 SA,A,PP,MB,JP,MS

(con.)
604 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Vegetative typesa

 to which

the species is adapted

Ease of cleaning seed

Ease of seeding

Ease of transplanting

Germination

Initial establishment

Final establishment

Persistence

Natural spread

Growth rate

Herbage yield

Availability of current growth

Palatability, early spring growth

Palatability, summer growth

Edible foliage retained fall and winter

Grazing tolerance

Seed production

Soil stability

Compatibility with other species

Shade tolerance

Flooding tolerance

Disease and insect resistance

Table 1 (Con.)

(con.)

Sagebrush, basin big 4d 35545555553354443233 JP,BS,SS,BG,BB,C,AW

Sagebrush, Bigelow 4d 35544454343344333233 JP,BS,WS,SS,BG,BB

Sagebrush, black 4d 45545554333554443114 MB,JP,MS,BS,SS

Sagebrush, bud 4d 22213322225125224124 BB,WS,SS

Sagebrush, early 4d 33444443333345232144 MB,MS

Sagebrush, fringed 4d 34344443332544334444 MB,JP,MS,BS,SS

Sagebrush, low 4d 34545554332235232244 JP,BS,WS,SS,BG

Sagebrush, mountain big 4d 3 5 4 4 5 5 5 5 5 5 3 4 5 5 3 4 4 3 3 3 SA,A,WM,PP,MB

Sagebrush, pygmy 4d 22322222223334234234 WS,SS,BB

Sagebrush, silver 4d 4 5 5 4 5 5 5 4 5 5 3 4 5 5 4 4 3 3 4 4 SA,A,MB,MS,WS

Sagebrush, stiff 4d 23323332233435233224 MS,BS,WS

Sagebrush, threetip 4d 35545555553354444334 MB,MS,BS,WS,JP,MS

Sagebrush, Wyoming big 4d 35545444443354343223 JP,BS,WS,SS,BG,BB,C,AW

Saltbush, Castle Valley

 clover 4 43433434445543321343 WS,SS,BG

Saltbush, fourwing 4 54443435543555545122 MB,JP,MS,BS,WS,SS, BG,BB,IS

Saltbush, Gardner 5 44333343344545343133 MB,JP,BS,WS,SS,BG,IS

Saltbush, mat 5 44323432445553211343 WS,SS,BG,BB

Saltbush, quailbush 4 43223454445423431343 WS,SS,BG,BB

Saltbush, shadscale 4 42212332333535244123 JP,BS,WS,SS,BG,BB

Sandsage 4d 35445444333344344224 JP,BS,WA,BB

Serviceberry, Saskatoon 3 53333433344434344423 SA,A,PP,MB,JP,MS

Serviceberry, Utah 3 53334433344434344324 PP,MB,JP,MS,BS,BB

Silverberry 3 45334423342334434344 WM,PP,MB

Snowberry, common 3 5 5 3 3 4 3 3 4 4 4 4 4 2 4 2 4 3 4 4 4 MB

Snowberry, longflower 3 55233434445324144224 A,JP,BS,WS,BB

Snowberry, mountain 3 55334544445425255444 SA,A,PP,MB,JP,MS

Snowberry, western 3 55334434344424244434 WM,MB,BS

Spiraea, birchleaf 3 3 4 4 3 5 5 3 4 4 5 2 2 5 4 2 4 4 3 4 3 A,WM,PP,MB,MS

Sumac, Rocky Mountain

 smooth 2 55224434443434444224 PP,MB,JP,MS

Sumac, skunkbush 2 55224533443435343234 PP,MB,JP,MS,BS,BB
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 605

Chapter 23 Shrubs of Other Families Thimbleberry 3 25224553342324154543 A,WM,PP,MB Virginsbower, western 1 24323443334414244324 A,PP,MB,JP,MS,BS,BB Willow, Booth 2 24424455455354344445 A,WM,PP,R Willow, coyote 2 24424445553344534334 WM,A,PP,IS,R Willow, Drummond 2 2 4 4 2 4 4 5 5 4 5 5 3 5 4 3 2 4 4 4 5 SA,A,PP,WM,R Willow, purpleosier 2 25424425454444454444 A,WM,PP,MB,R Willow, Scouler 2 2 5 4 2 4 4 4 5 5 5 4 4 4 4 4 5 4 4 3 4 A, WM,PP,R Willow, whiplash 2 25424445454444344444 A,PP,P-J,WM,R Winterfat, common 3 25544445334445335124 MB,JP,BS,WS,SS,BG,BB,IS Wormwood, oldman 4d 35114235333534154215 SA,A,PP,MB,MS aVegetative type or community to which the species is adapted: SA = Subalpine; WS = Wyoming big sagebrush; A = Aspen-conifer; SS = Shadscale saltbush; WM = Wet and semiwet meadows; BG = Black greasewood; PP = Ponderosa pine; BB = Blackbrush; MB = Mountain brush; IS = Inland saltgrass; JP = Juniper-pinyon; R = Riparian; MS = Mountain big sagebrush; C = Cheatgrass; BS = Basin big sagebrush; AW = Annual weeds. bKey to ratings: l = Poor–difficult; 2 = Fair; 3 = Medium; 4 = Good; 5 = Excellent–easy. cIn the flesh. dLess than 15 percent purity.

Vegetative types

a

 to which

the species is adapted

Ease of cleaning seed

Ease of seeding

Ease of transplanting

Germination

Initial establishment

Final establishment

Persistence

Natural spread

Growth rate

Herbage yield

Availability of current growth

Palatability, early spring growth

Palatability, summer growth

Edible foliage retained fall and winter

Grazing tolerance

Seed production

Soil stability

Compatibility with other species

Shade tolerance

Flooding tolerance

Disease and insect resistance

Table 1 (Con.)
606 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 2—Rocky Mountain maple may occur as

individual shrubs, or more often in small thickets in

mountain brush communities.

Twigs are usually gray with smooth, reddish-brown

bark. Flowers are monoecious or dioecious, appearing

in loose, racemose corymbs (Harrington 1964). Fruits

consist of two fused samaras about 1 to 1.5 inches (2.5

to 3.8 cm) long, with widely divergent wings, each

containing a single seed (Harrington 1964; Hitchcock

and Cronquist 1973; Olson and Gabriel 1974).

Ecological Relationships and Distribution—

Rocky Mountain maple occurs from Alaska south to

California, Arizona, New Mexico, and Nebraska (Little

1979; Welsh and others 1987). In the Pacific Northwest, Rocky Mountain maple is primarily a coastal

species, but it is distributed eastward into Idaho,

Montana, and western Colorado at elevations between

5,000 and 10,500 ft (1,500 and 3,200 m) (Hitchcock and

Cronquist 1973; Harrington 1964). It occurs at elevations of 5,000 to 9,000 ft (1,500 to 2,700 m) in conifer

forests of the Sierra Nevada Mountains, northern

Coast Ranges, and into western Nevada (Munz and

Keck 1959). Welsh and others (1987) reported the plant

occurs in all Utah counties at elevations between

5,440 and 10,300 ft (1,700 and 3,100 m).

Rocky Mountain maple is frequently a seral species,

but it is able to persist as an understory in climax

conifer stands due to its shade tolerance (Mueggler

1965; Ryan and Noste 1985). It occurs intermixed with

Douglas-fir, lodgepole pine, ponderosa pine, sprucefir, and quaking aspen. However, it does not occur as

a common understory species with aspen in Utah

(Mueggler 1988). It occurs along streambanks in mountainous areas, in moist canyon bottoms, and on slopes

(Mozingo 1987; Wasser 1982), primarily in moist habitats (Lotan 1986). This species is frequently associated with mallowleaf ninebark, Saskatoon serviceberry, Scouler willow, Greene’s mountain ash,

chokecherry, mountain lover, and Utah honeysuckle

(Atzet 1979; Johnson and Simon 1987; Mauk and

Henderson 1984; Steele and others 1981; USDA Forest Service 1973; Wasser 1982).

Within the Intermountain region, Rocky Mountain

maple occurs primarily in mountain brush communities. The species exists over a wide range of sites from

well-drained to moist situations. Plants grow intermixed with Gambel oak, chokecherry, and Woods rose

in mesic, brushland sites. At lower elevations, its

range overlaps with pinyon and juniper, although it

usually does not grow in mixtures with these conifers.

Where pinyon or juniper occur with maple, the conifers are usually restricted to the more arid sites. In

arid regions it is restricted to streams and moist

canyons (Mozingo 1987; Wasser 1982), existing as an

important species in riparian communities (Brown

and others 1977; Mozingo 1987; Myers 1987). It occupies both warm and cool wet areas.

In some situations, Rocky Mountain maple appears

to be dependent on the presence of a well-developed

soil, with high concentrations of organic matter in the

upper portions of the soil profile (Krajina and others

1982; Mueggler 1965). Plants also occur on rocky and

gravelly soils and on steep slopes with shallow soils,

where a considerable amount of litter has accumulated. At both upper and lower elevation limits, the

distribution of Rocky Mountain maple is influenced by

topographic features and aspects.

Rocky Mountain maple usually grows on sites that

receive between 16 and 30 inches (400 and 760 mm) of

annual precipitation. Mature stands will usually persist with limited dieoff during periods of extended

drought. A decrease in herbage production may occur

if the precipitation level falls below normal, but stand

density usually remains unchanged.

Rocky Mountain maple may occur in dense, restricted patches or clumps; yet, open diverse stands

often occupy an entire slope or small drainage (fig. 2).

When found on south or west aspects, it may be

confined to small patches surrounded by bunchgrasses

or more xeric species. Mountain brome and slender

wheatgrass frequently occur as an understory with

this species. These two grasses may increase or decline

dramatically from year to year, but the density of

overstory maple remains quite consistent.

Grazing, wildfires, and other impacts have not seriously reduced the distribution or density of Rocky

Mountain maple. Cutting and burning temporarily

reduce some stands, but natural recovery is quite

rapid. Weedy species have not seriously affected the

presence or vigor of Rocky Mountain maple even

though heavy grazing has removed the native understory in some areas.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 607

Chapter 23 Shrubs of Other Families

Rocky Mountain maple is particularly important in

many of the conifer forest communities. It is not seriously

diminished by a closing canopy (Antos 1977; Habeck

1970; Stickney 1985). It can recover immediately following fire (Cholewa and Johnson 1983; Mueggler 1965) or

logging due to its resprouting capabilities. Recovery is

from stem resprouting, not by root suckering or rhizome

formation (Haeussler and Coates 1986).

Plant Culture—Seeds usually mature in August or

September (Olson and Gabriel 1974). Some stands

produce seeds once every 2 to 3 years (Olson and

Gabriel 1974). However, individual plants scattered

throughout a stand usually produce some seeds each

year. In addition, certain plants have been observed to

produce a viable crop of seeds every year for 3- to 5-

year periods. Individual plants often produce an extraordinary amount of seeds. Seeds also tend to ripen

more uniformly on certain plants; consequently, seed

collection is usually confined to a few plants. Seeds

often ripen over a period of 1 month.

Seeds usually remain on the plants for 2 to 4 weeks

before being dislodged by wind. When mature, the

fruits turn from green to yellow or reddish brown.

Seeds can be collected by hand stripping or flailing the

bush. Green or immature seed should not be collected.

Seeds should be inspected for insect damage and

immature embryos before collection. Some collections

exhibit about 50 percent germination (Swingle 1939).

This percentage can be improved, however, if care is

taken to collect mature, well-developed seeds. There

are about 13,430 seeds per lb (29,600 per kg) (Olson

and Gabriel 1974).

The wings are usually detached from the fruits to aid

in handling and planting. Once the wings are removed, empty seeds can be removed by flotation. This

is advisable to permit regulation of seeding rates.

Seeds should be dried to a water content of 10 to 15

percent before storage (Olson and Gabriel 1974). Seeds

can be stored for 1 to 2 years in sealed containers at 35

to 41 ∞F (1.7 to 5 ∞C) without serious loss of viability

(MacArthur and Fraser 1963). Seeds have dormant

embryos that require 6 months of wet prechilling to

release dormancy (Shaw 1984). Consequently, fall

seeding is advised. However, wet prechilled seeds can

be spring planted, particularly under nursery conditions. Seeds dispersed on the soil surface do not maintain viability (Kramer 1984).

Cleaned seeds can be planted with most conventional seeders, but most plantings have been restricted

to small, carefully prepared areas where hand seeding

has been employed. Seeds should be planted at a depth

of 0.5 to 1.5 inches (1.3 to 3.8 cm). Broadcast planting

without any followup methods for covering the seeds is

not advised. Planting weedy sites should be avoided.

Seedlings have grown well and attained considerable

size in 1 to 2 years.

To date, most seedings have been confined to sites

where the species previously existed. When seeded

“offsite,” weak and erratic stands have developed.

Plants appear to be quite dependent on soil conditions

to survive and flourish. When seeded or transplanted

on mine wastes, plants have done poorly. However, if

sites are top soiled, better stands have established and

persisted.

Natural seedlings are usually not abundant. When

found, they are encountered in shady areas with little

competition (Steele and Geier-Hayes 1992). Under

these conditions seedlings have been reported to grow

slowly.

Rocky Mountain maple is easily cultured and has

been planted in commercial landscape programs.

Transplants or wildlings are used in conjunction with

other species in formal plantings.

Transplanting 1- to 3-year-old nursery stock has

improved stand establishment in areas with poor

soils or where some understory competition exists.

Older transplants are able to establish and begin

growth quite rapidly without a prolonged period of

adjustment.

Uses and Management—Rocky Mountain maple

has been planted to provide diversity and enhance

wildlife habitat. Though it is often confined to small

areas, it provides valuable thickets for protection and

concealment. It is a useful forage species and occurs in

important winter habitats for big game animals (Arno

and Hammerly 1977; Beetle 1962; Gaffney 1941; Keay

1977; Martinka 1976; Young and Robinette 1939).

Rocky Mountain maple normally supports a productive complex of understory herbs. These sites are

important areas for summer grazing by big game and

livestock. Stands of Rocky Mountain maple that recover following burning provide considerable forage

for big game, livestock, and nongame animals. In most

situations plants may grow too tall to be reached by

animals, yet they provide excellent cover (Johnson

and Simon 1987; Larson and Moir 1987). In more arid

situations, the shorter growing forms remain available to browsing animals. Rocky Mountain maple is a

palatable species, particularly to big game (Beetle

1962; Keay 1977; Martinka 1976; Mitchell 1950; Smith

1953), but is also used by other animals, including

ruffed and blue grouse (Steele and others 1981) and

small mammals (Martin and others 1951).

Rocky Mountain maple grows in moist areas. When

disturbed, these sites can produce sediment and increase runoff. These sites are important for watershed

stability and may require immediate attention if disturbed. Rocky Mountain maple is able to recover

following disturbances and can provide soil and

streambank protection.

Both Rocky Mountain maple and bigtooth maple are

attractive plants, particularly during the fall when
608 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

leaves turn red and yellow. Certain plants and clumps

exhibit early fall coloration, others are particularly

brilliant in color, and some may retain their leaves for

extended periods. Also, considerable differences in

growth habit and mature size occur among ecotypes.

Many native stands have high aesthetic appeal and

are important viewing areas and recreational sites.

These areas are frequently managed for scenic use.

Rocky Mountain maple has not been used as extensively in windbreaks and conservation plantings as

other maples.

Varieties and Ecotypes—None.

Family Anacardiaceae____________

Rhus aromatica

Skunkbush sumac

Description—Skunkbush sumac is an unusually

persistent and highly variable species (fig. 3). Welsh

and others (1987) described two varieties of this species complex that occur in Utah. Rhus aromatica var.

simplicifolia has simple leaves and occupies xeric

sites in southeastern and southwestern Utah. Rhus

aromatica var. trilobata leaves are trifoliolate, and

plants grow on mesic sites including streambanks,

seeps, and riparian communities. Shrubs occur singly

or in dense patches and may be connected by woody

rhizomes, which often exceed 20 ft (6.1 m) in length

(Sanford 1970). Root systems are deep and extensively

branched. Plants are deciduous, woody, spreading,

and moundlike to erect, often forming clumps or

thickets. They normally range from 2 to 7 ft (0.6 to

2.1 m) in height (Hitchcock and others 1961; Welsh

and others 1987); some may reach heights of 15 ft

(4.6 m) (Thornburg 1982). Crown diameter often exceeds the height and may be more than 30 ft (9.1 m)

(Plummer and others 1968; USDA Forest Service

1976c). Stems are numerous, woody, spreading, highly

branched, and brown hairy when young; they develop

a gray bloom with age (Barkley 1937; Wasser 1982).

Leaves are 1 to 3 inches (3 to 7 cm) long, trifoliate,

petiolate, and puberulent beneath. The terminal leaflet is fan shaped, shallowly to deeply incised or lobed,

and coarsely crenate. Lower leaflets are half as large

and shallowly crenate. The resinous, acrid sap of the

twigs and leaves is nonpoisonous, but produces a

disagreeable odor, hence the common name, skunkbush

sumac (Stubbendieck and others 1986). Plants are

polygamo-dioecious. The inflorescence consists of a

cluster of spikes located near the branch apex (Goodrich

and Neese 1986). Buds and flowers are inconspicuous,

pale yellow, and mostly imperfect. Fruits are orange

red, sticky, berrylike drupes, each containing a single

bony seed (Brinkman 1974g; Stubbendieck and others

1986; Welsh and others 1987).

Ecological Relationships and Distribution—

Skunkbush sumac is widespread and highly variable; it

ranges from Alberta south to Iowa and Mexico, and

west to southeastern Oregon and California (Harrington

1964). It is found at elevations from 3,500 to 8,000 ft

(1,100 to 2,400 m) (Hitchcock and others 1961; USDA

Forest Service 1937) on sites with annual precipitation between 10 and 20 inches (250 and 510 mm)

(Wasser 1982). It grows in the mountain brush and

pinyon-juniper types of the Intermountain region and

the plains and foothills of eastern Wyoming and Montana (Welsh and others 1987). It is a primary or

secondary species growing in sand hills of the plains

and Southwest (Wasser 1982). At lower elevations it

may be found with blackbrush, basin big sagebrush, or

Wyoming big sagebrush. It also occurs in openings in

oakbrush and ponderosa pine forests (Vories 1981).

Plants growing in rocky foothills are usually smaller

in stature and less productive than the more typical

upright forms of the mountain brush communities

(Welsh and others 1987). Skunkbush sumac is generally drought and cold tolerant, but these characteristics vary among ecotypes.

Skunkbush sumac is commonly found along stream

courses where it occurs intermixed with other shrubs

or as an understory with cottonwoods. Although dense

stands are usually not encountered, scattered plants

or thickets may be widely distributed in openings

along riparian areas; plants require full sunlight to

partial shade. On dry sites it occurs in swales and

drainageways where it may receive extra moisture. It

requires well-drained soils with deep water tables

(Wasser 1982), and although it tolerates short-term

Figure 3—Skunkbush sumac established from a

direct seeding provides important wildlife habitat.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 609

Chapter 23 Shrubs of Other Families

flooding where it occurs along streams, it is not capable of surviving long-term flooding.

Skunkbush sumac occurs on a wide range of soils

ranging from sandy textured soils to heavy clay-textured materials (Johnson and others 1966). In Utah, it

is particularly well adapted to heavy shale outcrops,

and sometimes salty outcrops, where few other shrubs

are adapted (Plummer 1977). It grows on both acidic

and basic soils including slightly alkaline soils (Johnson

and others 1966; Vories 1981). It occurs on deep fertile

soils and also on shallow, rocky, less-developed soils.

Plant Culture—Fruits of skunkbush sumac form

dense clusters near the ends of small branches. The

clusters are scattered over the surface of the entire

bush. Flowers appear from March to April (Brinkman

1974g). Fruits ripen from June to September (Plummer

and others 1968; Van Dersal 1938), and many remain

on the shrub until midwinter. Most seeds are removed

by birds or other animals (Brinkman 1974g).

Fruits may be harvested over a period of 2 to 3

months during early fall and winter, but they are more

difficult to clean if they have been allowed to dry on

the shrub. Fruits are collected by hand picking or

flailing the stems after leaves have fallen in early

winter. Fruits adhere to the plant and are not easily

dislodged. This characteristic interferes with seed

collection and results in high seed costs. Harvested

fruits are macerated using a Dybvig cleaner and

flushed with water to remove the pulp. The remaining

material, including the seeds, is then dried and fanned

to remove small debris. Seeds can be cleaned to purities exceeding 90 percent. There are approximately

20,000 cleaned seeds per lb (44,100 per kg) (Brinkman

1974g; Plummer and others 1968; USDA Forest Service 1948). Recommended laboratory standards for

seed purchases are 40 percent germination and 95

percent purity. Cleaned seeds remain viable for up to

5 years under open, dry storage.

Seed production varies among sites and ecotypes.

Most upland populations produce some seeds each

year, but abundant crops must be produced to justify

commercial harvesting. Seeds from many useful

ecotypes or collection sites normally are not harvested

or sold because production is quite low. Consequently,

poor seed-producing ecotypes are not collected and

used in restoration programs. Seeds from more limited, but productive, stands are normally harvested

and sold for extensive plantings. Thus, the most adapted

ecotypes are not always used.

Seed germination may be restricted by the presence

of a hard, impervious seedcoat and embryo dormancy.

Both forms of dormancy vary widely among seedlots.

Seedcoat permeability may be increased by a 20-

minute to 2-hour sulphuric acid scarification (Babb

1959; Brinkman 1974g; Glazebrook 1941; Heit 1968a).

A wet prechilling for 30 to 120 days is required to

release embryo dormancy (Babb 1959; Heit 1970;

Swingle 1939). Duration of acid treatment and wet

prechilling periods required vary among seedlots.

Physical rupturing of the seedcoat using mechanical

methods also increases germination (McKeever 1938),

but procedures have not been developed to treat large

seedlots. Fall seeding often provides the necessary

cold treatment to release embryo dormancy (Brinkman

1974g), but very erratic results have occurred, possibly due to differences in seedcoat imposed dormancy

(Boyd 1943; Monsen 1987; Monsen and McArthur

1985).

Erratic seed germination is the principal factor

preventing extensive seeding of this shrub. Differences in germinability are not entirely related to

ecotypes or site of collection. Plants from sites in

north-central Wyoming and south-central Montana

are in considerable demand for mine restoration, but

these ecotypes are difficult to germinate. Means to

stimulate or assure field germination are not well

understood, and until these problems are corrected,

use of skunkbush sumac will be adversely affected.

Seeds that have been properly pretreated will normally germinate in 30 to 45 days, once germination

begins (Monsen and Christensen 1975). Germination

is usually completed in 30 days under nursery conditions (Brinkman 1974g). This long germination period can adversely affect survival of wildland

plantings. Soil at many sites does not remain moist

long enough to ensure germination and seedling

establishment.

Different ecotypes of skunkbush sumac occur

throughout its range. Adapted seed sources must be

selected for range seedings or transplanting projects.

Scarified seed should be fall planted to provide overwinter wet prechilling; scarified seeds sown in the

spring must first be wet prechilled. Seed should be

planted approximately 0.5 inch (1.3 cm) deep or slightly

deeper in dry, coarse-textured soils. Skunkbush sumac

may be seeded at the rate of 2 to 4 lb per acre (2.2 to 4.5

kg per ha), depending on row spacings. It may be

broadcast on rough surfaces or in pits. If drill seeded,

it should be planted with other slow-growing shrub

species in rows separate from faster growing species.

Seedlings grow moderately well, but young plants

are not highly competitive with herbs (Monsen 1987).

Consequently, this species should not be seeded together with grasses. Young plants can be stunted by

herbaceous competition and may not recover (Monsen

1987). Established seedlings are quite hardy and resilient. Range or wildland seedings normally require 2 to

5 years to fully establish.

In the bareroot nursery seed should be planted

about 0.5 inch (1.3 cm) deep. A seeding density of 25

viable seeds per linear ft (82 viable seeds per linear m)

was recommended by Brinkman (1974g). At the Los
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Chapter 23 Shrubs of Other Families

Lunas Plant Material Center seeds are planted at a

density of 182 per linear ft (600 per linear m) of row to

yield 23 usable plants per linear ft (75 per linear m)

(Frazier 1979). Beds should be mulched to prevent

excessive drying. Plants may be lifted as 1-0 or 2-0

stock, depending on their growth rate. Field transplanted seedlings should be at least 8 to 11 inches (20.3

to 28 cm) tall. Container stock is most easily established from germinants and requires a 4- to 6-month

cropping time (Landis and Simonich 1984). Ferguson

and Frischknecht (1981) and Stevens (1981) reported

that large, well-rooted stock is easily transplanted.

Skunkbush sumac is grown by many commercial nurseries, and wildland plantings of bareroot stock have

been highly successful.

Doran (1957) reported that cuttings of Rhus root

well when taken in July and potted in a sand-peat

media. This provides a means of propagating collections from specific locations that are difficult to rear

from seed.

Field cultivation of skunkbush sumac has been successful in improving seed production. Producing seed

from agricultural fields or nursery sites could undoubtedly provide a means of marketing desired

ecotypes and providing a more stable market.

Uses and Management—Skunkbush sumac has

been extensively used in windbreaks, shelterbelts,

and conservation plantings (Brinkman 1974g; Hassell

1982; Swenson 1957). As an ornamental, it grows

rapidly and develops dense foliage (Johnson 1963;

Olson and Nagle 1965; Steger and Beck 1973). The

leaves and berries turn brilliant shades of red and

orange in the autumn. Mature plants withstand

burial by drifting sand and sprouting from the crown.

Plants are also able to survive when soil is eroded from

the upper portions of the root system. These characteristics, along with its cover value, low palatability,

ability to resprout after fire, longevity, and ability to

withstand climatic extremes, qualify sumac as a valuable species for revegetation (Johnson 1963; Olson

and Nagle 1965; Swenson 1957).

Skunkbush sumac requires little maintenance in

conservation plantings. Plants do not become decadent with age, but remain healthy and robust. Many

other shelterbelt species often decline in vigor or are

subject to crown damage from wind, insect attack, or

snowfall. Skunkbush sumac is generally not adversely

affected by these agents, but persists and becomes a

major species in older plantings.

Skunkbush sumac has been widely used in recreation sites, parkways, campgrounds, and roadway rest

areas. It frequently recovers by sprouting following

disturbances, and can be managed to assure the

presence of native species in recreational sites. It withstands traffic and campground activity; consequently,

it is used to screen landscape trails, roadways, and

structures. The plant is particularly important for

recreational sites in semiarid areas in the Intermountain region and the Southwest.

Selected ecotypes of skunkbush sumac have proven

well adapted to unstable disturbances. It has been

used to stabilize sand dunes (Kozlowski 1972), roadways, and mine sites (Hassell 1982; Monsen and

Christensen 1975). However, direct seedings and transplants establish and grow at only moderate rates for

the first few years. Although survival is often high,

plants do not provide extensive ground cover during

the establishment period. Once well established, annual growth increases, and the plants begin to provide

considerable ground cover and soil protection.

Skunkbush sumac can be used to stabilize disturbed

watersheds, including riparian sites. Its growth habit

provides excellent ground cover, and plants are able to

resist soil deposition. It is particularly useful for restoration of meadows and other riparian sites where

erosion and gully cutting have drained wetlands and

lowered water tables.

The spreading moundlike growth form of some

ecotypes and the ability of the shrub to form dense

thickets by root sprouting cause skunkbush sumac to

be a valuable cover plant for birds (Adkins 1980). In

addition, the fruits of skunkbush sumac are consumed

by many bird species (Brinkman 1974g), particularly

during winter months, as berries remain on the bush

(Swenson 1957). Consequently, the shrub has been

selectively planted in windbreaks and associated conservation plantings to provide forage and cover for

birds.

Skunkbush sumac has been most widely planted in

the Intermountain area for big game habitat. It is

often planted in mixtures with other more preferred

forage plants. The shrub is selectively planted on less

fertile sites in big sagebrush, pinyon-juniper, and

mountain brush communities where ground cover and

big game habitat are required (Plummer and others

1968). It grows slowly, normally requiring 10 to 20

years to attain mature stature (Monsen 1987), but it is

one of the most successful shrubs established from

rehabilitation plantings. Initial establishment and

ratings of young plantings are regarded as only fair,

but long-term performance ratings are good to excellent. Swenson (1957) and Adkins (1980) also reported

good to excellent survival for 20-year-old cover and

windbreak plantings of skunkbush sumac in eastern

Colorado and eastern Washington.

Palatability of skunkbush sumac is rated low for

livestock and fair to good for big game, depending on

the ecotype and other browse species available (Dayton 1931; Stanton 1974). It has been planted in mixtures for range and wildlife habitat to provide forage

and cover. In Montana and Wyoming it is considered

a preferred browse species and receives heavy winter
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Chapter 23 Shrubs of Other Families

use (Mueggler and Stewart 1980; Tweit and Houston

1980) and moderate use in the summer (Sanford

1970). Throughout other portions of its range, the

plant is only lightly grazed (Sanford 1970). Sheep

often browse the shrub at all seasons. Skunkbush

sumac is a particularly important forage plant in

midwinter during periods of deep snow cover. It occupies windblown slopes that often remain open and

available for browsing. Skunkbush sumac recovers

well from wildfires, sprouting vigorously from underground rhizomes (Dwyer and Pieper 1967; Wright and

others 1979). It is an important shrub in areas where

fires can be used to enhance vegetative conditions.

Varieties and Ecotypes—“Bighorn” skunkbush

sumac, developed by the Los Lunas Plant Materials

Center, is a tall, upright variety selected from trials

conducted at locations from New Mexico to Montana

and North Dakota (Thornburg 1982). Seed of this

variety does not exhibit embryo dormancy and requires only a 10 to 25 minute acid scarification (Frazier

1979). Populations of skunkbush sumac vary widely in

size, growth habit, leaf characteristics, and range of

adaptation; they provide many opportunities for future selection work.

Family Anacardiaceae____________

Rhus glabra

Rocky Mountain smooth sumac

Description—Rocky Mountain smooth sumac is a

small, little-branched, thicket-forming shrub (Van

Dersal 1938). Mature plants usually do not exceed 3 to

6 ft (0.9 to 1.8 m) in height. Root suckers develop from

the parental plant, and new shoots may appear 10 to

13 ft (3.0 to 4.0 m) from the base of the shrub (fig. 4).

Branchlets are glabrous and glaucous. The leaves are

mostly 4 to 12 inches (10 to 30 cm) long with 7 to 21

leaflets. Leaves are lanceolate, mainly 0.6 to 3 inches

(1.5 to 8 cm) long, acuminate, serrate, and range in

color from dark green to light green. The inflorescence

consists of a panicle about 4 to 8 inches (10 to 20 cm)

long, containing many white flowers. The fruit is a

globular, red, thinly fleshy drupe. It is about 0.2 inch

(4 mm) long and covered with a dense layer of short red

hairs (Harrington 1964; Welsh and others 1987).

Ecological Relationships and Distribution—

Rocky Mountain smooth sumac is widespread in North

America (Welsh and others 1987). It occurs from

British Columbia to California, Oregon, and Arizona,

and eastward to the Atlantic Coast (Munz and Keck

1959). It is common from Nevada and Mexico to New

Hampshire and Georgia (Davis 1952; Hitchcock and

Cronquist 1973). In Utah it occurs on a variety of sites

including desert shrub, riparian, pinyon-juniper, and

mountain brush communities at altitudes between

3,590 and 7,710 ft (1,100 and 2,350 m) (Welsh and

others 1987). Harrington (1964) reported that the

shrub occupies valleys and slopes throughout the

western portion of Colorado at elevations between

5,500 and 7,500 ft (1,700 and 2,300 m).

Rocky Mountain smooth sumac often exists in disjunct stands in dry places, usually in open areas

without overstory shade. Large patches or clumps

often occur alone without other shrubs, but intermingled in groupings with other plants. It grows on a

variety of soils including sandy as well as heavytextured materials. It is a climax species with various

bunchgrasses in the Northwest, although the native

grasses have often been replaced by annual cheatgrass

(Daubenmire 1970; Franklin and Dyrness 1973). This

species grows along streams and canyon bottoms aligning riparian communities, but does not occur in wet

areas. It occupies steep slopes and streambanks intermixed with numerous upright trees and shrubs, but

not as an understory in dense patches of vegetation. It

is extremely drought tolerant, and is encountered on

open south- and west-facing slopes with big sagebrush, antelope bitterbrush, and mountain brush species. Rocky Mountain smooth sumac also forms thickets in the prairie grasslands (Weaver and Fitzpatrick

1934).

Plant Culture—Rocky Mountain smooth sumac

normally produces an abundant seed crop. If grown on

favorable sites, plants usually produce heavy seed

crops each year. Even under arid situations, plants

produce some viable seeds. Plants flower from June to

August, and are not subjected to late spring frosts that

Figure 4—Rocky Mountain smooth sumac produces distinctive clusters of red drupes.
612 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

damage seed crops of many shrubs. The flowers are

small and borne in terminal or axillary clusters

(Brinkman 1974g). Initially, the developing fruits do

not appear to be very abundant, but as fruits ripen in

fall or early winter, large, dense clusters of red fruits

develop (Lovell 1964; Plummer and others 1968;

Radford and others 1964; Van Dersal 1938). The fruits

are easily collected by handpicking or beating the

shrub to dislodge them. Fruits are usually collected in

fall after the leaves have dropped and the fruits are

more visible. Fruits can persist on the bush until late

winter without loss of seed viability. Consequently,

seed collection may be delayed until late winter.

Once the fruits are harvested they should be cleaned

to remove the fruit coat and dry pulp. The fruit is a

small drupe having a single bony nutlet (Brinkman

1974g). The fruit is normally dry at the time of

harvest, and the flesh is often left on the seed; however, the dry fruit makes planting more difficult,

particularly when mixed with other seeds. When

cleaned, the seeds are more easily stored or seeded.

Seeds are cleaned using a macerator to grind away

the pulp. Water is used to float and separate the pulp

from the seeds (Brinkman 1974g; Swingle 1939). If the

fruits are dry, soaking with water is often required

before maceration. High yields of good seed are normally obtained from most seed collections. Cleaned

seeds consist of from 45 to 75 percent of the fruit or

material initially collected (Brinkman 1974g). Seed

numbers range between 23,000 and 48,000 per lb

(50,700 and 105,800 per kg) (Johnson and others 1966;

Lovell 1964; McKeever 1938; Plummer and others

1968; Swingle 1939).

Seed germination may be erratic (Monsen and

Christensen 1975), and inhibited by a hard impervious seedcoat (Heit 1967a; Johnson and others 1966)

and a dormant embryo. Pretreatment using sulfuric

acid or hot water to increase seedcoat permeability

has been employed (Johnson and others 1966), but

mixed results have occurred due to differences in

structure of the seedcoat (Brinkman 1974g; Heit 1967a;

Lovell 1964). Seeds can be wet prechilled at 40 ∞F

(4.4 ∞C) for 2 months (Brinkman 1974g) to relieve

embryo dormancy.

Seeds should be planted in late fall or early winter at

a depth of 0.75 to 1.5 inches (2 to 4 cm). Some seedlots

germinate much better than others, particularly if

pretreated. Attempting to predetermine planting

success from new seedlots is difficult. Nursery seedings

are often heavily sown to assure establishment of

satisfactory stands.

Success from seeding wildland sites has been extremely erratic due to poor or very erratic germination.

Excellent quality transplant stock can be grown as

bareroot or container stock. One-year-old bareroot

stock is easily cultured and field planted with good

success. Consequently, harsh sites can be successfully

planted with transplant materials. Rooted suckers

can be field dug and transplanted with good results if

lifted when dormant. Doran (1957) recommended root

cuttings be taken in December.

Uses and Management—Rocky Mountain smooth

sumac is a highly useful shrub. It is important for

wildlife habitat, ground cover, and conservation

plantings. However, its usefulness in large-scale restoration plantings has been limited by low or erratic

seed germination.

Plantings of Rocky Mountain smooth sumac on wildland sites have been enhanced by seeding the shrub

alone or with a few other species in well-prepared

seedbeds. Planting in late fall in areas free of competitive herbs has improved stand establishment. Once

established, small seedlings and young plants are very

hardy and survive periods of drought. New plantings

are not damaged by moderate grazing, insects, or

disease, and are able to develop normally under most

circumstances.

Seedling survival can be quite erratic. Some small

seedlings are quite vigorous and grow at moderate rates,

but natural thinning usually results in the loss of a

number of young plants. Plummer and others (1968)

ranked this shrub as low to fair for the development of

mature stands from direct seedings. Seedling losses

usually result from erratic seed germination; many

seedlings emerge in late spring when soil water is rapidly

being depleted. Such seedlings usually do not survive.

Plants form a single main stem during the first and

second growing seasons. During this time an extensive root system is developed. Plants that survive the

first growing season are quite persistent, and few

plants succumb after that date. Plants reach maturity

quickly; within 6 years the shrub is sufficiently developed to produce flowers and seed. Spread by underground rooting usually begins within 10 years after

planting. Thereafter, root suckering may occur each

year, but the pattern is somewhat erratic. Numerous

suckers may appear in some years, but not in others.

The size and area occupied by a clump of plants tend

to stabilize within 20 to 30 years. Natural spread

may continue after this time, but maximum spread is

usually attained within this time span. Natural spread

is not always restricted by the presence of other species. New shoots often emerge in areas occupied by

closed stands of native bunchgrasses, big sagebrush,

and antelope bitterbrush.

Rocky Mountain smooth sumac is an important

forage plant for wildlife. The fruits are eaten by a large

number of birds (Miller and others 1948; Stanton

1974). It is moderately browsed by big game in the

spring, fall, and winter months, but is more heavily

grazed in the summer (Hill and Harris 1943; Kufeld

and others 1973; Nemanic 1942; Plummer and others
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Chapter 23 Shrubs of Other Families

1968; Smith 1953; Snyder 1937). When this shrub

occurs on big game wintering ranges and is available,

it is heavily browsed when access to other species is

restricted by deep snow. Some populations are more

heavily grazed each year than others.

Rocky Mountain smooth sumac is relatively easy to

establish by transplanting on harsh, unstable sites

(Plummer and others 1968). Consequently, it has been

successfully used to plant mine disturbances, roadways, and unstable watersheds. Although it establishes well and spreads to provide soil stability, erect

open stands usually develop on semiarid sites. Plants

growing on more mesic sites develop dense protective

ground cover. Under most conditions, this species can

be used to provide soil protection, particularly if understory herbs are also planted.

Rocky Mountain smooth sumac is a useful shrub to

plant on unstable slopes, streambanks, and roadfills

where mass movement may occur. Plants naturally

spread by roots to furnish excellent site stability.

Unstable soils can be quickly stabilized by transplanting large stock (2-year-old plants) 2 to 3 ft (0.6 to 0.9 m)

apart. The plants spread rapidly and fully occupy the

area in 1 to 3 years.

Rocky Mountain smooth sumac has been used to

revegetate mine disturbances, as it will grow in soils

with low fertility. It does not establish or compete very

well if sites are heavily seeded to herbs and fertilized

at high rates.

This shrub is widely used as an ornamental because

of its low growth form, attractive fruits, and fall

coloration. It is used to stabilize and beautify steep

banks, roadways, parks, and recreation sites. It has

been cultivated since 1620 (Brinkman 1974g). Rocky

Mountain smooth sumac is also useful in windbreak

and conservation plantings. It withstands extreme

cold temperatures, and can be used on exposed, windblown sites to protect soils and entrap snow.

Rocky Mountain smooth sumac survives wildfires,

and is an important plant where burning may create

erosion and watershed problems. This species can

recover quickly from fire, and provides needed cover

the first year after burning. Along with Gambel oak,

maples, Saskatoon serviceberry, and chokecherry,

the species readily resprouts and protects steep

watersheds.

Varieties and Ecotypes—None.

Family Berberidaceae ____________

Mahonia aquifolium

Shining barberry

Description—Mature plants are 0.3 to 6.6 ft (0.1 to

2.0 m) or more tall. Leaflets are more than twice as

long as broad, glossy on the upper surface and dull

beneath (fig. 5), with more pronounced spiny tips than

Oregon grape. The species frequents moist, rich humus soils on the floor of conifer forests, but sometimes

grows on exposed rocky slopes, brushy hillsides, and in

aspen or alder stands. Marchant and Sherlock (1984)

report that it is rather slow growing during the first

season and slow to establish on wildland sites. The

species integrates with Oregon grape in northeast and

north-central Washington, thus the two taxa may be

varieties of a single species.

Distribution—Shining barberry is distributed from

British Columbia and northern Washington to northeastern Idaho, and south from the eastern edge of the

Cascade Mountains to the coast, as far as the southern

Willamette Valley.

Plant Culture—Shining barberry has not been

extensively used in wildland plantings. Transplants

have been grown for small plantings, but direct seedings

are rare. Plants normally produce some fruits or berries each year, and seeds can be collected and processed as described for Oregon grape. Seeds are difficult to germinate uniformly, and information is needed

to properly culture this species.

Uses and Management—Shining barberry is infrequently used for low-maintenance landscaping,

erosion control, and conservation plantings. It is grown

by some commercial nurseries, but is not commonly

used in most wildland restoration projects.

Varieties and Ecotypes—None.

Figure 5—Glossy, elongate, spine-tipped leaves

characterize shining barberry.
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Chapter 23 Shrubs of Other Families

Figure 7—Oregon grape, a perennial creeping shrub,

occurs on well-developed soils as an understory in

conifer forests.

Family Berberidaceae ____________

Mahonia fremontii

Fremont barberry

Description—Fremont barberry is a large shrub of

dry slopes and ridges in pinyon-juniper types of the

Southwest. It grows 3 to 12 ft (0.9 to 4 m) tall and

produces small, compound, spiny leaves and dark blue

or red berries. This shrub has yellow wood (fig. 6).

Ecological Relationships and Distribution—

Fremont barberry is distributed across the Southwest

from western Texas through Sonora to Baja California

and north, and from California through southern

Utah and Colorado. It grows on deep sandy soils in

valley bottoms and draws. It is also encountered amid

rocky outcrops and steep canyon slopes. A number of

individual bushes often grow together in small

patches, although this shrub does exist interspaced

with other woody species. It is found in the warmer

drier regions of southern Utah and northern Arizona.

It grows intermixed with Utah juniper, Stansbury

cliffrose, blackbrush, and Nevada ephedra. It is common in draws and washes that may receive infrequent

runoff.

Plant Culture—Berries are generally not produced

each year. Special climatic conditions must occur to

promote flowering and seed production. However,

bushes planted in more mesic situations in central

Utah have consistently produced abundant crops

most years.

Seeds are formed in a berry that dries at the time of

maturation. Seeds are large and easily extracted and

separated from the dry fruit. A hard seed coat restricts

exchange of water and air. In addition, most seeds

require some wet prechilling to germinate.

Both container and bareroot nursery stock have

been grown for restoration plantings. Plants have been

reared and used in cultivated gardens and nursery

sites. With some irrigation and culture, plants attain

large stature, provide distinct gray-green foliage, and

form an attractive display of red fruits.

Uses and Management—This shrub has been

used in some recreation plantings, but has potential to

restore disturbances in the southwest shrub complex.

Varieties and Ecotypes—None.

Family Berberidaceae ____________

Mahonia repens

Oregon grape, creeping barberry,

mountain holly, creeping mahonia

Description—Oregon grape is a perennial creeping shrub with evergreen hollylike leaves (fig. 7). It

seldom grows taller than 1 ft (30 cm). The shrub

spreads by stolons, rhizomes, and stem layering; individual plants sometimes attain a diameter of 4 to 6 ft

(1.8 to 3.3 m). Leaves are evergreen, leathery, and

pinnately compound with five to seven (rarely three

or nine) leaflets that are less than twice as long as

broad. Leaflets are spinulose, glossy to dull on the

upper surface, and dull to glaucous on the lower

surface. Perfect, regular flowers develop in fascicled

bracteate racemes in the leaf axils. The perianth

consists of five alternate whorls of three members

each; all are yellow. Those in the outer whorl are tiny

bracts; the next two whorls are the sepals, and the

Figure 6—Fremont barberry produces attractive red

fruit and small, leathery evergreen leaves. It grows in

a variety of southwestern shrub communities.
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Chapter 23 Shrubs of Other Families

inner whorls are the petals. Petals are bilobed with

two glands at the base. There are six stamens and a

one-celled ovary. Fruit is a one- to several-seeded,

deep blue, glaucous berry about 0.3 inch (8 mm) in

diameter (Harrington 1964; Welsh and others 1987).

Oregon grape growth is intermittent in winter and

more active in spring (Wasser 1982). Flowering occurs

from March to July. Fruits ripen from September to

October and may remain on the plant into winter. Seeds

are dispersed by birds and mammals (Rudolf 1974).

Ecological Relationships and Distribution—

Oregon grape ranges from British Columbia to Alberta

and North Dakota and southward on the east side of

the Cascade and Sierra Mountains through the Great

Basin, Arizona, and New Mexico. It grows with junipers, pinyon, aspen, mountain brush, lodgepole pine,

spruce-fir, and ponderosa pine at elevations ranging

from near sea level to 10,000 ft (3,000 m) (Harrington

1964; Hitchcock and others 1964; Wasser 1982; Welsh

and others 1987). It is abundant in coniferous forests

receiving more than 15 inches (38 cm) of annual

precipitation, and is fairly common on north-facing

slopes and other moist sites in plains, foothill

shrublands, and low-elevation woodlands. Oregon

grape tolerates full sun and partial to deep shade of

shrubland and coniferous forests. It requires welldrained, more-or-less neutral (pH 5.5 to 7.0) soils, and

is only weakly tolerant of salinity (Stark 1966). Oregon grape is intolerant of poor drainage and high

water tables. Although it grows in deep, mediumtextured soils, it is also common on shallow, rocky sites

(Bailey 1949; Harrington 1964; Plummer and others

1968; Wasser 1982).

Plant Culture—Good fruit crops occur erratically.

Fruits are most often collected by hand. Berries of

more upright populations can be beaten into containers (Rudolf 1974). Pulp is removed by macerating

the fruit in a Dybvig cleaner. The seeds and pulp are

dried, lightly chopped, and separated by screening.

Seed may be stored dry in air-sealed containers at 34 to

37 ∞F (1 to 3 ∞C) for 4 to 8 years without significant loss

of viability (Heit 1967c). Seeds kept under uncontrolled warehouse conditions have remained viable

for 13 years (Stevens and Jorgensen 1994). There are

71,000 seeds per lb (156,600 per kg) at 100 percent

purity. Minimum purchasing standards are 95 percent

purity and 85 percent germination.

Seed dormancy is complex and variable. A wet

prechilling period of 1 month at 34 ∞F (1 ∞C), followed

by 2 months at 70 ∞F (21 ∞C), and 6.5 months at 34 ∞F

(1 ∞C) relieves dormancy of some seedlots (Rudolph

1974; Wasser 1982). In some cases a 5-day water soak

may successfully substitute for wet prechilling (USDA

Forest Service 1972). Incubation of excised embryos at

68 ∞F (20 ∞C) for 10 to 14 days is recommended as a

relatively rapid means of testing germination.

Seed and dried berries may be fall sown in the field

or nursery, or wet prechilled seed may be planted in

spring. Fall seeding is preferred. Seedlings are vigorous. Bareroot seedlings may be transplanted after one

or two seasons of growth in nursery beds. Container

plants can also be established from seed (Landis and

Simonich 1984).

Seed germination is erratic and may lead to unpredictable stands, although establishment from planting stock is usually quite good. Creating depressions

for water catchments around planted seedlings, and

watering immediately following planting may be helpful, particularly on rocky sites.

Plants may be propagated vegetatively from cuttings taken from spring through fall and rooted on

mist benches (Hartmann and others 1990). Best results are obtained if basal cuts are made between the

nodes about 1.2 inches (3 cm) above the base of the

current year’s growth (Doran 1957). Rooting may be

hastened by treatment with 5,000 ppm IBA. Layers

are occasionally used for vegetative propagation.

Transplants are generally used instead of direct

seeding for landscape and erosion control purposes

to reduce the time until plants reach mature size.

Nursery-grown stock and wildings are easily transplanted. Large stock is more difficult to transplant

due to the development of thick stolons. Commercial

planting stock is widely available, but the seed or

transplants should originate from an area with

characteristics compatible with those of the planting site.

Uses and Management—Oregon grape has been

used in landscape planting as an evergreen ground

cover. It is quite useful because of its colorful flowers,

berries, and leaves. Under cultivation Oregon grape

will grow 2 ft (61 cm) tall. The prickly, hollylike leaves

are brilliantly red to bronze colored in winter and are

used in Christmas decorations. Its spreading habit

and ability to grow on fairly dry, exposed, rocky slopes,

make it a good candidate for mined lands, game

ranges, highways, and recreation areas. Ecotypes growing as an understory with ponderosa pine and other

conifers are, however, not well suited to dry, exposed sites.

The forage value of Oregon grape has been rated as

nearly worthless for livestock and wildlife in summer

(USDA Forest Service 1937). Smith (1953) ranked it

last of 33 shrubs tested as summer forage for mule

deer in northern Utah. However, both livestock and

wildlife use the plants when other vegetation is scarce,

particularly during winter (Monsen and Christensen

1975). Kufeld (1973) found Oregon grape valuable for

Rocky Mountain elk in fall and winter but of low value

in spring and summer when other vegetation was

available. Mule deer use it to a moderate extent in

winter, spring, and fall (Kufeld and others 1973). Patton

and Ertle (1982) reported that in the Southwest it
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Chapter 23 Shrubs of Other Families

Figure 8—Small multistemmed trees of thinleaf alder

border a small stream in eastern Idaho.

receives greatest use by elk in winter; mule deer use it

in the spring and fall. Plants are quite tolerant of

browsing. Once frosted in fall, Oregon grape berries

become more palatable and are used by birds and

small mammals (Thornburg 1982). The species is also

considered desirable for honey production. Plants contain several alkaloids, including berberine, which has

medicinal uses (Suess and Stermitz 1981). The inner

bark and wood of barberris species are yellow and

were used by Native Americans to prepare a yellow

dye.

Seedlings grown on adapted sites may be vigorous,

but require several years to reach mature size. Control

of rodents and rabbit populations has been necessary

to establish young plants in shelterbelts in the Midwest, as Oregon grape is preferred by cottontail rabbits (Swihart and Yahner 1983). Mature plants resprout

following burning, and are resistant to browsing.

Improved Varieties—None. A number of released

varieties have been developed for the ornamental

trade.

Family Betulaceae _______________

Alnus incana

Thinleaf alder

Description—Thinleaf alder is a deciduous, multistemmed shrub or small tree (fig. 8). Plants are usually 3 to 13 ft (0.9 to 4 m) tall (Welsh and others 1987),

but may attain heights of 40 ft (12.2 m) (Arno and

Hammerly 1977; Patterson and others 1985). The

bark is gray to brownish (Patterson and others 1985;

Preston 1948); twigs are puberulent and commonly

glandular (Welsh and others 1987). Leaves are broadly

elliptic or ovate-oblong, 0.4 to 1.2 inches (1 to 3 cm)

long with a dull green color and doubly dentate margins (Hitchcock and Cronquist 1973; Patterson and

others 1985). Male and female flowers are borne in

separate catkins on the same plant. The staminate

catkins are clustered at the ends of twigs, each 1 to 4

inches (2.5 to 10 cm) in length (Hitchcock and Cronquist

1973). The pistillate catkins arise from branches of the

previous season (Welsh and others 1987) and occur in

groups of three to nine on the ends of branches. These

eventually develop into small cones that are 3.6 to 5.2

inches (9 to 13 cm) in length (Hitchcock and Cronquist

1973; Patterson and others 1985). The fruit is a small

single-seeded nutlet with narrow wings (Haeussler

and Coates 1986; Mozingo 1987; Schopmeyer 1974a).

Ecological Relationships and Distribution—

Welsh and others (1987) recognized thinleaf alder as a

portion of a huge circumboreal complex, with Alnus

incana spp. rugosa var. occidentalis being the Old

World portion and Alnus incana spp. rugosa var.

rugosa representing the American plants. Thinleaf

alder, also referred to as mountain or river alder, is one

of approximately 30 species of this genus that occurs in

North America, Europe, and Asia (Schopmeyer 1974a).

It is widespread from Alaska and the Yukon south to

New Mexico and California, usually growing at elevations between 5,000 and 10,000 ft (1,500 and 3,000 m)

in the southern portions of its range (Harrington

1964).

Like most other alders, thinleaf alder occurs in

mountainous regions, often growing along streams

and in wetlands. It is adapted to drier sites and grows

as an understory with various conifers. It spreads

naturally onto disturbances and clearings caused by

wildfires, road construction, and related activities. At

lower elevations and on drier sites, thinleaf alder is

more restricted to wet areas along streams, seeps, and

moist mountain slopes (Arno and Hammerly 1977;

Hansen and others 1988a; Komarkova 1986). It does
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Chapter 23 Shrubs of Other Families

extend into big sagebrush communities at lower elevations growing along streams, seeps, and springs. It is

most prevalent in the Douglas-fir, ponderosa pine,

spruce-fir, and lodgepole pine communities within the

Intermountain region. It occupies sites that may be

seasonally flooded or where water tables remain near

the soil surface. Although often restricted to areas

with high water tables, it is also found on well-drained

sites. It is often abundant as an understory with

conifers, as it is shade tolerant (Haeussler and Coates

1986; Hansen and others 1988b; Kauffman and others

1985; Kovalchik 1987; Padgett and Youngblood 1986;

Youngblood and others 1985).

Thinleaf alder grows on well-developed, highly organic soils (Kovalchik 1987), but occurs on well-drained,

poorly developed, cobbly gravels and sandy textured

soils (Hansen and others 1988a; Kovalchik 1987). This

shrub is able to invade flooded or burned sites (Crane

1982; Zasada 1986). It responds as an early seral

species (Kauffman and others 1985; Zasada 1986), and

is very often abundant following disturbances. It is

commonly encountered dominating disturbed sites

including roadways and riparian habitats.

Plant Culture—Both staminate and pistillate catkins develop during the growing season and persist

through most winter months. The catkins begin growth

early in spring before leaves develop. Flowering occurs

in early March and April, depending on elevation and

site location. Fruits ripen in early fall (Schmidt and

Lotan 1980). Fruits are small winged nutlets borne in

pairs on bracts of small cones or strobiles—when

dispersed they are spread by wind or water (Haeussler

and Coates 1986; Mozingo 1987; Schopmeyer 1974a).

Thinleaf alder normally produces abundant seed crops

most years. Some individual plants bear heavy crops

every year. There are about 675,000 cleaned seeds per

lb (1,488,000 per kg) (Schopmeyer 1974a). Seed viability is quite variable. Schopmeyer (1974a) reported

some collections contain less than 5 percent viable

seed. However, good quality seed can be obtained by

collecting from healthy bushes early in the season

before seeds shatter. Bushes with disfigured cones or

infested with disease should be avoided. Also, seeds

from plants that are weakened by climatic stress

should not be harvested. Seed samples should be

cleaned and tested for germination before seeding

rates are determined. Seeds should not be stored in

open warehouses for more than 2 years because viability decreases rapidly after this time. If possible, airdried seeds should be stored in sealed containers at 34

to 38 ∞F (1 to 3 ∞C) (Schopmeyer 1974a). Seeds are

nondormant when collected and do not require wet

prechilling to germinate (Haeussler and Coates 1986).

Thinleaf alder seeds are small, lightweight, and

easily dispersed. Broadcast seeding, particularly using

aircraft, may create irregular stands, as seeds can be

carried considerable distances with strong winds. Seedlings developing from seeds broadcast on rough seedbeds establish well if the surface does not dry rapidly.

Seedlings do not establish well on compacted surfaces

and impervious soils. Thinleaf alder has been broadcast seeded in a slurry using a hydromulch, but with

limited success. Better stands have been achieved by

dry seeding.

Thinleaf alder seeds can be planted with most drill

seeders. If seeded alone in nursery beds, the seedlot may

require dilution with inert material to aid in regulating

the seeding rate. However, many precision nursery

seeders can plant pure alder seeds. Seeds should be

placed about 0.25 inch (0.6 cm) deep on firm, but

noncompacted, seedbeds. Planting on seedbeds having

considerable surface litter has been quite successful.

Within its range of climatic adaptation, thinleaf

alder will invade fresh disturbances. It responds as a

pioneer plant capable of establishing in open areas

(Zasada 1986). It is able to compete well with herbaceous species when seeded on both disturbed and

undisturbed sites. Seedlings develop quickly and can

establish amid considerable herbaceous competition.

New shrub seedlings often invade roadway and mine

disturbances where seeded herbs have previously established. On infertile soils, thinleaf alder may grow

more vigorously than the herbs.

Thinleaf alder is not universally adapted to all soil

disturbances and it is particularly sensitive to competition from some species. Although the shrub invades

areas seeded to grasses, its growth can be suppressed

by some herbaceous species, particularly smooth brome.

Planted seedlings that do not become inoculated with

soil organisms remain small and can be suppressed.

Although these factors are not well understood,

irregularities in plant performance on disturbed sites

can be expected.

Thinleaf alder can be easily propagated and planted

as bareroot nursery stock or container material (Platts

and others 1987). Transplants usually establish quickly

and grow considerably during the year of planting.

One-year-old transplants develop fibrous root systems that greatly enhance survival. Thinleaf alder is

reported to fix nitrogen (Haeussler and Coates 1986).

Plants inoculated with nitrogen-fixing organisms begin

growth soon after planting. Other plants that apparently lack these organisms grow slowly, but respond

dramatically if inoculated. Adding soil containing natural microflora to containerized stock appears beneficial. When plants are established on harsh sites,

fertilizer applications have benefitted initial growth.

Seedings of thinleaf alder on sites supporting a herbaceous cover are often successful in conifer forest communities. This shrub has frequently invaded roadways,

logging disturbances, and mine sites that were first

seeded to introduced and native herbs (Plummer 1977).
618 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

The shrub invades seeded areas where established

herbs are weakened due to decreases in soil fertility.

Its seedlings apparently are very competitive and

vigorous. Once established, few other plants grow as

rapidly.

Uses and Management—Thinleaf alder is particularly useful in providing cover and forage for wildlife. Abandoned logging roads and other disturbances

that are seeded to grasses often are invaded by thinleaf

alder, snowbrush ceanothus, chokecherry, and other

important browse plants. Game animals seek these

areas to graze the variety of plants. Animals also

utilize thinleaf alder in burned and cutover lands;

plants quickly recover and produce an abundance of

herbage. Use by big game animals varies among sites.

Light to moderate use has been reported by deer, elk,

and moose (Heale and Ormrod 1982; Kauffman and

others 1985; Knowlton 1960, Kufeld 1973). However,

Kufeld and others (1973) reported that alder is an

important browse for mule deer. Beaver and other

small animals and birds also utilize this species (Arno

and Hammerly 1977; Rue 1964). Thinleaf alder provides excellent cover for wildlife. It occupies riparian

zones, wetlands, and brush fields where animals concentrate. The establishment and growth traits of this

species cause it to be one of the most useful shrubs for

revegetation of riparian sites in the Intermountain

region.

Varying degrees of use by cattle and sheep are

reported for thinleaf alder (Dittberner and Olsen 1983;

Roath and Krueger 1982). Animals selectively graze

certain stands more heavily than others. Use is somewhat regulated by access, acceptance of associated

plants, and season of grazing. Livestock graze thinleaf

alder very heavily in many riparian areas. The plant

is frequently maintained in a stunted condition, but is

able to recover when grazing is well managed. New

plantings can be destroyed by heavy grazing.

Thinleaf alder is particularly useful as a ground

cover and erosion control plant. It is one of a few shrubs

that, as a seedling, is able to compete with seeded

herbs, and can be seeded in mixtures. It grows rapidly

and provides cover and herbage within a few years. It

is well adapted to a wide array of soil conditions,

particularly infertile sites. Its ability to fix nitrogen

apparently contributes to its adaptability to harsh

situations. The presence of thinleaf alder benefits the

growth of understory herbs, and thus improves ground

cover and stability on erodible sites. When transplanted onto open disturbances it grows rapidly and is

able to stabilize sites where seeding is difficult.

Thinleaf alder has been used successfully to revegetate road and logging disturbances in the Idaho

Batholith and on a number of mine sites. It has spread

onto abandoned mine disturbances, including rocky

dredge piles and exposed substrata.

Collections of thinleaf alder do not appear to be as

site specific as many other shrubs. Plantings have

been successful over a range of elevations and site

conditions. The species should not be planted on sites

where it does not normally occur, but it can be used on

mine or road disturbances within its natural range of

occurrence.

Thinleaf alder plants are not usually lost to insect

damage, climatic stress, or herbivore grazing. The species is long lived and compatible with overstory trees.

It is able to exist in open or shaded areas and can be

used as a low maintenance species in recreational

plantings. Thinleaf alder is an attractive shrub and

can be used as a screen or specimen plant for horticulture plantings. It remains vigorous and green the

entire summer and may be used to reduce fire spread

around recreational sites.

This species has been overlooked in many wildland

restoration programs. Although recognized as a rapiddeveloping species capable of producing considerable

biomass, it has not been emphasized for watershed or

wildlife plantings.

Varieties and Ecotypes—None.

Family Betulaceae _______________

Betula glandulosa

Bog birch

Description—Bog birch is a deciduous, highly variable, spreading to erect monoecious shrub ranging

from 0.3 to 19.7 ft (0.1 to 6 m) tall. Plants have one to

many main stems and deep, spreading root systems.

Alternative common names are glandular, resin,

swamp, marsh, and dwarf birch. Young branches are

densely puberulent and resinous with yellowish

wartlike, crystalline glands. Bark is reddish-brown to

gray or purplish and does not readily peel. Lenticels

are rarely conspicuous. Leaves are alternate, oval to

orbicular, elliptic, or sometimes obovate or ovate. They

are 0.4 to 0.8 inch (1 to 2 cm) in length, but range from

0.2 to 1.6 inches (0.5 to 4 cm) long, with finely serrate

or crenate-serrate edges (fig. 9). Blades are bright

green, thick and leathery, glandular on both surfaces,

glabrous above, and paler and puberulent beneath.

Staminate catkins are elongate and pendulous in

clusters of one to four. They develop in summer and

flower with or before leaf development the following

spring. Staminate flowers are three per cluster, each

with three perianth segments and two stamens subtended by three bracts. Pistillate catkins appear with

the leaves. They are conelike, erect, and solitary with

two or three flowers per bract. Fruits are narrowwinged, single-seeded samaras. Flowering occurs from

April to September. Samaras ripen from July to October (Brinkman 1974c; Hitchcock and others 1964;
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 619

Chapter 23 Shrubs of Other Families

Viereck and Little 1972; Welsh and others 1987); they

are dispersed by wind, gravity, and sometimes water.

They may also be blown for some distance across

crusted snow. Throughout much of the species range,

reproduction is primarily by seed. In the northernmost limits of its range, seedlings are rare and plants

spread primarily by layering (Hermanutz and others

1989; Weis and Hermanutz 1988).

Ecological Relationships and Distribution—

Bog birch is widely distributed from the interior of

Alaska across northern Canada to Labrador and

Greenland. In the West, it is found in mountainous

areas from coastal British Columbia to California

and Colorado at elevations from 4,000 to 11,000 ft

(1,200 to 3,400 m) (Dittberner and Olsen 1983;

Harrington 1964; Welsh and others 1987). Bog birch

occurs in wet meadows, fens, swamps, bogs (sphagnum and nonsphagnum), muskegs, moist-to-wet tundra, wet meadows, and along low gradient streams

and lakes. It is also found on sites where the water

table remains high from runoff in nearby uplands. Bog

birch maintains itself in these moist habitats and

appears to be a climax species under such conditions

(Marr 1961; Pojar and others 1984). In alpine areas it

may also be found in dry rocky habitats (Brayshaw

1976; Komarkova 1986). Bog birch is highly frost

tolerant and is widely distributed in permafrost areas

(Krajina and others 1982). Its shade tolerance is moderate to high.

Bog birch most often occurs on wetland sites in

lodgepole pine, Engelmann spruce, or subalpine fir

forest types (Hansen and others 1988b; Kovalchik

1987; Olson and Gerhart 1982; Pierce and Johnson

1986). It is frequently associated with alder, willows,

Woods rose, Douglas spiraea, blueberry, mountain huckleberry, and redosier dogwood. Herbaceous species

associated with bog birch include rushes, horsetails,

soft-leaved sedge, beaked sedge, bluejoint reedgrass,

and tufted hairgrass (Hitchcock and others 1964;

Kovalchik 1987; Patterson and others 1985; Welsh

and others 1987; Youngblood and others 1985).

Bog birch commonly grows in fens, swamps, or bogs

having large accumulations of organic matter derived

from sphagnum or nonsphagnum plant materials. On

flood plains it often grows in soils with textures ranging from silty to fine sandy loams to organic loam

textures (Kovalchik 1987). Mineral requirements of

bog birch may vary with population. Krajina and

others (1982) and Moss and Wellner (1953) reported

the species occurs in acidic or humic soils that are low

in nutrients, particularly calcium and magnesium.

They noted the absence of bog birch from swamps and

fens with high nutrient contents derived from adjacent uplands. However, Pojar and others (1984) noted

the reverse to be true.

Plant Culture—Most samaras are wind dispersed

in fall. Smaller quantities are released through the

winter and early spring as the catkins disintegrate on

the shrub. Many of these may be consumed by birds.

Samaras are sometimes collected from the surface of

crusted snow. Fruits are harvested, processed, and

stored as described for water birch. There are between

3 and 5 million cleaned seeds per lb (6,600,000 and

11,000,000 per kg) (Brinkman 1974c). Culture is as

described for water birch (Brinkman 1974c; Lotan

1981). Kelly (1970) reported that bog birch’s value in

rehabilitation is limited; it does not transplant easily.

In the northern portion of the species range, few viable

seeds are produced, and bog birch spreads primarily

by stem layering (Hermanutz and others 1989; Weis

and Hermanutz 1988).

Uses and Management—Bog birch is consumed

by many wildlife species. Moose, elk, and mule deer

make moderate to heavy use of the shrub in summer

and winter (Dorn 1970; Kufeld 1973; Kufeld and

others 1973; Zach and others 1982). Pine siskin, chickadees, kinglets, and many other birds use the catkins,

buds, and seeds (Brayshaw 1976; Komarkova 1986;

Pojar and others 1984; Stephens 1973). Livestock

make only light to moderate use of bog birch except in

late summer when the boggy soils dry out (Dayton

1931; Hansen and others 1988b; Kovalchik 1987).

Energy and protein value of the species are low.

Wetland areas supporting bog birch burn only infrequently due to the normally high water content of the

vegetation and soil (Crane 1982; Kovalchik 1987).

Such sites frequently act as firebreaks, but sometimes

burn during dry summers or in late fall. Shoot systems

may be destroyed but the plants resprout rapidly,

particularly if the organic layer of soil around the base

is not removed. Germination of wind-dispersed seeds

from surviving plants, or seeds arriving from offsite,

Figure 9—Stems of bog birch have reddish to purplish

bark and thick, serrate leaves.
620 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

might be facilitated or enhanced by the presence of

exposed mineral soils.

Varieties and Ecotypes—None.

Family Betulaceae _______________

Betula occidentalis

Water birch

Description—Water birch, also known as spring or

red birch, ranges from shrublike to treelike in growth

habit (fig. 10). Plants grow rapidly and are short lived.

They may attain heights of 36 ft (11 m), with clump

diameters of up to 20 ft (6 m), and major trunk

diameters of 1 ft (0.3 m). New stems develop from the

root crown, eventually forming dense, multistemmed

clumps with ascending branches and open crowns

(Hansen and others 1988a; Youngblood and others

1985). The thin, smooth bark is dark brown to black on

young trunks; it becomes lustrous coppery to brownish

red on older trunks. Lenticels are light colored and

vertical. The alternate, deciduous leaves are petiolate

with bright green stipules. They are glabrous to pubescent, glandular, ovate to suborbicular or slightly obovate, and 1 to 2 inches (2.5 to 5 cm) long. The blade apex

is rounded to acute, the base usually rounded, veins

prominent, and the margins sharply once or twice

serrate.

Male and female flowers develop in separate inflorescences on a single plant. Staminate catkins are

formed during the summer or fall and rapidly elongate

into tassel-like catkins 1 to 4 inches (2.5 to 10 cm) long

the next spring. Staminate flowers grow in clusters of

three; each consists of three perianth segments and

two stamens, all subtended by three bracts. Blooming

occurs as the leaves expand. Female flowers develop in

conelike catkins 1 to 1.5 inches (2.5 to 4 cm) long.

Water birch flowers in April and May. Fruits turn

brownish tan as they ripen in fall. Samaras are shed

from the catkins in fall and winter and are dispersed

by wind, gravity, and sometimes water. They may be

blown for considerable distances across crusted snow.

Weather conditions and foraging by seed-eating birds

contribute to overwinter disintegration of the catkins

(Brinkman 1974c; Lanner 1983).

Ecological Relationships and Distribution—

There are more than 40 species of Betula in the North

Temperate and Arctic Zones. Some taxa hybridize,

producing intermediate populations in zones of distribution overlap (Dugle 1966; Hitchcock and Cronquist

1973). Water birch is distributed from Alaska to California and east to Saskatchewan and New Mexico. It

forms dense thickets in riparian woodland communities and in moist areas of drier habitats (Arno and

Hammerly 1977; Hansen and others 1988a; Padgett

and Youngblood 1986; Welsh 1974). It is common

along streams, on steep slopes, or on alluvial terraces

from Engelmann spruce, ponderosa pine, and Douglasfir zones to sagebrush communities (Hansen and others

1988a,b; Padgett and Youngblood 1986; Youngblood

and others 1985). Communities tend to remain rather

stable in spite of annual flooding (Padgett and others

1989; Youngblood and others 1985). However, the

plants are susceptible to windthrow because they

grow on sites with high water tables and are usually

shallow rooted (Lanner 1983). In eastern Washington,

Oregon, western Idaho, and southern British Columbia, water birch hybridizes with paper birch, producing many localized intermediate forms (Dugle 1966;

Hitchcock and others 1973). It is closely related to bog

birch (Betula glandulosa), and in western Canada

hybridizes with this species (Dugle 1966).

Water birch is listed as a dominant species in a

number of communities or habitat types of the Intermountain region (Hansen and others 1988a,b; Olson

and Gerhart 1982; Padgett and others 1989;

Youngblood and others 1985). It occurs with many

woody riparian tree and shrub species including cottonwoods, maples, aspen, thinleaf alder, willows, alder,

and redosier dogwood (Lanner 1983; Olson and

Gerhart 1982; Padgett and others 1989). It is also

associated with many shrubs that are facultative

riparian species such as Woods rose, Nutka rose,

chokecherry, Saskatoon serviceberry, currant, and

skunkbush sumac (Hansen and others 1988a,b; Kelly

1970; Olson and Gerhart 1982; Padgett and others

1989; Youngblood and others 1985).

Water birch is adapted to soils ranging in texture

from silty to sandy to coarse-textured types that contain

at least 35 percent rock fragments. These often have

Figure 10—Water birch forms attractive foliage

and flowers as it grows along valley bottoms and

riparian sites.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 621

Chapter 23 Shrubs of Other Families

thin profiles and overlie river cobbles or other rocky

substrates (Hansen and others 1988b; Padgett and

others 1989). Nutritional requirements for magnesium and calcium are relatively high (Krajina and

others 1982).

Plant Culture—Seeds should be collected from

selected young, accessible, highly productive trees by

hand stripping the catkins into bags before they begin

to disintegrate. Fruits are sometimes shed on snow

and may be hand collected.

Catkins should be thoroughly dried prior to extraction. The tiny samaras are separated from the catkins

by screening them through a round-holed screen and

fanning to remove the remaining bracts (Brinkman

1974c). Samaras should be dried prior to storage at 36

to 38 ∞F (2.2 to 3.3 ∞C).

Seed fill and quality vary greatly among years. Fill

should be estimated prior to harvest by observing

seeds under transmitted light (Patterson and Bruce

1931). Wet prechilling for 4 to 8 weeks releases embryo

dormancy. Dormancy may also be overcome by germinating seeds in light (Brinkman 1974c; Yelenosky

1961).

Plantings are generally established from bareroot or

container stock rather than from seed. Emergence of

seedlings from direct seedings on wildlands is erratic.

Only 15 to 20 percent of the planted seeds of these

species usually produce usable seedlings. Seeds planted

in nursery beds or containers must be covered as

lightly as possible (Hartmann and others 1990). Covering may not be necessary if the soil surface is kept

moist. Newly emerged seedlings are susceptible to

damping off at low temperatures (Marchant and

Sherlock 1984). They benefit from shade for the first

2 or 3 months (Brinkman 1974c). Seedlings develop

rapidly, producing dense root systems, and may be

transplanted after one or two growing seasons

(Brinkman 1974c). Cuttings are difficult to root, but

leafy cuttings will root in summer if planted under

glass and treated with IBA (Doran 1957; Hartmann

and others 1990). Low-growing forms are easily propagated by layering.

Seed or cuttings used to produce planting stock

should be harvested from areas near the planting site.

Seedlings should be planted early in spring while

native birch near the planting site is still dormant.

Seedlings planted on adapted sites with good soil

water establish well and grow rapidly if protected from

browsing and competition.

Uses and Management—Water birch provides

food and cover for many wildlife species (Kufeld 1973;

Gullion 1964; Rue 1964; Stark 1966). It is useful for

streambank stabilization in adapted areas. It grows

rapidly, producing a dense, fibrous, but shallow root

system. It is useful in shelterbelts if adequate water is

available (Cook 1981), and has some potential as an

ornamental. The hard wood is used locally for firewood

and fenceposts (Lanner 1983).

Dense thickets or corridors of water birch provide thermal, hiding, and travel cover for a wide array of wildlife

species (Hansen and others 1988a). Palatability of

water birch to big game is usually low. It may be used

when other browse is limited (Hansen and others

1988b; Kufeld 1973; Kufeld and others 1973). Its

presence in many riparian woodland communities

contributes to their structural diversity and provides

habitats for many bird species (Lanner 1983;

Youngblood and others 1985). Seeds, catkins, buds,

and sap of water birch are used by many birds including

grouse, redpolls, pine siskin, chickadees, kinglets, rednapped sapsucker, and broad tailed hummingbirds

(Brinkman 1974c; Gullion 1964; Platts and others 1987).

Water birch is not highly palatable to livestock,

although it is used to some extent by sheep and goats

(Youngblood and others 1985). Dittberner and Olsen

(1983) rated energy and protein values as fair. Productivity is often reduced by flooding and sedimentation,

and the nearly impenetrable monotypic stands reduce

livestock access (Hansen and others 1988a).

Water birch is usually associated with riparian

communities that act as firebreaks and burn only

during late summers or dry years. Shoots will burn in

intense fires. Burned plants generally resprout from

the root crown, which is often protected to some extent

if the soil is wet. New plants may also develop from

seed delivered by wind or water from areas outside the

burn. Germination is favored by exposure of mineral

soils and reduced competition following fires (Crane

1982; Hansen and others 1988a,b).

In Utah, water birch is recommended for landscape

plantings in residential areas developed on historic

mule deer winter range. Many traditionally used landscape plants in these areas have been heavily browsed

and damaged by mule deer. Water birch is normally

browsed only moderately by mule deer and recovers

quickly (Austin and Hash 1988).

The root systems of water birch stabilize banks; overhanging branches shade the water and supply organic

matter, which improve fish habitat (Youngblood and

others 1985). Soils on steep streambanks may be

susceptible to erosion, especially along trails used by

livestock and wildlife. Heavy recreational use associated with fishing may also increase sloughing rates

(Hansen and others 1988a). Stands should, therefore,

be maintained for their streambank stabilization value

(Hansen and others 1989; Youngblood and others

1985).

Varieties and Ecotypes—None.
622 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Family Betulaceae _______________

Betula papyrifera

Paper birch

Description—Paper birch is a treelike species growing from 60 to 130 ft (18.3 to 39.6 m) tall and 2 to 3 ft

(60 to 90 cm) in diameter from a deep root system

(fig. 11). Alternative names are canoe, white, and

silvery birch. Trees are distinguished by their grayish

white to coppery bark with long brown horizontal

lines. The outer bark often peels in long strips; the

inner bark is orange. The leaves are ovate to acuminate and doubly serrate, usually with tufts of hairs in

the vein axils on the lower surface. Leaves are 2 to 5

inches (5 to 13 cm) long and 1 to 2 inches (2.5 to 6 cm)

wide. Male flowers develop in early spring on long,

yellowish, pendulous catkins. Female cones are 1 to 2

inches (2.5 to 6 cm) long, hanging from slender stalks.

Fruits are translucent samaras with the wings slightly

broader than the nutlet. Paper birch and water birch

hybridize widely in areas where their ranges overlap

(Hitchcock and others 1961; Marchant and Sherlock

1984; Welsh and others 1987).

Ecological Relationships and Distribution—

Paper birch is distributed from Alaska east to

Newfoundland and south across the northern tier of

States including Washington, northern Idaho, and

Montana. It occurs in riparian habitats, dry bogs,

swamps, and poorly drained acidic soils with pH as low

as four (Marchant and Sherlock 1984). It also grows on

open slopes, rock slides, forest openings, and disturbances. It occupies soils with silty loam, sandy, and

gravely textures. It is an aggressive pioneer species on

logged or otherwise disturbed areas, and rapidly invades areas where mineral soils are exposed.

Plant Culture—Culture and management are

largely as described for water birch (Brinkman 1974c).

Cones are more accessible, abundant, and easily harvested from young trees. Minimum seed-bearing age

is about 15 years (Brinkman 1974c). Catkins may open

during dry, cold periods from late fall well into winter;

when large numbers of samaras are deposited on the

surface of crusted snow.

Seeds are cleaned using, a number seven sieve to

separate fruits from chaff (Marchant and Sherlock

1984). There are 610,000 to 4,120,000 cleaned seeds

per lb (1,344,800 to 9,083,000 per kg); the average is

1,380,000 seeds per lb (3,042,000 per kg). Longevity of

seeds dried to a water content of 1 to 5 percent and

stored at room temperature is about 1.5 to 2 years.

Viability tends to decrease rapidly at greater water

contents, even if seeds are kept in cold storage

(Brinkman 1974c).

Seeds require a wet prechilling treatment of at least

30 days or fall sowing to produce nursery stock. Surface-sown seeds exposed to light will germinate without cold wet prechilling. Seedlings are subject to damping off if crowded or if temperatures are cool. They are

resistant to frost but not shading (Krajina and others

1982). Propagation from cuttings is not recommended

(Marchant and Sherlock 1984).

Uses and Management—Seedlings have been

used for revegetating riparian sites and forest disturbances, improving wildlife cover, landscaping recreational sites, and in ornamental plantings. Plants

have established and survived well, but have not

been extensively used in wildland plantings. Young

plants grow quite rapidly and compete well with

understory competition. Paper birch resprouts following fire and is capable of withstanding both periodic flooding and drought (Marchant and Sherlock

1984). It is browsed to some extent by wildlife, horses,

and cattle. Fruits are eaten by many birds.

Figure 11—White-barked, multiple-stemmed paper Varieties and Ecotypes—None.

birch is often found growing in a forest openings.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 623

Chapter 23 Shrubs of Other Families

Family Caprifoliaceae ____________

Lonicera ciliosa, Orange honeysuckle

Lonicera involucrata, Bearberry honeysuckle

Lonicera utahensis, Utah honeysuckle

Introduction—There are about 180 honeysuckle

species distributed across the Northern Hemisphere,

Africa, Java, and the Philippines (Brinkman 1974f),

although only about 20 species occur in North America

(Hitchcock and others 1959). All are widely planted for

their fragrant flowers and ornamental fruits. Some

species provide food and cover for wildlife and are

valuable for erosion control and shelterbelt plantings

(Adkins 1980; Brinkman 1974f; Plummer 1977;

Plummer and others 1968; USDA Forest Service

1985a).

Different species of honeysuckle possess important

traits that are useful for range and wildland enhancement. Some of the more important characteristics

include consistent and abundant fruit production, low

stature, cover for birds and small mammals, excellent

establishment traits from direct seeding and transplanting, rapid growth, and excellent windbreak attributes and conservation features.

Plant Culture—All three species described in this

chapter produce small berries and seeds. Berries are

usually dislodged by stripping or beating the stems

when fruits are mature. Seeds are extracted from the

berries by macerating the pulp and flushing with

water. Once seeds are cleaned, they are easily seeded

with conventional seeders. Seedlings develop quite

successfully, but most plantings have been completed

using bareroot or container transplants.

Family Caprifoliaceae

Lonicera ciliosa

Orange honeysuckle

Description—Orange honeysuckle is a native trailing or twining vine. Stems are branched and may

become 20 ft (6.1 m) long (Hitchcock and others 1959).

Although plants normally grow in wet sites in aspen

and conifer forests, they are also encountered in open

areas on rocky and shallow soils. This vine naturally

occurs in southwestern Canada, from Oregon to California, and east to Idaho and Montana (Hitchcock and

others 1959). Individual plants can spread over large

areas on fertile or disturbed soils. They root at the

junction of each node if the stem remains in contact

with a wet soil surface (Monsen 1975). This species is

easily transplanted from stem cuttings.

Uses and Management—The orange yellow to

orange red flowers and foliage are very attractive and

could be used in roadside, summer home, and recreational landscaping. In northern Idaho this species

has survived serious air pollution created by smeltering

operations, while a number of other native shrubs

have succumbed (Monsen 1975).

Varieties and Ecotypes—None.

Family Caprifoliaceae ____________

Lonicera involucrata

Bearberry honeysuckle

Description—Bearberry honeysuckle is a native

shrub that is found throughout the Intermountain

West from Alaska to Quebec and south to California,

New Mexico, and Mexico (Viereck and Little 1972;

Welsh and others 1987). It has yellow to purplish

tinged flowers (fig. 12) and black fruit. There are about

326,000 seeds per lb (718,700 per kg) at 100 percent

purity. This shrub grows to 2 to 6 ft (0.6 to 1.8 m) tall.

It occurs on fertile soils along streambanks and in

riparian sites in the aspen, Douglas-fir, and spruce-fir

communities.

Uses and Management—Bearberry honeysuckle

has been planted with some success in conservation

and wildlife projects. It has not done well when planted

on disturbed sites. This species is used by large and

small game and livestock during spring and summer

Figure 12—Bearberry honeysuckle, widespread in

Western North America, is used in conservation

plantings because of its attractive flowers.
624 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 14—Blue elderberry growing at the base of

a rocky slope in eastern Idaho.

months. It has potential for restoring disturbances

within its natural range, but its seeds are not easily

harvested, thus seed availability is limited.

Varieties and Ecotypes—None.

Family Caprifoliaceae ____________

Lonicera utahensis

Utah honeysuckle

Description and Distribution—Utah honeysuckle is a native shrub that grows 3 to 6 ft (0.9 to 1.8 m)

tall (Welsh and others 1987) (fig. 13). It has pale

yellow to yellowish-white flowers and red fruit. It

occurs from British Columbia and Alberta, south to

California and Wyoming. It is fairly common in the

Intermountain West within mountain brush, ponderosa pine, aspen, and spruce-fir communities.

Uses and Management—This shrub has good fire

tolerance, and is used extensively by livestock and

game during the summer and fall seasons. Utah honeysuckle does well under heavy grazing and survives

heavy trampling. It has excellent ornamental characteristics including showy flowers and fruit. The plant

establishes well from transplanting and can withstand

considerable traffic and other abuses. It should be

considered for use in restoration projects, recreational

areas, summer homes, and administrative sites.

Varieties and Ecotypes—None.

Family Caprifoliaceae ____________

Sambucus cerulea

Blue elderberry

Description—Blue elderberry (fig. 14) is a large

spreading shrub or small tree growing to 10 ft (3.0 m)

tall. It has distinctive opposite branches and leaves,

pithy young stems, and older grayish stems with

irregularly furrowed and ridged bark. Flowers are

white and borne in large flat-topped clusters (compound cymes). Fruits are black to blue (Hitchcock and

others 1959; Welsh and others 1987). The root system

is generally fibrous and spreading, but a thick taproot

may be present (Haeussler and Coates 1986; Welsh

and others 1987).

Ecological Relationships and Distribution—

Blue elderberry is most common on moist, well-drained

sunny sites, but it is also somewhat shade tolerant

(Dayton 1931; Plummer and others 1968; Van Dersal

1938) and occurs at elevations between 4,250 and

8,200 ft (1,300 and 2,500 m) (USDA Forest Service

1937). It ranges from British Columbia and western

Alaska south to California, Arizona, New Mexico,

Texas, and northern Mexico (Dayton 1931; Harrington

1964; Hitchcock and others 1959; Little 1979; Powell

1988). It may be found on a wide range of soil types,

growing from the sagebrush grass zone to well above

the mountain brush and ponderosa pine communities.

It also grows in openings in aspen and spruce-fir

communities (Welsh and others 1987). Blue elderberry is found in canyon bottoms, along streams, and

on sites that are wet during the spring. In riparian

areas, blue elderberry can occur in dense stands; in

other areas individual plants are generally widely

spaced. Plants are often abundant in burned areas.

Figure 13—Distinctive flowers and leaves of Utah

honeysuckle, an understory shrub that grows

with mountain brush, ponderosa pine, aspen, and

conifer forests.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 625

Chapter 23 Shrubs of Other Families

Plant Culture—Blue elderberry generally produces

a good seed crop every year. Berrylike fruits are

produced in large clusters. Each fruit contains three to

five seeds (Hitchcock and others 1959). Fruit clusters

are easily collected by cutting them from the stem.

Seeds are extracted from the fruit by using a Dybvig

macerator. The resulting pulp and seeds are dried,

then lightly chopped and separated using an air

screen fanning machine. Unfilled seeds can be

removed by flotation. At 100 percent purity there are

about 217,000 seeds per lb (478,500 per kg). Acceptable purity is 95 percent and germination 50 percent.

Good viability is retained for up to 16 years (Haeussler

and Coats 1986).

Germination of blue elderberry is inhibited by a hard

seedcoat and embryo dormancy (Plummer and others

1968). Without artificial treatment, germination of

planted seeds may occur sporadically over one to three

growing seasons. Scarification in acid or a 2-month

warm pretreatment followed by a wet prechilling of

up to 5 months improve germination (Brinkman

1974j). Fall seeding is recommended to permit overwinter wet prechilling (Plummer and others 1968).

Seeds in the soil seedbank often germinate readily

following fire (Heit 1967a).

Elderberries can be established by direct seeding in

favorable spots (Plummer and others 1968). Seed may

be hand planted in pits on burns, chained areas, or

disturbed riparian sites. Elderberry seed may also be

planted alone or with other shrubs using Hanson or

thimble seeders. Elderberries should be planted separately from grasses to protect the shrub seedlings from

competition. Seeds should be covered 0.25 inch (6 mm)

deep on a firm seedbed.

Blue elderberry is easily propagated, planted, and

established using container or bareroot stock (Plummer

and others 1968). Plants grow quickly in nursery beds,

and often develop a thick taproot and spreading root

system. Root pruning can eliminate or reduce this

problem. Blue elderberry can also be established from

cuttings (Plummer and others 1968).

Blue elderberry shrubs begin growth slowly in early

spring, although rapid vegetative development occurs

in June or July. The number of stems and buds, as well

as total productivity, fluctuate greatly from year to

year. Older branches die back during the winter. New

growth develops from large vegetative buds on the

rhizomes.

Uses and Management—Blue elderberry has been

used successfully in soil stabilization (Plummer and

others 1968), riparian plantings (Carson and Edgerton

1989; Goldner 1984), streambank stabilization planting (Lines and others 1979), and for revegetation of

mined areas (Ferguson and Frischknecht 1985;

Hungerford 1984; Monsen 1984). Fruits of blue

elderberry are gathered and used for making wine,

jellies, candy, pies, and sauces (Mozingo 1987; Powell

1988).

Blue elderberry is grazed by livestock and wildlife

species throughout the year (Dittberner and Olsen

1983; Kufeld 1973; Kufeld and others 1973; Martin

and others 1951; Robinette 1972). However, use varies

by season (Kufeld 1973; Sampson and Jesperson 1963;

Smith and Hubbard 1954; Steele and Geier-Hayes

1987). Spring and summer use are light. Many mature

plants grow so tall that by late summer much of the

foliage is out of reach of grazing animals. Palatability

and use increase dramatically after the fruit ripens

and the foliage is frosted (Dayton 1931). Plants are

heavily used by cattle, deer, and elk in fall; they are

especially palatable to sheep. Use decreases in the

winter after leaf fall, but plants provide important

herbage. The berries and dried fruit, which often

remain on the shrubs through the winter, are consumed and dispersed by birds and many other animals. Buds are used by grouse, rodents, and big game

animals in winter. The shrub provides cover for big

game and nongame species, and perching and nesting

sites for birds (Brown and others 1977; Dittberner and

Olsen 1983; Gray and Greaves 1984). Blue elderberry

is persistent and recovers well from moderate grazing

and trampling (Monsen 1984; Plummer and others

1968; Van Dyne and Payne 1964). Continual, heavy

grazing can be detrimental. Seeded areas should not

be grazed for two or three growing seasons following

seeding. Buds and branches may be defoliated by

grasshoppers during mid or late summer during years

when population densities are high. Plants generally

recover the following year.

Blue elderberry exhibits good fire tolerance (Aro

1971) and is able to resprout from the root following

fire (Little 1979; Preston 1948; Steen 1965; Van Dersal

1938). Germination of seed buried in the soil is enhanced

by fire (Heit 1967a; Morgan and Neuenschwander

1988).

Varieties and Ecotypes—None. There are many

ecotypes within the species. Drought-tolerant, shrubby

populations associated with sagebrush types do well

with only 12 inches (30 cm) of annual precipitation.

Tall, almost treelike populations also occur. Some

populations are consistently good seed producers, while

others produce erratic seed crops.

Family Caprifoliaceae ____________

Sambucus racemosa

Red elderberry

Description—Red elderberry is a native circumboreal shrub (Hitchcock and others 1959). Two natural

varieties occur in the Western United States: S. r. var.

microbotrys has red fruits and is widely distributed,
626 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

ranging from Wyoming and northern California south

to Mexico. S. r. var. melanocarpa produces black

fruits, and occurs primarily in the Rocky Mountains

from Canada south to the Pacific Northwest and

Wyoming (Welch and others 1987).

Red elderberry has distinctive opposite branches

and leaves. Leaves are pinnately compound, leaflets

are five to seven serrate, pointed, 1.2 to 5.9 inches (3 to

15 cm) long, and 0.4 to 2.4 inches (1 to 6 cm) broad.

Plants are fairly short, usually about 3 to 6 ft (0.9 to

1.8 m) in height. Stems have warty bark with brown

pith; they have a rank odor if bruised and crushed

(Kelly 1970; Viereck and Little 1972). The numerous

white to cream-colored flowers are borne in terminal

pyramidal clusters (cymes) less than 3 inches (8 cm)

wide. The fruit is a red drupe that ripens in September.

Each drupe contains two to four seeds (Hitchcock and

others 1959; USDA Forest Servcie 1985; Welsh and

others 1987).

Ecological Relationships and Distributions—

Red elderberry occurs in subalpine, spruce-fir, lodgepole pine, and aspen communities (fig. 15). It is distributed across North America from Alaska to

Newfoundland and south to New Mexico and Georgia

(Great Basin Flora Association 1986; Viereck and

Little 1972; Worley and Nixon 1974). It prefers full to

intermediate sunlight, and is not found in full shade

(Haeussler and Coats 1986). Greatest areas of abundance are in openings where winter snow accumulates

and remains late in the spring, riparian areas, meadows, parks, and other wet areas. Red elderberry grows

in association with tall and intermediate forbs and

grass communities. It occurs singly and in patches of

various sizes.

Plant Culture—Red elderberry flowers in midsummer. Fruits ripen in mid to late September. Good

seed crops are produced most years. Fruit clusters are

harvested by cutting them from the stems. Seeds are

extracted with a Dybvig macerator. Seeds and pulp

are dried, lightly chopped, and processed through an

air screen fanning machine to separate seeds and

debris. At 100 percent purity there are about 286,000

seeds per lb (630,500 per kg). Acceptable seed purity is

95 percent and germination is 50 percent. Success

from direct seeding can be erratic (Plummer and

others 1968). Germination requires wet prechilling;

therefore, fall seeding is recommended. Seed should

be covered lightly and planted in areas where competition will be light for a few years. Red elderberry can

be propagated by cuttings, either hardwood cuttings

started in winter or softwood cuttings taken during

spring and summer (Doran 1957; Ritter and McKee

1964).

Most natural reproduction occurs vegetatively from

buds on stems, rhizomes, and root crowns following

fire or mechanical damage (Conrad and McDonough

1972; Van Dersal 1938). Some reproduction also occurs from seed. Seeds are dispersed by birds and

rodents and can remain dormant in the soil for long

periods. Seeds scarified by fire often germinate in

large numbers during the first season following burning (Heit 1967a; Hungerford 1984; Kramer 1984).

Uses and Management—Red elderberry is used

seasonally by sheep, cattle, deer, bear, porcupine,

and elk (Conrad and McDonough 1972; Dayton 1931;

Kufeld 1973; Kufeld and others 1973; Martin and

others 1951; Ritter and McKee 1964; Smith 1953;

Steele and Geier-Hayes 1987; Zager 1980). It is one of

the most palatable browse species for elk in Idaho and

Montana (Gaffney 1941; Young and Robinette 1939;

Zager 1980). Palatability is excellent for sheep following frost.

All top growth is consumed or dies to ground level

each fall. Fruits mature in late August and September. They are consumed by birds, rodents, livestock,

and big game. Seed is cached and used by rodents.

During winter porcupines and mice feed on the buds

and bark (Conrad and McDonough 1972). Red elderberry provides valuable nesting and perching habitat

and food in the form of berries and buds for a large

number of birds (Denslow 1987; Gullion 1964; Martin

and others 1951; Van Dersal 1938).

Red elderberry is used to stabilize disturbed and

erosion-prone areas, especially on aspen and upper

spruce-fir sites, riparian areas, seeps, and other moist

sites (Platts and others 1987; Worley and Nixon 1974).

Bareroot stock and container-grown stock have been

used successfully for planting disturbed riparian areas in aspen and ponderosa pine forests (Platt and

others 1987). Red elderberry is used as a colorful

Figure 15—Red elderberry frequently occurs in forested openings and subalpine communities.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 627

Chapter 23 Shrubs of Other Families

ornamental (Kelly 1970; Ritter and McKee 1964), and

should be considered when planting recreational sites,

summer homes, and ski areas.

Varieties and Ecotypes—None. There is, however, considerable variation within the species.

Family Caprifoliaceae ____________

Symphoricarpos albus

Common or white snowberry

Description—Common snowberry is an erect, multibranched shrub usually 3.3 to 6.6 ft (1 to 2 m) tall, but

ranging from 1.6 to 9.8 ft (0.5 to 3 m) in height (fig. 16).

Plants are fibrous rooted with the densely branched

root system concentrated near the soil surface. The

species is slightly rhizomatous with sprouts from the

thick, spreading rhizomes forming dense thickets.

The slender upright stems are hollow, smooth, and

brownish. Twigs are glabrous and obscurely puberulent. Leaves are opposite, thin, pale green, elliptic to

elliptic ovate, and entire or with a few coarse, irregular

teeth. They are 0.6 to 2 inches (1.5 to 5 cm) long and 0.4

to 1.4 inches (1 to 3.5 cm) wide with petioles 0.1 to 0.2

inch (2 to 4 mm) long. Leaves on sterile shoots may be

somewhat larger. Racemes are terminal or sometimes

in the upper leaf axils. They are dense, subsessile and

few flowered. The perfect flowers are small, pinkish,

and bell shaped. The five-parted corolla is 0.2 to 0.3

inch (5 to 7 mm) long and nearly as wide. The interior

is densely hairy. Corolla lobes range in length from

one-half to nearly equaling the tube length. Anthers

alternate with the petals and nearly equal the filaments in length. There is a single style. The ovary is

four locular. Generally two locules produce a single

mature seed each. The white pulpy fruits are subglobose

or ellipsoid and 0.3 and 0.6 inch (8 to 15 mm) in

diameter. Seeds are elliptical, flattened, creamy white,

smooth, and 0.16 to 0.2 inch (4 to 5 mm) in length with

fleshy endosperm and a tiny, poorly developed embryo

(Evans 1974; Hitchcock and others 1959; Hopkins and

Rawlings 1985; Kovalchik and others 1988; Marchant

and Sherlock 1984; Viereck and Little 1972; Welsh

and others 1987).

Ecological Relationships and Distribution—

Common snowberry occur across the United States

and Canada except in the far north of Alaska and

Canada and the Southern States. It naturally occurs

associated with conifer species, in shade and in openings with full sunlight.

Plant Culture—Common snowberry seeds are white

and average 76,000 per lb (167,500/kg). One hundred lb

(45 kg) of fresh fruit will yield 3 to 5 lb (1.4 to 2.3 kg) of

cleaned seed. Good seed crops are produced in 4 out of

5 years. Flowering occurs from June through July.

Fruits remain on the plant into winter or until eaten.

Fruits are processed through a Dybvig macerator, dried,

and run across an air screen separator. Unfilled seeds

are removed by flotation. Germination is delayed by

dormancy, which can be overcome with wet prechilling

or scarification with sulfuric acid. Germination will

occur over a number of years. Young plants grow slowly;

however, they can mature and flower in 3 or 4 years.

Uses and Management—Common snowberry is

an important forage plant in the West, especially for

sheep and deer, which make considerable use of it year

around. Elk strip the leaves in the fall. Plants can be

lost as a result of excessive use.

This species has been used extensively as an ornamental. It can be directly seeded or transplanted in

restoration projects. It does not do well on disturbed

sites such as mines and road fills and cuts.

Varieties and Ecotypes—There are a number of

ornamental varieties and selections, but none are

available for use on rangelands and disturbed sites.

Family Caprifoliaceae ____________

Symphoricarpos longiflorus

Desert snowberry

Description—Desert snowberry is a native, longlived, deciduous shrub. Plants are low growing, erect

to arching, and range in height from 1.6 to 5 ft (0.5 to

1.5 m). The ability of the species to spread by rhizomes

is poorly known. Plants exhibit wide variation in leaf

and twig pubescence. Young stems are persistent,

glabrous to pubescent, and tend to spread at right

Figure 16—Open, branching, scattered, pale-green

leaves, and shedding bark are distinctive characteristics of common snowberry.
628 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 17—Western snowberry with ovate leaves

and small white flowers is widespread in aspen and

conifer forests where it often occurs in dense thickets.

angles, giving the plant a thorny appearance. The firm

leaves are simple and opposite, 0.2 to 0.6 inch (6 to 15

mm) long and 0.1 to 0.3 inch (2 to 7 mm) wide, oval to

lanceolate or oblanceolate, and entire or with one or

two teeth. The fragrant flowers are solitary, paired in

the leaf axils, or in small, few-flowered, terminal

racemes. The calyx and corolla are five lobed and

regular. The slender corolla tubes are 0.4 to 0.7 inch

(1.0 to 1.8 cm) long, salverform, glabrous, and pale to

deep pink. Lobes are much shorter than the tube and

rather abruptly spreading. Anthers are sessile, alternating with the corolla lobes. The single style is stiff

and hairy above the center. The ovary is four loculed;

the two functional locules each contain a single seed.

The fruit is a white, waxy, two-seeded berry about 0.3

to 0.4 inch (8 to 10 mm) in diameter. Seeds are 0.2 inch

(5 mm) long and acute at the base (Cronquist and

others 1984; Davis 1952; Evans 1974; Welsh and

others 1987).

Ecological Relationships and Distribution—

Desert snowberry occurs on deep, well-drained soils in

desert-shrub, pinyon-juniper, sagebrush, mountain

bush and ponderosa pine communities from Oregon to

Colorado, and south from southern California to Texas.

It is the most drought tolerant of the snowberries. It

grows intermixed with other species, but not in closed

stands.

Plant Culture—Desert snowberry produces white,

two-seeded berries. There are an average of 70,000

seeds per lb (154,000/kg). One hundred lb (45 kg) of

fresh fruit will yield 5 to 7 lb (2.3 to 3.2 kg) of cleaned

seed. Good seed crops are produced in about 2 out of 5

years, depending on spring and summer storms. Flowering occurs in June to mid July. Berries are harvested

by hand stripping or beating. Fruits are extracted by

maceration through a Dybvig cleaner. Unfilled seeds

are removed by flotation. Germination can be erratic.

Dormancy can be overcome with wet prechilling and

scarification with sulfuric acid. Fall seeding will permit prechilling to occur. Young plants can mature and

produce seed in 3 to 4 years and reach full stature in

5 to 6 years.

Uses and Management—Desert snowberry can be

an important browse plant for sheep and deer, and to

a lesser extent for cattle. It is fairly resistant to

grazing. On many overgrazed sagebrush ranges in

which the understory species have been eliminated,

desert snowberry and sagebrush will remain. Excessive use will result in death of plants.

This species can be seeded or transplanted. It does

not do well on severely disturbed sites. It does best on

soils with well-developed, undisturbed horizons.

Varieties and Ecotypes—None.

Family Caprifoliaceae ____________

Symphoricarpos occidentalis

Western snowberry, wolfberry

Description—Western snowberry is an erect shrub,

branching weakly from a woody base and short main

branches. Plants are 1 to 3.3 ft (0.3 to 1 m) tall,

spreading freely from rhizomes, and often forming

dense thickets that exclude other vegetation. Roots

are densely branched, shallow, and intermixed with

stout creeping rhizomes. Young twigs are puberulent

to glabrous. Petioles are 0.1 to 0.4 inch (3 to 10 mm)

long. The opposite leaves (fig. 17) are thick, elliptic to

ovate, entire or with a few, coarse, blunt, irregular

teeth. Most are 0.8 to 3.1 inches (2 to 8 cm) long and 0.4

to 2 inches (1 to 5 cm) wide with somewhat revolute

margins. Leaves on sterile shoots are often larger,

frequently more lobed, glabrous above, and usually

hirsute puberulent beneath, at least along the main

veins. Few to many-flowered spikelike racemes are

produced at the ends of the twigs and in the upper leaf

axils. The five-merous corollas are often wider than

long, pink to rose colored, and densely hairy within.

Lobes are arcuate spreading. Anthers alternate with

the petals and are shorter than the filaments. The

style may be hairy near the middle, but is occasionally

glabrous. The ovary is four loculed; two locules usually

produce one mature seed each. Fruits are subglobose

and greenish white, turning black after frost. The two

seeds are white, somewhat ellipsoid, and flat on one

side with a poorly developed embryo (Cronquist and

others 1984; Davis 1952; Hansen and others 1988b;

Hitchcock and others 1959; Wasser 1982; Welsh and

others 1987).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 629

Chapter 23 Shrubs of Other Families

Ecological Relationships and Distribution—

Western snowberry is widespread from British Columbia to Manitoba, south to Washington, Idaho, and

Colorado. In Utah it is most often associated with

riparian communities, mainly with cottonwood and

willow species (Welsh and others 1987). It does especially well on sites with high water tables. It is also

found in the mountain brush, open conifer sites, and

upper pinyon-Rocky Mountain juniper communities,

especially on east- and north-facing slopes. It competes well with understory herbivore species.

Plant Culture—Fruits of western snowberry are

collected by hand stripping. Seeds are extracted by

maceration through a Dybvig cleaner. There are about

74,000 seeds per lb (163,000 seed/kg). One hundred lb

(45 kg) of fresh fruits will normally yield 8 to 10 lb (3.6

to 4.5 kg) of seed. Good seed crops can be expected 3 out

of 5 years. Flowering occurs from the first of June

through late July. Fruits ripen from mid-September to

mid-October. Considerable seed dormancy can exist.

This can be overcome with moist prechilling and

scarification with sulfuric acid. Fall seeding will help

to overcome dormancy. Seedling emergence, however,

can be erratic over a few years. Plants can be established from nursery-grown container and bareroot

stock. Wilding stock can be obtained early in spring

while the ground is still moist and before plants break

dormancy.

Uses and Management—Western snowberry can

be used extensively by sheep and deer, and a little less

by cattle. Elk prefer it in the fall after it has been

frozen. It can receive heavy browsing and trampling in

riparian areas. Excessive use can result in plant death.

This species has been used fairly extensively in restoration projects, especially as transplants.

Varieties and Ecotypes—None.

Family Caprifoliaceae ____________

Symphoricarpos oreophilus

Mountain snowberry

Description—Mountain snowberry is a spreading

to erect branching shrub 4.9 to 6.6 ft (1.5 to 2 m) tall

(fig. 18). Leaves and twigs are densely puberulent to

glabrous and sometimes glaucous. The thin opposite

leaves are elliptic to ovate, entire, or with a few teeth

or lobes. They range from 0.4 to 2 inches (1 to 5 cm) in

length and 0.1 to 1 inch (0.3 to 2.5 cm) in width. Leaves

on sterile shoots may be larger (Mozingo 1987). The

cream-colored to pinkish flowers have an unpleasant

odor. They are solitary or paired on short, drooping

pedicels in the upper leaf axils, or borne on short, fewflowered terminal racemes. The corolla and calyx are

five- or occasionally four-lobed. The corolla is longer

than wide, elongate campanulate to tubular funnelform. The corolla lobes are one-fourth to one-half as

long as the tubular portion of the corolla. The interior

of the tube is sometimes hairy below the level of

filament insertion, but may be glabrous. The filaments are equal to or shorter than the anthers. The

ovary is four-locular. Two locules normally contain

several abortive ovules each; the remaining two locules each contain one normal pendulous ovule. Fruits

are waxy, white ellipsoid, and about 0.3 to 0.4 inch

(7 to 10 mm) long. The hard nutlets or seeds are 0.2 to

0.3 inch (4 to 6.5 mm) long (Cronquist and others 1984;

Hitchcock and others 1959; Welsh and others 1987).

Ecological Relationships and Distribution—

Mountain snowberry is common in midelevation sagebrush, ponderosa pine, Gambel oak, aspen, Douglasfir, lodgepole pine, and spruce fir communities in the

Intermountain region. It can be found growing from

British Columbia to Montana, and south to California,

Arizona, and New Mexico (Welsh and others 1987).

Plants may be locally abundant, and in some cases

form nearly pure stands. Mountain snowberry will

dominate aspen stands. This shrub does well in full

sunlight and shade.

Plant Culture—Mountain snowberry seeds are

collected by hand stripping the two-seeded fruits from

the stems. Good seed crops are produced 3 out of 5

years. Seeds are cleaned in a Dybvig macerator. Unfilled seeds are removed by floatation. Germination is

delayed by the impermeable endocarp and an immature embryo. Pretreatments include a 3- to 4-month

wet prechilling retreatment. Scarification with sulfuric acid is also used to break the dormancy imposed by

the endocarp (Evans 1974). The embryo develops to

Figure 18—Mountain snowberry is commonly

encountered in open parks and conifer forests.
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Chapter 23 Shrubs of Other Families

Figure 19—Redosier dogwood with its distinctive

red bark, opposite leaves, and stoloniferous habit

grows in riparian areas and on other moist sites.

maturity during an extended period of wet

prechilling. There are about 75,000 seeds per lb

(165,000/kg). One hundred lb (45 kg) of fresh fruits

will yield 6 to 10 lb (2.7 to 4.5 kg) of seed. Flowering

occurs through June and July. Fruits mature from

mid-September through October.

Successful establishment may be achieved by fall

planting. Seeds may be hand planted or drilled. Seeds

should not be directly planted with grasses because

snowberry seedlings develop very slowly. Mature plants

will, however, develop in 3 to 4 years. Mountain

snowberry is easily established using nursery-grown

bareroot or container stock. Seedlings propagated in

the nursery or greenhouse grow relatively rapidly,

producing extensively branched root systems. Softwood cuttings taken during the period of flowering or

vegetative growth are easily rooted and transplanted

(Everett and others 1978a; Mirov and Kraebel 1939).

Uses and Management—This species does not

grow well on disturbed roadways, mine sites, or similar disturbances unless areas are top soiled. It has

been seeded and transplanted fairly extensively on

such sites with good success.

Mountain snowberry is low growing, and is accessible to all classes of livestock and wildlife. This shrub

is an important source of forage in the early spring, as

it is one of the first shrub species to leaf out. In areas

where snowberry is a dominant species, it often provides important forage throughout the spring and fall.

The persistent, fleshy fruits of all snowberry species

are taken by a variety of birds. Field mice scatter the

seeds, and plants are widely established from mice

caches.

Varieties and Ecotypes—None.

Family Cornaceae _______________

Cornus stolonifera

Redosier dogwood

Description—There are about 30 species of dogwood; most occur in the Temperate Zone of the Northern Hemisphere. Three species occur in the Rocky

Mountain region. Redosier dogwood, named for its red

to purplish bark is also known as creek dogwood. It is

a deciduous, short-lived, rapidly growing, thicket-forming shrub common to moist places (fig. 19). Its spreading clumps of slender stems rise to 15 ft (4.6 m) in

height and 20 ft (6 m) in width from a central crown

and spreading root system. Stem layers form on decumbent to prostrate stems that simulate stolons. The

degree of vegetative spread varies widely. Bark of

young branches is bright red to purplish, later turning

gray green. Lenticels are diamond shaped. Leaves are

petiolate, opposite, entire, elliptic-ovate to obovate

and usually acuminate, 1 to 4 inches (2.5 to 10 cm) long

and about two-thirds as wide. They are distinctively

pinnately veined and sometimes wrinkled, sparsely

strigillose and greenish on the upper surface, and

paler with pubescence along the veins and sometimes

covering the lower surface. Flowers are clustered in

flat-topped, ebracteate cymes. Penduncles and

branches are strigose to conspicuously spreading pubescent. Flowers consist of four very small sepals, four

white petals, 0.1 to 0.2 inches (2 to 4 mm) in length,

four stamens, and a two-carpellate pistil. The white to

bluish drupes are berrylike, glabrous to pubescent,

and 0.28 to 0.35 inches (7 to 9 mm) in diameter. Flesh

is thin and succulent or mealy. There are usually one

or sometimes two bony stones. A smooth rather than

grooved stone is a major feature separating C. s. var.

stolonifera from C. s. var. occidentalis, although they

are widely hybridized (Hitchcock and others 1961).

Redosier dogwood flowers from May to July, depending

on location. There may be a second period of flowering

later in the summer. Fruits ripen from July to September and remain on the plant for 8 or 10 weeks (Olsen

and Nagle 1965; Orme and Leege 1980; Stark 1966).

Ecological Relationships and Distribution—

Redosier dogwood is widely distributed across North

America, occurring from Alaska to Newfoundland

and south from California to the Northeastern States.

The species also occurs in central Mexico. It grows at

elevations from 1,500 to 10,000 ft (460 to 3,030 m)

(Harrington 1964; Thornburg 1982) on moist sites,

usually along streams, swamps, low meadows, river

and creek banks, fields, and woods in areas receiving

more than 19 inches (48 cm) of annual precipitation.
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Chapter 23 Shrubs of Other Families

It is one of the earliest shrubby plants to invade bogs

and swamps due to the ability of its roots to live

immersed in water (Conway 1949). It is frequently

associated with stands of alder, birch, and willow, and

in moderately moist areas, with mountain maple,

blackberries, or wild roses. Large plants grow singly in

grasslands, incapable of spreading by layering due to

the dense ground cover. Redosier dogwood, highly

adaptable to a range of soil types, is found on sphagnum mats with a pH of 3.2 (Jewel and Brown 1929) and

alkaline soils of pH 8 (Van Dersal 1938). It grows on

soils with textures ranging from silty to clayey. In

Wisconsin it is recommended for planting on soils with

a pH of 5.0 to 6.0, base exchange capacity of 6.0 me per

100 g, and total nitrogen of at least 0.7 percent (Wilde

1946). Roots of redosier dogwood may be associated

with vesicular arbuscular mycorrhiza (Barnhill 1981).

Redosier dogwood will grow in full sun or in shaded

areas. In full sun, the plants are dense and compact

with many lateral branches. In shade, the growth

habit is more open and sprawling with few axillary

branches, larger leaves, thinner and less curled blades,

and lower leaf/total weight ratios. Shrubs growing in

full sun tend to develop richer purple coloring in the

fall (Sheppard and Pellitt 1976).

Use of redosier dogwood in landscaping, along highways, and on mine spoils has led to a number of

physiological studies to determine the effects of short

days, water stress, defoliation, and low temperature in

initiating and maintaining various degrees of frost

hardiness (Chen and Li 1978).

Lumis and others (1973) found that of 20 deciduous

shrubs selected from those growing along highways in

Ontario, redosier dogwood exhibited the greatest sensitivity to aerial drift of deicing salt because its buds

are exposed in winter. Affected trees exhibited twig

dieback, a tufted growth pattern, and a greater susceptibility of flowering buds than vegetative buds.

Heale and Ormrod (1982) evaluated plants used for

revegetation of toxic wastes high in nickel and copper

at primary base metal smelters and found redosier

dogwood exhibited typical symptoms. It was sensitive

to both levels of nickel tested (2 and 10 mg per 1), but

was sensitive only to the highest copper level tested

(20 mg per 1, but not 4 mg per 1).

Uses and Management—Redosier dogwood seedlings establish and grow rapidly on wet sites, making

it a useful species for streambank, erosion control, and

windbreak plantings. It can withstand moderate browsing, but repeated heavy browsing leads to plant loss

(Swihart and Yahner 1983). New plantings begin

producing fruit in 3 to 5 years.

Berries of redosier dogwood provide nutrients for

pheasants, grouse, and many other birds in early winter (Stokes 1977). In New England redosier dogwood

seeds were found in the diets of 95 bird species (McKenny

1939). Beaver, bears, and mountain goats use the fruit,

wood, and foliage (Murie 1951; Rue 1964). Rabbits,

moose, deer, and elk take twigs and exposed winter

buds. Plants on winter ranges sometimes suffer from

excessive browsing. Kufeld (1973) rated the forage

value of redosier dogwood as valuable in winter and fall

and highly valuable in summer. It receives moderate

fall and winter use and heavy summer use from mule

deer (Kufeld and others 1973). The species provides

dense summer cover and partial winter cover for birds

and other small animals.

Redosier dogwood is recommended for streambank

plantings to stabilize eroding banks and provide shade.

It may be used to provide an attractive row in windbreak and conservation plantings in areas with adequate water availability (Cook 1981; Olson and Nagle

1965; Stokes 1977). Varieties selected for ornamental

use generally have a more compact growth form. In

landscapes it provides obvious color and a graceful

growth form during all seasons (Stokes 1977; Sutton

and Johnson 1974).

Plant Culture—Fruits are collected as soon as

they are ripe to reduce losses to birds. Seeds should

be fully mature when harvested; some planting

failures have been attributed to use of immature seeds

(Smithberg 1974). Embryo dormancy is overcome by a

60- to 90-day wet prechilling at 35 to 41 ∞F (2 to 5 ∞C).

A tetrazolium chloride or embryo excision test is normally used to indicate seed quality and preclude the

lengthy wet prechilling requirement (Heit 1955;

Brinkmann 1974d). The double-celled fruits sometimes contain two viable embryos, which may contribute to high germination percentages.

Field seeding is best accomplished by spot seeding

untreated seed in fall or wet prechilled seed in spring

on favorable microsites. Seedlings must be protected

from competition and browsing. Peterson (1953) found

that laboratory wet prechilling produced better greenhouse germination than wet prechilling through exposure to winter conditions, or combinations of each type

of wet prechilling with scarification. Laboratory wet

prechilling also provided the greatest resistance to

damping off.

Nursery seeding is accomplished by seeding freshly

collected fruits. Dry stored fruits should be soaked in

water prior to planting (Heit 1968a). Seeds may also be

wet prechilled in the laboratory for spring planting.

Seed should be drilled at a density of 40 viable seeds

per ft2 (430 per m2

). Seedlings grow rapidly under

favorable conditions. One- or two-year-old bareroot

stock may be used for outplantings (Shaw 1984).

Plants may also be grown as containerized seedlings

(Landis and Simonich 1984).

Redosier dogwood reproduces vegetatively from stolons or layering (Peterson 1953; Smithberg 1974).

Layering may be induced by pressing branches to the
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Chapter 23 Shrubs of Other Families

Figure 20—A 10-year-old planting of Arizona cypress provides wildlife cover in south-central Idaho.

ground with hooks and covering them with loose soil.

Rooted shoots are later severed, dug, and transplanted.

Layers, as well as hardwood or softwood cuttings, may

be used for vegetative propagation. Dormant cuttings

may be rooted immediately in the field in moist areas,

or rooted in a nursery or greenhouse prior to field

planting (Doran 1957). Planting stock is generally

available commercially, but origin should be matched

as closely as possible to the planting site.

Varieties and Ecotypes—None.

Family Cupressaceae ____________

Cupressus arizonica

Arizona cypress

Description—Arizona cypress is an erect, somewhat pyramidal, evergreen tree with fragrant, resinous foliage (Johnson 1974c; Welsh and others 1987).

Five different varieties were described by Johnson

(1974c); these differ somewhat in erectness and

crown spread (Sudworth 1915; Wolf and Wagener

1948). Trees are mainly 15 to 70 ft (5 to 21 m) in

height (Johnson 1974c; Welsh and others 1987).

Trunks are 5 to 20 inches (13 to 51 cm) thick; twigs

are stout, branching at nearly right angles; the bark

is scaly or furrowed and usually grayish. Leaves are

scalelike, sharply pointed, blue green, and 0.1 inch

(2 mm) long. Cones are globose, 0.6 to 1 inch (1.5 to

2.5 cm) thick, short-stalked, hard and woody, with six

to eight flattened scales bearing a hard point in the

center. Seeds are 0.1 inch (2 mm) long and purplish

brown (Welsh and others 1987). Plants are monecious.

Staminate and ovulate strobili are produced on the

ends of the branches. Staminate strobili are long and

cylindrical, turning yellow as pollen ripens (Johnson

1974c). Ovulate strobili are small, usually less than

0.3 inch (6 mm) long, and green, with six to 12

distinctly arranged scales (Johnson 1974c).

Ecological Relationships and Distribution—

Arizona cypress is distributed from Texas to Arizona

and Mexico (Welsh and others 1987). The varieties

described by Johnson (1974c) occur in somewhat different locations, mostly in north-central Arizona, southern California, and Mexico. This plant has commonly

been grown as an ornamental and Christmas tree

throughout much of the West. It has proven adaptable

to areas outside its native range, and has been planted

on numerous sites including many range and wildland

communities (fig. 20). It is best adapted to sites receiving at least 14 to 16 inches (36 to 41 cm) of annual

precipitation. Normal occurrence is on north slopes,

coves, benches, and canyon bottoms (Sudworth 1915),

particularly moist, sheltered canyon bottoms.

Plant Culture—Throughout its natural range,

cones develop in late fall and pollen is shed in October

and November (Johnson 1974c); seeds mature 15 to 18

months later. At the time of maturity cones are about

1.2 inches (3 cm) diameter. Because cones require

approximately 2 years to mature, numerous cones of

different ages are borne on a single tree. Some trees

produce abundant female cones; other trees are much

less productive. Trees less than 10 years of age normally do not produce many cones. Insects often damage the cones; consequently, seeds should be carefully

inspected to assure healthy cones and viable seeds are

collected. Ripe cones are dark brown or deep purplish

brown (Sudworth 1915; Wolf and Wagener 1948).

Seeds are harvested by handpicking or beating the

tree to dislodge the hard woody cones onto canvas

tarps. Trees can be damaged from excessive flailing.

Cones are normally harvested in early to late fall. The

cones are serotinous; they open when mature to shed

the seeds. The scales surrounding the cones form a

tight cluster that can be difficult to open unless quite

dry. Drying the seeds normally causes natural shattering in 1 to 2 months. Johnson (1974c) suggested

storing the cones for several days in a refrigerator to

aid in drying and to prevent cone hardening. Seeds

normally separate freely from the cones. Approximately 55,050 cleaned seeds per lb (121,400 per kg)

were reported by Johnson (1974c) for collections of

C. a. variety glabra. Between 90 and 100 seeds are

produced per cone. Considerably larger numbers are

reported for other varieties (Goggans and Posey 1968;

Johnson 1974c; Wolf and Wagener 1948). Seeds are

flattened or lens shaped with short wings (Johnson

1974c).

Seeds wet prechilled for 30 days at 34 ∞F (1.1 ∞C) will

germinate readily at incubation temperatures near
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Chapter 23 Shrubs of Other Families

70 ∞F (21.1 ∞C) (Johnson 1974c), but most authors

recommend incubation at alternating temperatures of

86 ∞F (30.0 ∞C) day, and 68 ∞F (20.0 ∞C) night (International Seed Testing Association 1966; Stein 1966;

USDA Forest Service 1948). Seeds may be infested

with mold or bacteria that reduce germination. Treatment with Captan (N-[(trichloromethyl) thio]-4-

cyclohexene-1,2-dicarborimide) or other fungicides will

retard fungal development (Johnson 1974c). Because

young seedlings are also susceptible to damping off

fungi, seedbeds or nursery-grown containers can also

be pretreated with a fungicide.

Arizona cypress has been established by fall seeding

in a well-prepared seedbed. Seedlings are sensitive to

herbaceous competition, and require a weed-free seedbed for 1 to 2 years. Established seedlings are vigorous

and grow rapidly when well protected. Plants may

attain heights of 2.5 to 4 ft (0.8 to 1.2 m) in 2 to 3 years.

They continue to grow rapidly, but may not reach

mature stature for 40 years.

Seedlings and transplant stock are easily grown,

and the plant is sold by many commercial nurseries.

Small stock 6 to 10 inches (15 to 25 cm) high and large

ball and burlap trees are usually available.

Uses and Management—Arizona cypress is an

important resource in its native range. It provides

lumber, wildlife habitat, and watershed protection

(Johnson 1974c). This tree has been particularly useful for landscape and erosion control plantings in

southern California on a range of soil types at elevations below 6,000 ft (1,800 m) (Horton 1949). It is well

suited as a ground cover or landscape plant because it

grows rapidly, is long-lived, and has an attractive

pyramidal growth form (Horton 1949).

Within the Intermountain States, this tree has been

used for conservation and wildlife habitat improvement projects. It has been successfully used as a

windbreak or cover plant to protect structures and

shelter range and farmlands. It has been used to

provide cover in mixed plantings with other trees and

shrubs. Being evergreen, it is particularly important

as winter cover. It provides a dense amount of foliage

from the base to the top of the plant. It has survived

wind and heavy snowstorms, but some individuals

and populations lack cold tolerance to harsh conditions in mountainous regions in the West.

This tree species has been useful for wildlife habitat

improvement in Utah (Plummer and others 1968). It

establishes and grows well, particularly from transplanting in mountain brush, pinyon-juniper, and big

sagebrush communities. Seed germination is somewhat erratic, and irregular stands have established

from direct seedings. Natural spread by seedling

recruitment is poor from all plantings within the

Intermountain area.

This species has received considerable browsing by

big game at all planting sites in the Intermountain

area. Game animals use the plant heavily in summer,

and in lesser amounts in the spring, fall, and winter.

The foliage is quite palatable, and heavy browsing and

highlining of the trees normally occurs. Heavy browsing can prune or eliminate leaf and small stem growth

from the lower portion of the plant.

Arizona cypress has demonstrated mixed success on

mine sites and other disturbances. It is not adapted to

infertile mine wastes unless top soiled. It does not

serve well as a pioneer species on barren disturbances.

Young plants succumb quickly if planted on marginally adapted sites.

Varieties and Ecotypes—None.

Family Cupressaceae ____________

Juniperus horizontalis

Creeping juniper

Description—Creeping juniper is a native, decumbent to procumbent or prostrate shrub that normally

attains heights of 8 to 12 inches (20 to 31 cm) (Bifoss

1947; Great Plains Flora Association 1986; Welsh and

others 1987) (fig. 21). Plants develop in clumps or mats

that may exceed 23 ft (7.0 m) in diameter (Hitchcock

and others 1969; Stephens 1973). The decumbent or

prostrate stems root at the nodes forming well-rooted

clumps. Leaves are mostly 0.04 to 0.16 inch (1 to 4 mm)

long, decurrent, opposite, scalelike or awl shaped, and

acute to spinulose tipped. Cones mature the first

season; they are green when immature and ripen to

blue purple or blue black. They are glaucous, 0.2 to

Figure 21—Creeping juniper growing as an understory with aspen.
634 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

0.4 inch (5 to 10 mm) thick, and mostly three- to fiveseeded (Welsh and others 1987). Seeds are reddish

brown and ovoid to spherical (Hitchcock and others

1969). Plants are dioecious, and although mature

cones may be produced in one season (Welsh and

others 1987), 2 years are normally required (Great

Plains Flora Association 1986; Miller 1978).

Ecological Relationships and Distribution—

Creeping juniper is widely distributed in North America

(Little 1979). It occurs from Montana to northern

Colorado and Utah, and east to northeastern Iowa,

Nebraska, Illinois, and New York (Little 1979). It is

abundant throughout Canada (Couchman and Von

Rudloff 1965), and occurs from Alaska and the Yukon,

east to the Atlantic, and south to British Columbia

(Welsh and others 1987). The shrub is widely distributed

over a number of different plant communities. It

grows in association with grass and broadleaf forbs

at low elevations in the northern Great Plains (Girard

1985; Hansen and others 1984; Johnston 1987). It is

also associated with conifer forests, aspen, and limber

pine (Girard 1985) at high elevations and on more

moist sites (fig. 21).

Plant Culture—Staminate cones are produced at

the tips of the branches in early spring. Pollination of

pistillate cones normally occurs in April and May

(Miller 1978). Pistillate cones form globose, berrylike

fruits (Great Plains Flora Association 1986; Hitchcock

and others 1969; Miller 1978). As they ripen they

change in color from green to blue black or bluish

purple (Stephens 1973). Cones normally remain on

the plant until the late fall or early winter of the second

year. When fully mature they drop from the plant.

Seeds are harvested by stripping or picking the

ripened cones. Cones can also be dislodged by flailing

the branches and dislodging them onto canvases spread

on the ground. It is advisable to collect the cones as

soon as they ripen. Collecting green cones or immature

seeds is not advised, as they are difficult to separate

from fully ripened seeds during cleaning. Freshly

collected cones should be spread to dry, but can be

stored and seeded as dry cones or as cleaned seeds

(Johnsen and Alexander 1974). Seeds are normally

extracted from the cones by maceration using a Dybvig

seed cleaner. The debris is floated away in water

(Johnsen 1959; Johnsen and Alexander 1974). If cones

are completely dry before cleaning, they may require

soaking for a few days to aid in separation. The

macerated material is dried and then fanned to separate seeds from the trash. Seeds should be stored in

sealed containers at 20 to 40 ∞F (–6.7 to 4.4 ∞C) with

relative humidity at about 10 percent (Johnsen and

Alexander 1974; Jones 1962).

Germination of creeping juniper seed is delayed by

embryo dormancy and an impermeable seedcoat.

Seeds also require a period of dry afterripening (Pack

1921). To enhance germination, seeds of most junipers are subjected to various pretreatments including

wet prechilling at 40 ∞F (4.4 ∞C) for 30 to 120 days

(Barton 1951; Johnsen and Alexander 1974).

When planted in the fall, seed dormancy can usually

be broken by overwinter wet prechilling. Seedbeds

should be covered with mulch to ensure the soil remains moist during the time of germination (Johnsen

and Alexander 1974). Seeds are normally covered to a

depth of at least 0.25 inch (6.4 mm) in a firm seedbed

(Stoeckeler and Slabaugh 1965). Seedbeds are frequently shaded during the first growing season to

protect the soil surface from rapid drying and reduce

the chance of damage from early spring frosts (USDA

Forest Service 1948).

Creeping juniper can be propagated from stem cuttings. Portions of the stem that contact the soil and

form roots can be cut and separated into individual

plants. The stem cutting and excised root system can

be planted in containers, grown to a suitable size, and

field planted. Stem cuttings can also be rooted easily

under greenhouse conditions (Zorg 1954). Stem cuttings taken in early winter (November and December)

root readily without the use of rooting compounds

(Doran 1957; Kiplinger 1938).

Plantings of 2-year-old bareroot stock have established and survived better than younger, smaller stock.

Field lifting and transplanting must be completed in

early spring when plants are dormant. Although transplanting may be done later in the season, supplemental irrigation is usually required to assure survival.

Uses and Management—Creeping juniper has

not been used extensively in large revegetation projects.

Although recognized as an important ground cover

and a locally useful forage plant for wildlife, it has

been difficult to establish on wildland sites. This

species has been able to persist following heavy grazing when more palatable shrubs and herbs have been

eliminated. Consequently, rehabilitation programs

often do not include this shrub. Creeping juniper

normally occurs as scattered plants, occupying ridge

crests, rocky slopes, ledges, washes, or areas that are

difficult to plant with conventional equipment. Although site accessibility limits planting in some areas,

this plant is important for ground cover and wildlife

habitat and should not be overlooked.

Creeping juniper provides dense cover on harsh

sites. Its low evergreen growth habit provides diversity and cover throughout the entire year. Ecotypes

that are semierect or upright furnish better habitat for

wildlife. This shrub grows on sites where few other

plants are adapted. Because creeping juniper grows on

isolated wildlife ranges, it is important to maintain its

presence. It occurs in most plant communities and big

game winter ranges on sites where soil protection and
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Chapter 23 Shrubs of Other Families

forage diversity are critical for protecting watershed

and wildlife values. Consequently, it is important that

techniques are developed to better accommodate its

propagation and field planting.

Creeping juniper normally receives only limited use

by big game, but locally heavy winter browsing has

been reported (Frischknecht 1975; Miller 1978). Dusek

(1975) found that creeping juniper provided a major

portion of mule deer diets during March and April in

north-central Montana. This plant provides important fall, winter, and spring forage for big game (Dusek

1971; Kufeld and others 1973; Lovaas 1958; Martinka

1968; McKean 1954; Schallenberger 1966). It is more

commonly used by deer, but Dirschl (1963) found that

creeping juniper provides about 10 percent of the

winter diet of pronghorns, and was the dominant

forage from December to March in some parts of

Saskatchewan. Some ecotypes in the Intermountain

States exhibit considerable differences in use by game

animals. Selections that receive moderate or heavy

use are consistently browsed at these levels, although

climatic, edaphic, or other factors may contribute to

selective use by game.

Birds and other wildlife seek sites occupied by

this shrub. The cones provide food for many game

birds and mammals. They remain on the shrub for 1 or

2 years and provide 20 to 40 percent of the total food

consumed by sharptail grouse (Miller 1978). Not all

plants produce heavy seed crops. Annual production is

often quite low, but fruits that do develop are used

during winter and early spring periods when other

foods are less available.

Creeping juniper begins growth very early in spring.

Game animals are attracted to the shrub in late

winter and early spring when other forages are less

palatable. The plant grows in areas where game may

be forced to concentrate during periods of deep snow

and adverse weather conditions. At these critical

periods, it provides protection, cover, and forage that

are crucial for the survival of game animals.

Creeping juniper has been used to restore disturbed

watersheds and mine sites. It is extremely difficult to

establish by direct seeding because of germination

problems. Transplanting has been the most successful

means of establishment; however, costs associated

with transplanting limit its usefulness. Transplants

are difficult to establish on semiarid sites. Small

transplants root slowly and require frequent irrigation to fully establish. Thus, it has been difficult to

plant sites that do not remain moist through the year

of planting. Once established and well rooted, this

species survives very well. New plantings are not able

to compete well with broadleaf herbs and grasses, and

the shrub should be planted alone.

This shrub occupies areas in Wyoming, Montana,

and Colorado where considerable surface mining

occurs. Use of this shrub has been encouraged where

efforts are made to restore native communities. Creeping juniper has established slowly on mine spoils, and

appears sensitive to soil conditions. Although it naturally occurs on infertile sites, it is not well suited to all

mine wastes and spoil materials. It does best on sites

that have been top soiled or contain soil fines capable

of holding soil moisture. It has been less successful on

rocky, coarse mine wastes and exposed substrata. To

be successful on mine disturbances, a balanced amount

of soil nutrients must be provided.

This shrub can be used to restore mine spoils where

irregular topography is created. If topsoil or surface

soil material is reapplied, the plant can be successfully

planted to re-create natural communities and landscapes-disturbed areas to match adjacent native

plant associations.

Creeping juniper can also be used to stabilize watersheds and erodible sites. It establishes and grows

slowly, a major disadvantage in stabilizing erosive

soils and steep slopes. However, once established, it is

able to withstand windy conditions, and provides

excellent ground cover under very exposed and harsh

situations. This shrub is also very useful as a landscape species. It is widely used in horticultural

plantings in areas including recreational sites, campgrounds, parks, and associated sites where little maintenance is provided. It is particularly useful in planting parks and recreational sites where use of native

species is emphasized. It can withstand heavy traffic,

and is used to align walkways, roadways, and trails.

Varieties and Ecotypes—Various selections have

been developed for horticultural uses that exhibit

different colors, or vary in growth habit, leaf texture,

rooting habits, and growth rates. However, none have

been released for wildland situations.

Family Cupressaceae ____________

Juniperus osteosperma

Utah juniper

Description—Utah juniper is a small tree normally with a single stem and several short branches

that originate near the ground (fig. 22). Individual

trees may exceed 21 ft (6.4 m) in height (Goodrich and

Neese 1986). They have thin, fibrous bark that shreds

from the trunk and branches (Welsh and others 1987).

Welsh and others (1987) described the mature leaves

as typically opposite, scalelike, 0.04 to 0.12 inch (1 to

3 mm) long, and yellowish green. Juvenile leaves are

decurrent, awl-shaped, sharp, and 0.1 to 0.3 inch (2 to

8 mm) long. Plants are monoecious; staminate cones

are yellowish brown and 0.1 to 0.2 inch (3 to 4 mm)

long; ovulate cones are subglobose, 0.2 to 0.5 inch (6 to
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Chapter 23 Shrubs of Other Families

12 mm) thick and brownish or blue to blue purple at

maturity. There are one to two seeds per cone (Arnold

and others 1964; Johnsen and Alexander 1974; Tueller

and Clark 1975; Welsh and others 1987).

Ecological Relationships and Distribution—

Utah juniper occurs from Montana and Wyoming,

south to California, Arizona, and New Mexico (Griffin

and Critchfield 1972; Hitchcock and Cronquist 1973;

Welsh and others 1987; Zarn 1977). It grows with

singleleaf pinyon to form vast woodlands, typically

existing between more xeric cold desert shrub communities at lower elevations and mountain brush or

ponderosa pine at higher elevations. It normally occurs

between elevations of 6,000 to 7,500 ft (1,800 to 2,300 m)

(Goodrich and Neese 1986), although it is not uncommon between elevations of 2,790 to 8,000 ft (850 to

2,400 m) (Welsh and others 1987).

Utah juniper is the dominant species, along with a

number of shrubs and grasses, on many sites that have

been subjected to heavy domestic grazing (Clary 1975a)

and reduction of wildfires. These management practices have resulted in an increase of trees, coupled

with a decrease of understory plants. Pinyon-juniper

stands have not only increased in density, but have

spread to occupy adjacent shrub and grass communities (West and others 1979). The spread of Utah

juniper may be regulated by periods of drought and the

occurrence of years of favorable precipitation

(Blackburn and Tueller 1970).

Utah juniper is more tolerant of arid regions in the

Great Basin (Tausch and others 1981) than singleleaf

pinyon; it is the only tree species present in many

areas. It forms dominant stands throughout much of

southern Utah and Nevada. Treshow and Allan (1979)

concluded that the distribution of Utah juniper is

primarily regulated by precipitation. Tree stands normally occupy sites that receive from 10 to 25 inches

(254 to 635 cm) of annual rainfall (Blackburn and

Tueller 1970). Utah juniper is more tolerant of fire and

spring frost, and performs better with low summer

precipitation than singleleaf pinyon pine (Tueller and

others 1979; West and others 1979), yet these factors

do not fully explain the distribution and composition of

the tree species and communities. At higher elevations, singleleaf pinyon may dominate and is generally

more abundant.

Utah juniper is widely adapted to a variety of soil

types and parent materials. It grows on limestone,

sandstone, granite, and mixed materials (Clary and

Jameson 1981). Utah juniper is particularly well

adapted to rocky, well-drained, coarse-textured soils

(Eckert 1957).

Plant Culture—Good seed crops are normally produced annually by most wildland stands, although

abundant seed crops may only occur every 2 to 6 years.

Seeds require about 2 years to mature. Cones appear

from mid-April to mid-May, and normally mature in

late fall or early winter the second year after flowering

(Johnsen and Alexander 1974). Cones of different ages

are normally found on the same plant. Mature cones

are darker in color and are obviously drier than immature cones (Zarn 1977).

Most wildland stands produce commercially

harvestable crops, although selected individuals or

small groups of trees may be the primary producers.

Seed collection, cleaning, and storage practices are

similar to those described for creeping juniper and

discussed by Johnsen and Alexander (1974).

As with other junipers, the seed coats are impermeable and embryos are dormant (Johnsen 1959; Johnsen

and Alexander 1974). Seeds are difficult to germinate

without pretreatment. Johnsen and Alexander (1974)

recommended a 120-day warm pretreatment at 68 to

86 ∞F (20 to 30 ∞C) alternating diurnal temperatures

followed by wet prechilling for 120 days at 41 ∞F (5.0 ∞C).

Incubation at alternating day and night temperatures

of 68 to 86 ∞F (20.0 to 30.0 ∞C) for 70 days resulted in

less than 50 percent germination.

Although Utah juniper spreads well by natural

seeding, it is a difficult plant to propagate and it

establishes poorly by artificial seeding. Field plantings

establish erratically and unpredictably. Erratic seed

germination is the primary factor affecting poor seeding establishment. Pretreatments to stimulate and

regulate seed germination have been somewhat helpful when seeds are germinated under controlled laboratory conditions. However, pretreatment practices

have not significantly improved seedling emergence

from wildland plantings. Seedling establishment in

nurseries has been enhanced by prechilling seeds or by

Figure 22—Stands of Utah juniper furnish considerable cover for a variety of wildlife following

chaining and seeding with grasses and forbs.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 637

Chapter 23 Shrubs of Other Families

mulching and irrigating the seedbeds, but erratic

stands still result (Johnsen and Alexander 1974).

Seedlings of Utah juniper that emerge in early

spring and survive spring frosts are most likely to

survive their first season. Seedlings that germinate

late in spring, as soil moisture is depleted, are less

likely to survive. New seedlings grow slowly and

normally require 2 to 4 years to attain large stature.

During this period of establishment, mortality is very

high. Young plants that do survive for 1 to 4 years

become hardy, and can persist with considerable herbaceous competition and adverse climatic conditions.

Blackburn and Tueller (1970) concluded that spread of

juniper is favored by years of good seed production

followed by 6 years of average or above-average precipitation. Seedling establishment is difficult to predict. Numerous chaining and burning projects have

cleared areas and created openings where new seedlings may establish. Chaining and other activities

bury surface-deposited seeds, but new tree seedlings

do not always invade the treated sites. New tree

seedlings may not appear for many years after treatment, and increases may occur after understory species have established a dense cover.

Tree seedling survival is affected by the presence

and density of understory herbs. Monsen and others

(1987) found that native understory species, including

shrubs, have prevented tree seedling encroachment in

central Utah. Seeding introduced grasses and broadleaf herbs on pinyon-juniper sites reduces the establishment of tree seedlings, but the influence of individual herbaceous species is not fully understood. The

presence of native herbs also limits tree seedling

survival.

Tree recruitment is undoubtedly site influenced.

Natural recruitment frequently occurs very rapidly on

both treated and untreated sites where mature trees

have been removed by burning or chaining. On adjacent treated areas, tree recovery has been delayed for

extended periods even though all sites receive similar

amounts of precipitation. In addition, at some Utah

locations, Utah juniper stands cleared of mature trees

at different times, have little tree recruitment, but

other sites have experienced considerable tree recovery each year after treatment.

When direct seeded, Utah juniper should be planted

at a depth of 0.5 to 1.0 inch (1.3 to 2.5 cm) and sown

in late summer or fall (Heit 1967b; Johnsen and

Alexander 1974; Stoecker and Slabaugh 1965). Hand

seeding in small openings where the soil surface is

protected by litter, rock, or other debris tends to

improve survival.

Utah juniper is widely used in landscape plantings

as 2- to 5-year-old container transplants (Johnsen

and Alexander 1974). Bareroot transplants have also

been cultivated for field plantings. Survival of larger

2- to 3-year-old stock has exceeded that of smaller

transplants. Small transplants grow slowly and do not

compete as well as larger stock. Utah juniper transplants do not survive well if the primary root of young

plants is damaged. This impact is less evident for

larger stock.

Utah juniper can be propagated from stem cuttings

taken from November through February, but it is

difficult to root (Doran 1957; Snyder 1954). Treatment

with root-inducing substances may or may not be

beneficial (Doran 1957; Zorg 1954).

Uses and Management—Utah juniper is widely

regarded as an important plant for big game winter

ranges. The plant provides important cover and forage for wildlife (Frischknecht 1975). Nearly all big

game animals utilize pinyon-juniper woodlands.

This species is not regarded as highly palatable to big

game by most authors (Brotherson and Osayande

1980). Rosenstock and others (1989) found that mule

deer depended heavily on juniper during winter

months, and it is a principal part of their diet during

the midwinter period. The species may be regarded as

an emergency winter forage plant (Gullion 1964),

with only fair energy and protein values (Dittberner

and Olsen 1983), but Kufeld and others (1973) concluded that juniper is moderately to heavily grazed by

big game during most seasons. The fruits are widely

used by numerous birds (Turkowski and Watkins

1976). Big game and livestock graze the understory of

pinyon-juniper sites heavily in spring, fall, and winter.

Utah juniper provides essential cover for big game,

particularly during the winter months. Animals seek

cover and protection, and current chaining practices

are now being carefully evaluated to assure that adequate sized openings, escapeways, and concealment

cover are retained.

Game animals, livestock grazing, and recreational

uses of pinyon-juniper sites have increased dramatically in many locations. Game animals have been

forced to use pinyon-juniper areas in the spring, fall,

and winter, as access to and use of natural ranges have

been diminished by the construction of homes, roads,

farms, ranches, and commercial developments. Many

pinyon-juniper sites have been converted to shrub/

grass communities and managed to prevent tree recruitment. Pinyon and juniper have also spread to

occupy sites that are void of competitive understory

species. The encroachment has been facilitated by

the reduction or control of wildfires. Consequently,

pinyon-juniper sites have become increasingly important to the management of wildlife populations.

Chaining, burning, and seeding of pinyon-juniper

areas for livestock grazing may not be economically

practical in all situations, but restoration and improvement for wildlife habitat improvement is beneficial.
638 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Pinyon and juniper communities are an important

component of many watersheds. These sites may not

receive great amounts of precipitation each year, but

plants grow on steep slopes that are subjected to runoff

and erosion from high-intensity storms. Closed stands

of pinyon-juniper and communities that lack suitable

understory provide poor ground cover and are high

sediment producers. These sites should be managed to

prevent infrequent, but damaging erosion and runoff

from unpredictable storm events. Pinyon-juniper woodlands are also subjected to wildfires and other impacts

that can diminish tree cover. If the tree communities

lack a suitable understory, the burned or disturbed

sites become vulnerable to erosion; this significantly

impacts downstream resources. Consequently, it is

important to maintain a diverse understory of shrubs

and herbs that are capable of recovering following

these natural disturbances.

Pinyon and juniper communities have been invaded

by cheatgrass, even where closed tree stands exist. As

mature trees die from fires, insects, or disease, the

annual grass spreads quickly, and once in place prevents the recovery of nearly all native species. Extensive areas of pinyon-juniper woodlands throughout

the Intermountain region occur with only remnant

amounts of native shrubs and herbs, but with scattered and suppressed amount of cheatgrass. As these

sites eventually burn, the sites are converted to annual grass communities. The annual grass ranges

burn frequently, resulting in a loss of wildlife habitat.

To prevent the conversion of pinyon-juniper sites to

annual grass, sites can be artificially seeded with

desirable understory natives following burning or

other disturbances. In addition, potential problem

areas can be treated by controlled burns or chaining

followed by seeding. This practice has been extensively employed in Utah and other Intermountain

States. This problem has not been fully understood

and addressed in land management programs.

Deterioration of pinyon-juniper communities within

the Intermountain area has had a significant impact

on big game populations. Most pinyon-juniper sites

have been heavily impacted by previous grazing by

livestock and wildlife. This has resulted in a loss of

shrub and herb understory. Trees have increased and

once in place prevent the return of the desired understory. Consequently, extensive chaining, clearing, and

seeding projects have been employed to reduce tree

density and allow seeded species to establish. Numerous native shrubs, broadleaf herbs, and grasses have

been developed to convert tree-dominated sites into

sites dominated by a more diverse array of species.

Varieties and Ecotypes—None.

Family Cupressaceae ____________

Juniperus scopulorum

Rocky Mountain juniper

Description—Rocky Mountain juniper is typically

taller than Utah juniper, normally reaching 10 to

20 ft (3.0 to 6.1 m) in height (Welsh and others 1987),

but heights of 20 to 50 ft (6.1 to 15.2 m) are not

uncommon (Johnsen and Alexander 1974). Crowns

are conical to pyramidal or, less commonly, rounded.

The bark is thin, fibrous, and shredded. Leaves are

opposite, but sometimes in threes, scalelike, 0.02 to

1.2 inches (0.5 to 30 mm) long, green or blue green,

sometimes with dorsal resin glands. Juvenile leaves

are needlelike and 0.1 to 0.3 inch (3 to 8 mm) long

(Welsh and others 1987). Plants are generally monoecious with staminate cones 0.08 to 0.12 inch (2 to 3 mm)

long; female cones develop strobili or berrylike structures that are 0.2 to 0.3 inch (4 to 8 mm) thick at

maturity (Johnsen and Alexander 1974; Zarn 1977).

Arnow and others (1980) distinguished Rocky Mountain juniper from Utah juniper by leaf shape; Rocky

Mountain juniper has entire and opposite leaves rather

than minutely toothed and somewhat whorled leaves.

Ecological Relationships and Distribution—

Rocky Mountain juniper occurs throughout central

British Columbia and Alberta, and much of the Pacific

Northwest, Rocky Mountains, and Great Basin. It is

found from the northern Great Plains, south to Nevada, Arizona, New Mexico, and Texas (Fowells 1965;

Hitchcock and Cronquist 1973; Pieper 1983; Welsh

and others 1987; Zarn 1977). It is commonly encountered in canyons and cool exposures at higher elevations than Utah juniper, normally occurring as scattered plants rather than extensive stands (Goodrich

and Neese 1986). It usually grows with mountain big

sagebrush, mountain brush, ponderosa pine, aspen, and

Douglas-fir at elevations between 7,000 and 9,000 ft

(2,100 and 2,700 m) (Fowells 1965; Goodrich and

Neese 1986). At higher elevations it grows with

Engelmann spruce and subalpine fir. Although a

typical mountain species, it often occurs at low elevations in valley bottoms with cold air drainage (Welsh

and others 1987). It is most common in open woodlands or with big sagebrush-grass communities.

Throughout central Utah it is particularly abundant

with Gambel oak. The erect columnar variety is more

common in Utah on open aspects with mountain big

sagebrush. It is also common in northern Wyoming

and southern Montana.

Plant Culture—Cones appear from mid-April to midJune, mature from mid-September to mid-December
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 639

Chapter 23 Shrubs of Other Families

the second year after pollination, and persist on the

plant for 2 to 3 years (Fowells 1965; Johnsen and

Alexander 1974). Seeds are harvested, cleaned, and

stored as described for other junipers (Johnsen and

Alexander 1974). Seed production is erratic, but some

seeds are produced every year; abundant crops are

produced about every 2 to 5 years. Seeds are collected

by hand stripping or beating the limbs to dislodge the

cones onto canvas spread on the ground. Seeds are

relatively small compared with those of other junipers; the count ranges between 17,850 and 42,100

seeds per lb (39,350 and 92,800/kg) (Johnsen and

Alexander 1974).

Germination is delayed because of embryo dormancy

and hard impermeable seedcoats; seed pretreatments

are recommended to hasten germination. Untreated

seeds may require 14 to 16 months to germinate

(Afanasiev and Cress 1942). Johnsen and Alexander

(1974) reported that Rocky Mountain juniper seeds

require a warm pretreatment at 68 ∞F (20 ∞C) for 45 to

90 days followed by wet prechilling to initiate

germination.

Johnsen and Alexander (1987) suggested different

methods of pretreatment and seeding. These include:

(1) cleaning the seed followed by warm/cold wet pretreatment and fall seeding; (2) storage of the cones

(berries) for 1 year before cleaning, wet prechilling,

and fall or spring seeding; and (3) a warm pretreament

outdoors from spring to the time of fall sowing.

Seeds should be planted in a firm seedbed at a depth

of 0.25 to 0.5 inch (6 to 13 mm). Addition of a surface

mulch is helpful. When planting wildland sites, spots

should be selected that are somewhat protected and

have variable soil conditions. Seedlings do not emerge

from bare, open, compacted, or crusted surfaces.

Young Rocky Mountain juniper plants normally

grow faster than Utah juniper until they reach mature

stature. Fowells (1965) reported fairly uniform growth

to an age of 40 years, at which time the growth rate

declines. Plants live 250 to 300 years (Tueller and

Clark 1975).

This species is easier to rear as bareroot or container

stock than Utah juniper. Seeds are somewhat less

dormant, and plants usually grow faster when irrigated and fertilized. Commercial growers often produce this species and are able to rear the plant with

good success. Transplant stock of various ages is

produced and sold as potted container stock or balland-burlap plants. Plants are usually 3 to 4 years of

age before they are of sufficient size for commercial

sales.

Uses and Management—Rocky Mountain juniper

(fig. 23) is commonly used for conservation, windbreak, and horticultural purposes. This species has

been useful for improving upland game bird habitat on

farmlands (Miller and others 1948). Its pyramidal

evergreen growth habit provides attractive yearlong

protection. The plant is sufficiently drought tolerant

to persist on sites receiving 12 to 14 inches (30.5 to 35.6

cm) of precipitation. It is better adapted to alkaline soil

conditions than most other junipers (Miller and others

1948). Rocky Mountain juniper is particularly useful

when planted in combination with other trees and

shrubs as a windbreak or screen. It withstands harsh,

cold temperatures, and can be used to protect roadways, buildings, and other facilities.

This species is also of importance as a cover and

forage plant for big game. It receives moderate use in

winter, spring, and fall (Kufeld and others 1973).

However, considerable differences are noted among

sites and game animals. Some plants are grazed

heavily, but most trees are not grazed extensively.

Livestock generally make little use of this plant.

Rocky Mountain juniper furnishes excellent cover

for a variety of wildlife. It frequently grows in association with various shrubs that provide winter cover. Big

game animals often use stands of Rocky Mountain

juniper for cover and protection. Sites that are used for

this purpose should be managed to maintain appropriate cover and habitat.

Rocky Mountain juniper frequently grows in canyon

bottoms, streams, seeps, and other riparian areas. It

provides excellent cover and structural diversity. Plants

also normally support a herbaceous understory if sites

are properly managed. Small pockets of Rocky Mountain juniper, intermixed with mountain big sagebrush,

are important areas for wildlife. The diversity provided by this plant is important for many upland sites

where lower growing shrubs occur.

Few other tree species are able to grow where this

native normally exists. Because it is relatively slow to

establish and mature, removal of this species should

be avoided. Restoration projects should be designed to

Figure 23—Young Rocky Mountain juniper trees

grow in association with mountain big sagebrush.
640 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 24—Silverberry grows along stream bottoms

with willows and other riparian species in eastern Utah.

leave this species in areas of high importance. Rocky

Mountain juniper normally grows on sites that are

moist enough to support understory herbs. Herb and

shrub seedlings are better able to establish with this

overstory tree than with Utah juniper. Direct seeding

of herbs can be recommended on areas where scattered plants exist. Complete removal of the trees is not

required to accomplish seedling establishment of desired native understory species.

Rocky Mountain juniper has been planted on mine

disturbances with mixed success. Small 1- to 2-yearold container stock has not established or persisted

well on mine wastes in southeastern Idaho and southern Montana. Larger stock, 2- to 6-year-old potted

plants, survive better than smaller stock. Supplemental irrigation for 2 to 3 years significantly improves

establishment on harsh sites. Once plants are well

rooted, they persist well.

Varieties and Ecotypes—None.

Family Elaeagnaceae ____________

Elaeagnus commutata

Silverberry

Description—Silverberry is a native, deciduous

shrub, mostly 3 to 6.5 ft (0.9 to 2.0 m) tall, with peltate

scales. Plants vary in growth habit, often occurring as

thicket-forming shrubs up to 12 ft (3.7 m) tall or

frequently as small trees (Johnson and Anderson

1980). Stems are stoloniferous and unarmed. Leaves

are 0.5 to 2.8 inches (1.3 to 7.1 cm long), 0.2 to 1.2

inches (6 to 31 mm) wide, elliptic to oblanceolate,

acute to obtuse or rounded and silvery on both sides

with brown scales. Flowers are one to four per axil,

0.4 to 0.6 inch (1.0 to 1.5 cm) long, and yellowish. The

fruit is drupelike, consisting of a dry indehiscent

achene enveloped by a fleshy persistent hypanthium.

Fruits are 0.2 to 0.4 inch (6 to 10 mm) long, mealy, and

silvery (Welsh and others 1987).

Ecological Relationships and Distribution—

Silverberry occurs from Quebec to the Yukon, and

south to Minnesota, Wyoming, Utah, and northern

Colorado (Harrington 1964). In Utah, it is usually

restricted to riparian zones at elevations between

6,000 and 8,000 ft (1,800 and 2,400 m) (Welsh and

others 1987). Plants occur in areas receiving 18 to 20

inches (45.7 to 50.8 cm) of annual precipitation, but

usually require supplemental water even on moist

sites (fig. 24). Silverberry is very winter hardy and

drought tolerant. In eastern Utah, silverberry grows

along streams and dry hillsides (Goodrich and Neese

1986). In the interior of Alaska it grows on rocky,

south-facing slopes and sandbars (Viereck and Little

1972), often forming thickets along major streams.

Plant Culture—Silverberry plants flower in early

summer. The large, silver-green drupelike fruits ripen

in late fall and early winter. This shrub produces

abundant seed crops at intervals of 1 to 2 years. Ripe

fruits can be picked by hand, or the bushes can be

beaten to dislodge the fruits onto canvas spread on the

ground (Olson 1974). The fruits are easier to clean

before they dry. Seeds are extracted from the fruit by

maceration in a Dybvig separator, and water is used to

grind and float the pulp away (Heit 1968a; Olson

1974). Between 2,700 and 4,600 cleaned seeds are

reported per lb (5,950 to 10,140 per kg) of pure seed

(Olson 1974).

Silverberry has not been widely used in range and

wildlife projects. Seeds should be planted 0.5 to 1 inch

(1.3 to 2.5 cm) deep in a firm seedbed. Seedling growth

rate is fairly rapid, and young plants persist quite well

if planted in adapted areas. Plants also establish well

from bareroot or container-grown stock. Silverberry is

reported to have stoloniferous stems (Johnson and

Anderson 1980), and can spread from underground

rootstocks (Viereck and Little 1972). Vegetative spread

has not occurred in wildlife plantings in Utah.

Uses and Management—Silverberry has been selectively used throughout the Western United States.

It was propagated as early as 1813 (Olson 1974) for

windbreaks and shrub row plantings, but has not

been used extensively in recent programs. The species has been recommended for windbreak and wildlife plantings in Wyoming (Johnson and Anderson

1980). Plants are grown as ornamentals and windbreaks in the interior of Alaska (Viereck and Little

1972). Plants produce leaves having an unusual silver-green cast. Leaves persist on the shrub until

early winter, thus furnishing an attractive background screen.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 641

Chapter 23 Shrubs of Other Families

In Utah, big game have made moderate use of the

shrub during the spring and fall months, but they

seem to prefer the plant much more as summer

browse. Reports by Kufeld and others (1973) and

Dusek (1971) described limited fall grazing but increased summer browsing by big game. The shrub has

been transplanted with good success on mine disturbances in southeastern Idaho, and has received only

limited use by mule deer.

Silverberry appears to have particular value for

riparian habitat improvement within its native range.

It transplants very well and grows rapidly even with

some understory competition. It is able to establish

and grow on moist and well-drained soils, typical of

disturbed riparian sites. It provides considerable cover

for animals and soil protection. It is a well-rooted plant

that can provide excellent soil stability, but it does not

spread as does Russian-olive to become a serious weed.

Silverberry has considerable value as an ornamental plant for residential and recreational sites. It can

be used to enhance parkways, campgrounds, and recreational sites, particularly near streams and semiwet

areas. It can be used to screen and protect structures,

roadways, trails, and campgrounds. It does not attract

heavy use by wildlife and can be used where animal

concentration is to be avoided.

Varieties and Ecotypes—None.

Family Elaeagnaceae ____________

Shepherdia argenta

Silver buffaloberry

Description—Silver buffaloberry is a deciduous,

thorny shrub with a spreading to ascending growth

habitat that attains heights of 6 to 10 ft (1.8 to 3.0 m)

(Thilenius and others 1974a; Welsh and others 1987).

Goodrich and Neese (1986) reported that plants may

reach heights exceeding 22 ft (6.7 m). Branchlets are

covered with silvery peltate scales and terminate in

sharp thorns. Leaves are opposite, petiolate, 0.2 to 2.4

inches (0.5 to 6 cm) long, and 0.12 to 0.55 inch (3 to 14

mm) wide. They are oblong, elliptic or lanceolate, and

silver gray on both sides. Plants are dioecious, with

axillary flowers that can be perfect or imperfect. Flowers are 0.1 to 0.16 inch (2.5 to 4 mm) long with

yellowish perianths. The red fruit is drupelike, 0.16 to

0.28 inch (4 to 7 mm) long, and edible, though tart

(Goodrich and Neese 1986; Sampson and Jespersen

1963; Welsh and others 1987). Staminate flowers have

eight stamens alternating with glandular thickenings

at the base of the perianth lobes (Welsh and others

1987). The perianth of pistillate flowers is short and

tubular, investing the ovary.

Ecological Relationships and Distribution—Silver buffaloberry is distributed from British Columbia

to Saskatchewan, and south to California, Nevada,

New Mexico, and North Dakota. It occurs along

streams in sagebrush and pinyon-juniper woodlands

(Sampson and Jesperson 1963). It is widely scattered

throughout Utah at elevations between 4,590 and

7,510 ft (1,400 and 2,290 m) along streambanks, terraces, and on moist or wet sites (Welsh and others

1987). It occupies ditch banks, abandoned fields, disturbed fence lines and powerlines (Goodrich and Neese

1986). In Wyoming it grows in bottomlands, on moist

hillsides, and on streambanks at elevations up to

7,000 ft (2,100 m) (Johnson and Anderson 1980).

Plant Culture—Plants flower in spring, beginning

in early April and continuing into June. Fruits ripen

from June to August, and they remain on the bush

until December (Plummer and others 1968; Thilenius

and others 1974a; Van Dersal 1938). Ripe fruits are

most easily collected by flailing the plant, causing the

fruits to drop onto canvas spread on the ground. Fruits

may be handpicked, but the thorny branches hinder

this approach.

Stems, twigs, and leaves must be separated from the

fruits before they are macerated and the pulp floated

away with water. The seeds (cleaned achenes) are

processed with a fanning mill to separate them from

small debris. Seedlots can be cleaned to a high purity.

There are between 18,000 and 67,000 seeds per lb

(39,700 and 147,700 per kg); the average is 41,000

seeds per lb (90,390 per kg) (Thilenius and others

1974a). Mirov and Kraebel 1939 and Plummer and

others (1968) reported the average number of cleaned

seeds per lb is between 10,800 and 18,000 (23,810 and

39,680 per kg). Cleaned seeds can be stored in open

warehouses for at least 5 years without appreciable

loss of viability (Plummer and others 1968).

Both embryo dormancy and hard seedcoats restrict

or delay germination. Thilenius and others (1974a)

found that wet prechilling for 60 to 90 days, followed

by exposure to alternating day and night temperatures from 86 to 69 ∞F (30.0 to 20.0 ∞C) for 30 to 60 days,

resulted in 26 to 93 percent germination.

Seeds sown in fall will germinate in spring. Seeds

should be planted at a depth of 0.5 to 1 inch (1.3 to

2.5 cm). Seedlings and young plants have established

moderately well on range or conservation plantings.

Seedling growth is only fair, but as plants reach 1 or 2

years of age, they become well established and are very

persistent. Seeds are easily handled and planted with

most conventional planters. Bareroot or containergrown transplants establish better than direct

seedings. The shrub is not browsed very heavily, nor

adversely affected by insects, consequently it is able to

establish without serious delays (Plummer and others
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Chapter 23 Shrubs of Other Families

Figure 25—Silver buffaloberry has been successfully

established on mine disturbances in southeastern

Idaho.

1968). Silver buffaloberry grows moderately well with

herbaceous understory species. If 1-year-old transplant stock is planted, very good survival and excellent

growth can be attained. The shrub is produced by

many commercial nurseries. Root sprouts can be propagated with good success.

Uses and Management—Silver buffaloberry has

been widely planted for shelterbelts, game habitat,

cover, and watershed protection (Thilenius and others

1974a). It has been used in ornamental plantings

because of its gray, willowlike foliage (Sampson and

Jespersen 1963). The thorny nature of this plant

limits its use in heavily populated areas. However, it

can be planted in recreation sites to screen areas,

direct foot traffic, and control or restrict use of campgrounds and parks. It is used as the outer row in

multirow shelterbelt plantings, but it is often planted

alone where a low, dense cover is required (Johnson

and Anderson 1980). The shrub is highly susceptible to

heart rot, and breakage by wind and snow results as

the wood becomes brittle (George 1953). In the Midwest, high winds have uprooted cultivated trees; consequently, this species is not recommended for windbreak plantings, particularly near buildings and

walkways. (George 1953).

Silver buffaloberry is recommended for use in pinyonjuniper, big sagebrush, inland saltgrass, and wet

meadow communities of Utah for wildlife habitat

improvement (Plummer and others 1968). It is especially useful on sandy soils and in moist sites

(Thornburg 1982). The thorny thickets created by this

shrub provide cover and nesting sites for birds and

other animals; quail (Martin and others 1951) and

some small mammals take the drupelike berries

(Hall 1946). Big game animals make considerable

use of the shrub during the summer but only moderate use in the winter (Compton 1966; Dusek 1971;

Kufeld and others 1973; Lamb 1968; McKean 1954;

Plummer and others 1968).

Although silver buffaloberry is normally recommended for moist site plantings, it has performed well

on mine disturbances in southeastern Idaho, Montana, and Wyoming (fig. 25). It has established and

grown well on sites receiving 16 to 24 inches (40.6 to

61.0 cm) of annual precipitation. The species has

established from transplant stock on mixed, infertile

mine wastes and exposed substrata. It has exhibited

adaptability to coarse-textured spoils, rocky substrata,

and mixed heavy-textured soils. It fixes nitrogen

(USDA Forest Service 1948), which explains its excellent vigor and growth on infertile mine spoils.

Thornburg (1982) reported that a number of collections are being evaluated for use on mined lands in the

northern Great Plains.

This species is used to restore riparian disturbances;

it is well suited to both mesic and dry sites. It provides

overstory cover, thickets, and low ground cover. It is

adapted to streambanks, sandbars, unstable soils,

and moist sites in pinyon-juniper and sagebrush

communities where few upright species grow. It transplants very well and can be used to populate open,

exposed sites.

Varieties and Ecotypes—None.

Family Elaeagnaceae ____________

Shepherdia canadensis

Russet buffaloberry, soapberry

Description—Russet buffaloberry is a native, thornless, small to medium, spreading shrub 3 to 9 ft (0.9 to

2.7 m) tall (Thilenius and others 1974a). Branches are

covered with brown peltate scales. Leaves vary in

length from slightly less than 0.25 to 3 inches (0.5 to

8 cm). They are ovate to lanceolate, rounded apically

and basally, green above and slightly pale beneath.

Flowers vary from 1 to several per axil, and 0.08 to 0.12

inch (2 to 3 mm) long. Fruits are 0.16 to 0.28 inch (4

to 7 mm) long, red, and quite succulent (fig. 26). When

mixed with water the fruits produce a soapy solution

(Welsh and others 1987).

Ecological Relationships and Distribution—

Russet buffaloberry occurs as an understory in aspen,

fir, lodgepole pine, spruce, open woodlands, and old

burns (Goodrich and Neese 1982; Viereck and Little

1972). It often forms dense thickets on gravel bars

bordering streams. It is distributed from east-central

Alaska to Newfoundland, and south to Maine, New

York, Michigan, New Mexico, and Oregon (Viereck

and Little 1972). It is widespread throughout Utah at

elevations between 6,690 and 10,500 ft (2,000 and
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 643

Chapter 23 Shrubs of Other Families

3,200 m) (Welsh and others 1987), and it occupies

similar elevations in Colorado (Harrington 1964).

Plant Culture—Thilenius and others (1974a) reported that this shrub is well suited for dry, rocky

banks, but experimental plantings in central Idaho on

mine and roadway disturbances have not been successful. Transplant stock propagated from native

stands has not established well when planted on

adjacent disturbances. The species appears sensitive

to shade, soil disturbances, and competition from

understory herbs. It has not established well on mine

spoils or on streambanks that have been subjected to

soil loss from flooding. Planting success may be significantly improved with the use of better quality stock,

but this species has been less widely used and has not

been as successful in revegetation plantings as silver

buffaloberry.

Uses and Management—Russet buffaloberry occurs at rather high elevations and on sites where logging,

road construction, and mining have created considerable disturbances; consequently, it has been used for

revegetation of these sites. It provides useful ground

cover and improves diversity in lodgepole pine forests

where other understory and shrubs are lacking.

Varieties and Ecotypes—None.

Family Elaeagnaceae ____________

Shepherdia rotundifolia

Roundleaf buffaloberry

Description—Roundleaf buffaloberry is a short

statured plant growing 3 to 6.5 ft (0.9 to 2.0 m) in

height. The twigs are stellate hairy with white or

yellowish trichomes. Leaves are ovate to oval, rounded

apically, silvery green above, pale beneath, and clothed

with trichomes that provide a silvery appearance.

Flowers are yellowish and very attractive. Fruits are

0.2 to 0.31 inch (5 to 8 mm) long, drupelike, and orange

to red (Welsh and others 1987). This is an attractive

shrub with distinct silver-green foliage, thick leaves,

and colorful fruits and flowers.

Ecological Relationships and Distribution—

Roundleaf buffaloberry is native to the blackbrush,

ephedra, shadscale, pinyon-juniper, and ponderosa

pine communities of southeastern Utah and Arizona

(Welsh and others 1987). It is not reported from Colorado. This shrub is endemic to the Colorado Plateau

(Welsh and others 1987), and occupies important areas

where restoration of native shrubs is desirable (fig. 27).

Investigative studies have been conducted to develop

methods to culture and establish this species by artificial plantings.

Plant Culture—Seeds of roundleaf buffaloberry

are relatively large compared to those of other

buffaloberry species. There are about 6,855 cleaned

seeds per lb (15,110 per kg) (Plummer and others

1968). Abundant seedcrops are produced infrequently,

usually once every 3 to 10 years, although some fruits

may be produced on individual plants more often.

Plants flower early in spring, and fruits usually ripen

in early to late July (Plummer and others 1968). The

plants are very attractive because of the color contrast

between the fruit and leaves. Berries are processed by

maceration to release and separate the seeds. Seeds

germinate erratically following extended periods of

wet prechilling. Young plants grow slowly, even if

protected from competition. Nursery and greenhousegrown stock also grow slowly, although older plants

grow somewhat faster.

Figure 26—Fruits of russet buffaloberry are brilliant red and highly attractive. Seedlings establish

well on mine sites and other disturbances.

Figure 27—Roundleaf buffaloberry is a unique,

attractive shrub common to arid regions of the

Colorado River drainage.
644 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 28—Nevada ephedra generally grows on dry

sites in pinyon juniper, big sagebrush, Indian ricegrass,

rabbitbrush, black sagebrush, blackbrush, salt desert,

and creosote communities.

Uses and Management—This species is most important in the unique desert shrublands of southern

Utah and Arizona. It provides useful ground cover,

aesthetics, and limited, but seasonal forage for big

game (Plummer and others 1968). Attempts to culture

this species have only resulted in fair success; but with

further research, this species may be of more use for

restoration of recreational sites and native shrub

communities. It has been successfully used in landscape plantings. It grows well with limited irrigation,

and once established produces considerable foliage. It

demonstrates utility in recreation plantings for screening campgrounds and related sites. It also provides a

useful and attractive native shrub that can be used to

restore native communities in highly used recreation

sites.

Some collections of roundleaf buffaloberry have not

survived when transferred to more northern climates

in central Utah. This species normally grows with a

limited understory and cannot compete well if planted

with a dense cover of herbs. Plants are difficult to

establish from transplanting or direct seeding.

Varieties and Ecotypes—None.

Family Ephedraceae _____________

Ephedra nevadensis

Nevada ephedra

Description—Nevada ephedra or jointfir is a member of the gymnosperm family Ephedraceae. This

dioecious shrub grows to 5 ft (1.5 m) tall and has

evergreen stems with paired scalelike leaves 0.08 to

0.2 inch (2 to 5 mm) long (Welsh and others 1987).

One to several male cones are borne at the nodes.

Female cones are borne solitary at nodes and produce

a pair of pale-brown to yellowish-green seeds (Welsh

and others 1987). The stems of Nevada ephedra are

divergent, coarse, bluish-green to gray, and produced

on older wood. In contrast, green ephedra produces

erect, parallel, broomlike stems that are green to

yellow green in color (Cronquist and others 1972;

Dayton 1931).

Ecological Relationships and Distribution—

Nevada ephedra is a native perennial shrub that

occurs throughout the Intermountain West in juniperpinyon, big sagebrush, Indian ricegrass, rabbitbrush,

black sagebrush, blackbrush, salt desert, and cresotebush types in central and southern Utah, Nevada,

Arizona, California, and Mexico (Cronquist and others

1972). It frequently grows with green ephedra; however, it is usually more abundant on drier sites and in

salt desert, black sagebrush, and blackbrush types.

This species is generally found growing in areas with

7 to 14 inches (18 to 36 cm) of annual precipitation.

Nevada ephedra has good tolerance to soil salinity and

can be found growing in well-drained to fairly poorly

drained soils ranging from rocky, sandy loams to fairly

heavy clays (Dayton 1931; Plummer 1977). Nevada

ephedra occurs in open stands of scattered plants, but

commonly forms dense, slowly spreading colonies.

Individual plants may be connected by underground

stolons that develop when branches are covered with

windblown sand or silt (Wallace and Romney 1972).

Plant Culture—Seed crops are produced erratically (Cronquist and others 1972; Wallace and Romney 1972). Generally, one good seed crop is produced

every 5 to 6 years (fig. 28). Consequently, seed must be

harvested and stored for anticipated planting needs.

Within the Great Basin, cones appear in April to early

May, and seeds ripen in July (Dayton 1931; Turner

and Randall 1987). Farther south, cones appear in

March and seed matures in June. Seeds are easily

collected, cleaned, and seeded. Mature seeds must be

collected quickly before they drop from the shrub or

are gathered by rodents. They are collected by beating

the cones into a container. Establishment and maintenance of seed orchards on adapted sites might alleviate seed supply problems. Four to 6 years should be

allowed for stand establishment before significant

seed crops are produced. Seeds can be easily separated

from debris with an air screen cleaner, providing seeds

with better than 99 percent purity. There are about

20,000 Nevada ephedra seeds per lb (44,100 per kg) at

100 percent purity.

Seed remains viable for long periods when stored air

dry at room temperature. Seedlots of Nevada ephedra
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 645

Chapter 23 Shrubs of Other Families

exhibited 89 percent germination following 15 years of

storage (Stevens and others 1981a). With 25 years of

storage, germination was 77 percent (Stevens and

Jorgensen 1994). Seeds germinate under a wide range

of temperatures without wet prechilling or other pretreatments (Young and others 1977). Seeds of Nevada

ephedra will germinate somewhat faster than seeds of

green ephedra. Nevada ephedra can be seeded separately or in conjunction with other shrubs, grasses,

and forbs within its native range. When drill seeded,

Nevada ephedra and most other shrubs do best if

planted in separate rows from competitive perennial

grasses. Seeding Nevada ephedra through a seed

dribbler or thimble seeder provides good results. Nevada ephedra does best when seeded on a firm seedbed

and covered to 0.5 inch (1.3 cm) depth.

In most of the Intermountain West, Nevada ephedra

should be seeded in fall to allow for effective use of

spring soil moisture and precipitation (Plummer and

others 1968). In the more southerly regions, good

results have been obtained with midsummer seeding,

just prior to the summer storm period. Plants often

require 5 to 10 years to attain a height of more than

2 ft (61 cm) (Plummer and others 1968), yet seedlings

emerge and establish quickly. Seedlings withstand

extreme drought once established.

Nevada ephedra can be transplanted with moderate

success (Plummer and others 1977). Roots of bareroot

stock are generally weakly developed and are very

fragile and easily damaged. The root systems of containerized stock are sometimes too small to bind the

root plug together. The plug must be carefully removed from the container to avoid breaking the roots.

Low root-to-shoot ratios are common for both types of

stock, and survival rates are frequently less than 40

percent. Wieland and others (1971) found that Ephedra species could be propagated from rooted cuttings.

Everett and others (1978a), however, reported little

success in rooting cuttings of either Nevada ephedra

or green ephedra.

Uses and Management—Nevada ephedra provides forage and cover for cattle, sheep, deer, and

antelope (Dayton 1931; Dittberner and Olsen 1983;

Kufeld and others 1973; Smith and Beale 1980; Stevens

1983b; Welsh and others 1987). It is reported to be the

most important forage plant of the ephedra species

(USDA Forest Service 1937). Use is often difficult to

detect or measure because browsing causes the young

stems to break off at the joints. Nevada ephedra can

be an important source of browse during drought and

winter periods. Quail and cottontail rabbits use the

species for food and cover (Stanton 1974). Seeds are

readily gathered and eaten by rodents and birds.

Rodents cache seeds, which can then emerge in clusters of new seedlings.

Nevada ephedra has been used to stabilize rocky and

sandy areas (Plummer 1977). It is well adapted for use

as a low-maintenance, low-water-requiring ornamental for home sites, parks, recreational areas, and

administrative areas. It establishes slowly and has

not done well on exposed substrata at mine sites or on

road disturbances. Plants should not be grazed for two

to three growing seasons following seeding. Grazing

and trampling should be avoided until shrubs become

firmly established and are 5 to 8 inches (13 to 20 cm)

tall.

Varieties and Ecotypes—None.

Family Ephedraceae _____________

Ephedra viridis

Green ephedra

Description—Green ephedra is also called Brigham

tea, Mormon tea, or jointfir. This dioecious shrub has

evergreen stems, can grow to 4 ft (1.2 m) tall, and

produces a deep fibrous root system (Cronquist and

others 1972; Wasser 1982). Green ephedra produces

erect, parallel, jointed, broomlike stems that are

green to yellow green (Cronquist and others 1972).

Leaves are small ligulelike scales, and are opposite at

stem nodes. Plants are dioecous; male cones are compound, borne at the nodes or terminal points on the

stems. Female cones are solitary or whorled, and

sessile or peduncled (Wasser 1932; Welsh and others

1987). Cones are formed in pairs and are slightly boat

shaped.

There are two varieties of green ephedra. E. v. var.

viridis has sessile female cones, nonviscid stems, and

grows to 6 ft (1.8 m) tall. E. v. var. viscida has

pedunculate female cones and viscid stems. It occurs

on sandy soils where it may be grasslike in appearance; it is often covered with sand with only small

stems protruding (Kartesz and Kartesz 1980; Welsh

and others 1987).

Ecological Relationships and Distribution—

Green ephedra is a native, perennial shrub that occurs

on shallow to medium depth soils, on sandy or rocky

slopes and in valleys (Plummer and others 1968;

Wasser 1982) throughout the Intermountain West

(Cronquist and others 1972). It occurs in juniperpinyon-sagebrush, rabbitbrush, black sagebrush,

blackbrush, salt desert shrub, ponderosa pine, and

mountain brush types (Plummer and others 1968;

Stevens 1983b) (fig. 29). Green ephedra frequently

grows with Nevada ephedra; however, it is usually

more abundant on mesic sites with 9 to 14 inches (23

to 36 cm) of annual precipitation. Green ephedra

occurs as dense, individual bushes and in large stands.
646 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

It sprouts readily from the root or crown following fire

(Young and Evans 1974, 1978b).

Plant Culture—Cones develop in April and May;

seeds ripen in July and early August. The large seeds

are easily collected, cleaned, and planted. Mature

seeds must be collected quickly because they are

dislodged from the plant by wind. Seeds are also

quickly gathered by rodents and birds. Heavy seed

crops are produced erratically (Plummer and others

1968). Generally, one good seed crop is produced every

5 years. Consequently, seed must be harvested and

stored for later use. Seeds are harvested by dislodging

them into a container. They can be easily separated

from debris with an air screen cleaner. Seed lots

usually are 99 percent pure. There are about 25,000

green ephedra seeds per lb (55,100 per kg) at l00

percent purity.

Seed remains viable for long periods when stored air

dry at room temperatures. Stevens and others (1981a)

found that seedlots of green ephedra exhibited 87

percent germination following 15 years of storage.

With 25 years of storage, germination declined to 2

percent (Stevens and Jorgensen 1994). Seeds will

germinate under a wide temperature range without

wet prechilling or other pretreatments (Young and

others 1977).

Green ephedra can be successfully seeded with other

shrubs, grasses, and forbs within its native range. If

drill seeded, it should be planted in separate rows to

reduce competition with perennial grasses. Seeding

green ephedra with a seed dribbler or thimble seeder

has resulted in good success. Seeds establish best

when planted at a depth of 0.5 inch (1.3 cm) on a firm

seedbed. Rodents often gather planted seeds, and can

limit the success of direct seedings.

In the Intermountain region, green ephedra should

be seeded in the fall. (Plummer and others 1968;

Vories 1981). Seedlings that emerge early and quickly

can best withstand extreme drought. Both green ephedra and Nevada ephedra can be transplanted with

only moderate success (Luke and Monsen 1984). The

roots of bareroot stock and container stock develop

slowly and are easily damaged. Survival rates are

frequently less than 40 percent.

Uses and Management—Green ephedra provides

forage and cover for cattle, sheep, deer, and antelope

(Dayton 1931; Dittberner and Olsen 1983; Kufeld and

others 1973; Smith and Beale 1980; Stevens 1983a;

Welsh and others 1987). It is also grazed by bighorn

sheep, rabbits, and quail (Stanton 1974). Green ephedra is heavily hedged by sheep, particularly during

winter periods. Because of its evergreen growth habit

and abundant herbage that protrudes above the snow,

green ephedra is an important species on winter game

and livestock ranges. Seeds are gathered and eaten by

rodents and birds, which tend to spread the species.

Both green ephedra and Nevada ephedra can be

used to stabilize rocky, sandy, and disturbed areas

(Plummer 1977; USDA Forest Service 1976; Wasser

1982; Young and others 1979a). However, green ephedra persists much better on disturbances and exhibits

adaptation to a wider range of soil conditions.

Although both green ephedra and Nevada ephedra

have been used in ornamental and landscape plantings

(USDA Forest Service 1976; Wasser 1982), green ephedra has survived much better, and is less sensitive to

supplemental irrigation and cultivation.

Varieties and Ecotypes—None.

Family Ericaceae ________________

Arctostaphylos uva-ursi

Bearberry manzanita

Description—Bearberry manzanita is a prostrate,

mat-forming, evergreen shrub with stoloniferous stems

(Hitchcock and Cronquist 1973; Morris and others

1962; Noste and Bushey 1987; Welsh and others 1987).

The branches are ascending; the internodes are usually apparent, puberulent, and sometimes glandular.

The exfoliating bark has a dull brown to red color.

(fig. 30). Leaves are 0.2 to 1.1 inches (0.6 to 2.7 cm)

long, 0.1 to 0.5 inch (3 to 12 mm) wide, oblanceolate to

spatulate, and glabrous or puberulent, especially on

the margins. The inflorescence is racemose. The axis

and bracts are glandular. Flowers are pink to white,

and 0.16 to 0.2 inch (4 to 5.2 mm) long. Fruits are 0.2

to 0.4 inch (6 to 11 mm) thick, globose, bright red, and

Figure 29—Green ephedra exists as an important component of many shrub communities

in the foothill regions of central Utah.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 647

Chapter 23 Shrubs of Other Families

berrylike with separable nutlets. The four to 10 nutlets are fused or partially fused, 0.12 inch (3 mm) long,

0.08 inch (2 mm) wide, and light brown with a very

thick seedcoat (Belcher 1985; Welsh and others 1987).

Many roots appear near the soil surface to form a

dense, fibrous system (Berndt and Gibbons 1958).

Numerous runners extend laterally from the crown

to a distance in excess of 3 ft (0.9 m) (McLean 1967).

Bearberry manzanita is able to regenerate by resprouting from stolons (Berndt and Gibbons 1958). Rowe

(1983) reported that sprouting also occurs from large

lignotubers.

Ecological Relationships and Distribution—

Bearberry manzanita is circumpolar in distribution,

occurring in North America from Labrador to Alaska,

and south to northern California, and the mountains

of New Mexico, Virginia, Illinois, and Nebraska (Berg

1974; Sampson and Jespersen 1963). It is common in

ponderosa pine and Douglas-fir forests in mountainous regions, and it also occurs in oak woodlands and

chaparral types in Arizona and California. Cooper and

others (1987), Mueggler and Campbell (1986), and

Youngblood and Mauk (1985) reported bearberry

manzanita was present in many of the conifer forest

and aspen types in Utah and northern Idaho. In Utah

it is encountered in conifer forests at 7,000 to 11,500 ft

(2,140 to 3,510 m) in the north and north-central

portions of the State (Welsh and others 1987).

Bearberry manzanita exists on a variety of soil types

including weakly developed soils and coarse-textured

materials (Carmichael and others 1978). It is usually

regarded as a late seral species. However, it does not

form an abundant understory in forest communities

in Utah and northern Idaho (Cooper and others 1987;

Mueggler and Campbell 1986; Youngblood and Mauk

1985). It persists in open and shaded areas for extended

periods, and thus remains as a component of various

successional stages of community development.

Bearberry manzanita is generally not considered

fire tolerant (McKell and others 1972), but some plants

are able to survive burning. Following fire, new shoots

develop from portions of the plant that did not burn.

Seeds stored in the soil are stimulated to germinate by

the heat treatment provided from burning (Carmichael

and others 1978), and numerous seedlings may appear

1 to 5 years after a burn (Pase and Pond 1964).

Plant Culture—Flowers appear from March to

May; fruits ripen from June to August and are dispersed from August to March (Berg 1974; Everett

1957; McMinn 1951; Munz and Keck 1959).

Fruits are collected by hand stripping, but some can

be dislodged by beating the bush with a paddle.

Seeds are easier to clean when the fruit is still green.

If the fruit is allowed to dry, the fruit coat and fleshy

material are difficult to remove. Cleaning is accomplished using a macerater to grind and remove the

fleshy material from the seeds. Grinding is frequently conducted using water to flush away the

fleshy material. After grinding, the material is dried

and screened with a fanning mill to separate the seeds

from other materials. Certain seedlots have a large

number of empty seeds; these can be removed by

flotation or use of a gravity table separator. Seeds can

be stored for long periods at room or warehouse temperatures (Glazebrook 1941). Bearberry manzanita

produces between 26,800 to 58,000 clean seeds per lb

(59,080 to 127,870 per kg) (Berg 1974; Glazebrook

1941; McKeever 1938; Plummer and others 1968).

Seedcoats are thick, hard, and impermeable to water; however, a small opening extends from the surface

of the seedcoat to the embryo. This is plugged with a

waxlike substance that may be removed by soaking

the seeds in sulfuric acid. Wet prechilling treatments

are required to overcome the embryo dormancy. In

addition, the embryo requires afterrippening

(Glazebrook 1941). Most authors recommend various

periods of acid scarification, followed by a warm pretreatment and a period of wet prechilling. Berg (1974)

and Milstein and Milstein (1976) recommended 2 to 5

hours of acid scarification and planting in summer, or

a warm pretreatment at 25 ∞C (77.0 ∞F) for 60 to 120

days, then a wet prechill for 60 to 90 days. McLean

(1967) suggested an acid soak for 7 hours, followed by

a warm, wet pretreatment at 68 ∞F (20.0 ∞C) for 90

days, and a wet prechill at 34 ∞F (1.1 ∞C) for 90 days.

Giersbach (1937) recommended an acid soak for 3 to 5

hours, then overwinter storage outdoors in mulch

flats.

Following treatment, seeds are usually incubated at

high temperatures. Milstein and Milstein (1976) recommended incubating pretreated seeds at a constant

temperature of 78 to 80 ∞F (25.6 to 26.7 ∞C). Berg (1974)

Figure 30—Bearberry manzanita is a mat-forming,

evergreen shrub that provides excellent ground cover.
648 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

suggested either constant temperature incubation at

77 ∞F (25.0 ∞C), or alternating 86 ∞F (30.0 ∞C) day, and

68 ∞F (20.0 ∞C) night. These treatments produced 30 to

61 percent germination in 15 to 30 days. Germination

exceeding 90 percent has been reported by Glazebrook

(1941).

Early summer planting, following acid soaking, is

recommended by Berg (1974), Milstein and Milstein

(1976), and Swingle (1939), but early fall seeding of

treated seeds is also employed. Seeds are small and

should be planted approximately 0.5 inch (1.3 cm)

deep. Seed germination will normally occur in 20 to 40

days (Milstein and Milstein 1976). On wildland sites,

seedling emergence has been observed to occur in 5 to

10 days. Seedlings are slow developing and can be

suppressed by competitive vegetation. Also, seedbeds

often dry rapidly and some seedlings fail to survive.

Plants can be easily propagated by stem tip cuttings

(Berg 1974). Doran (1957) reported that cuttings taken

in late fall or winter (October to February) root well.

Eighty-eight percent of cuttings taken in October

rooted in 22 weeks, but if treated with IBA, 3 mg per g

of talc, 96 percent rooting occurred in 15 weeks. Stem

cuttings taken in February rooted best, 90 percent in

11 weeks. Cuttings are rooted under greenhouse conditions using bottom heat at 76 ∞F (24.4 ∞C). Once stem

cuttings are rooted, they survive field planting very

well.

Bearberry manzanita is widely used for horticultural plantings. Considerable information is available

related to vegetative propagation and culture

(Darbyshire 1971; Dehgan 1972; Dehgan and others

1975; Hildreth 1969; Leopold and Kriedeman 1975).

Dehgan and others (1977) recommended stem cuttings be taken in winter or early spring and treated

with low IBA concentrations. Rooted cuttings survive

better if hardened off gradually, starting with complete shade and progressing to an open situation.

Potting material and pots should be well drained to

avoid overwatering.

Uses and Management—Various authors consider the genus, Arctostaphylos, to provide quality

browse (Conrad 1987), but reports by Cada (1971),

Cooperrider (1969), Cowan (1947), Gullion (1964), Hill

and Harris (1943), Kamps (1969), Kufeld and others

(1973), and Lovaas (1958) indicated that the plant

receives moderate use by wildlife. In general, it is one

of the earliest shrubs to initiate growth in spring.

Leaves are also maintained on the shrub during winter, and game animals are attracted to the plant

during the winter and spring periods. It often grows on

open sites, not heavily covered by snow, that are

important winter grazing locations. Game animals

frequently utilize the shrub, although it may not be a

major portion of their diet.

Bearberry manzanita is widely used for landscape

plantings, and numerous selections are available.

Because it grows as a low-spreading plant, it has been

used in erosion control and watershed plantings. It

requires special care to assure establishment by transplanting as container or bareroot stock. Transplants

must be dormant when field planted or they are

susceptible to damage by frost or drought. Dormant

stock that is field planted early in the spring survives

well.

Small rooted transplants usually grow slowly, and

often fail to survive if planted on unstable slopes or

surfaces. Larger transplants (2-year-old stock) survive harsh conditions much better and grow faster.

However, growth is usually slow and plants fail to

provide immediate ground cover. Normally 3 to 5

years are required before plants spread and occupy a

large area. If this species is used to control erosion,

transplants should be spaced 1.5 to 2.5 ft (46 to 76 cm)

apart, depending on surface conditions.

Collections acquired from wildland sites can be

propagated for field plantings, but have demonstrated

adaptability to specific site conditions. Materials growing in shaded areas beneath ponderosa pine perform

erratically when transplanted in open areas at similar

elevations. Transplants also grow poorly on road and

mine disturbances, although poor performance may

be due to poor planting stock. However, this species

appears to be sensitive to soil conditions and has not

survived on mine disturbances.

Bearberry manzanita slowly invades road and logging disturbances, but only in areas where some topsoil has accumulated. Although use is questionable on

harsh disturbances, its growth habit is an important

feature. Topsoiling and use of soil amendments significantly enhance survival and growth rates.

This plant has adapted well when used for recreational sites, landscaping summer homes, and other

low-maintenance landscape areas. Considerable differences in ecotypes and growth forms are available to

provide diversity in landscape plantings. It could be

more widely used within the Intermountain area for

this purpose.

Varieties and Ecotypes—Various horticultural

selections are available, but materials have not been

selected for wildland uses.

Family Fagacae _________________

Quercus gambelii

Gambel oak

Description—Gambel oak (fig. 31) is a member of

the white oak subgenus Lepidobalanus. Most taxonomists consider Gambel oak part of the Quercus

undulata complex, which hybridize readily (Welsh
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Chapter 23 Shrubs of Other Families

1986). Gambel oak is a native, broadleaf, deciduous

shrub that typically forms dense stands (Neilson and

Wullstein 1983). It generally grows from 3 to 20 ft (0.9

to 6 m) in height, forming spreading thickets connected by underground rhizomes (Christensen 1949).

In the southern end of its range it can be treelike and

as much as 20 to 25 ft (6 to 7.6 m) tall (Wagstaff 1985).

Leaves are alternate and broadly obovate to oblong

lanceolate, with considerable variation among clones.

Leaves are somewhat leathery, yellow green to dark

green above and pale yellowish and densely hairy

below. They turn a deep-reddish color in fall.

This monoecious, wind-pollinated shrub bears red

pistillate flowers in the upper leaf axils. Male flowers

grow in pendent catkins that are borne singly or in

groups at the base of new shoots (Harper and others

1985). Fruits are brown acorns enclosed in a cap.

Acorns form and mature in 1 year (Harper and others

1985). Reproduction is primarily vegetative from an

extensive, freely branching, underground network of

lignotubers (enlarged, stemlike structures covered by

numerous adventitious buds), roots, and rhizomes

(Tiedemann and others 1987). Lignotubers account for

up to 72 percent of the total belowground biomass

(Tiedemann and others 1987).

Ecological Relationships and Distribution—

Gambel oak is widely distributed throughout the Southern and Central Rockies. In the Intermountain West,

Gambel oak occurs in valleys, foothills, on alluvial

fans, and on lower mountain slopes between sagebrush, pinyon-juniper, aspen, and spruce-fir types.

When it is not the dominant species, it can be found

growing as a codominant with big sagebrush, pinyon,

juniper, ponderosa pine, mountain mahogany, bitterbrush, cliffrose, aspen, snowberry, spruce, and fir.

Gambel oak occurs between 4,000 and 9,000 ft

(1,200 and 2,700 m). Pure stands occur along the

Wasatch Mountains, Wasatch Plateau, and west slope

of the Rocky Mountians in western Colorado in nearly

continuous belts (oakbelt) between 5,500 and 7,000 ft

(1,700 and 2,100 m) (Cottam and others 1959). Gambel

oak generally requires annual precipitation between

15 and 22 inches (38.1 and 55.9 cm) (Harper and

others 1985). However, it does occur in some areas

where annual precipitation is as low as 12 inches

(30.5 cm). The most common soils in which Gambel

oak occurs are calcarcous, heavy, and fine-grained

loams. It can also be found on sandy, gravelly, and clay

loams, and alluvial sands (Christensen 1955; Tucker

and Muller 1958).

Gambel oak occurs as a climax species on many sites,

and as a mid or early seral species on others (Harper

and others 1985). In some areas Gambel oak is considered the ecological equivalent of ponderosa pine (Harper

and others 1985).

Plant Culture—Gambel oak reproduces both vegetatively and sexually. Clones expand through both

radial and lateral growth (Christensen 1955). Rate of

vegetative spread ranges from 1.5 to 12 inches (381 to

360 mm) annually; the average is 4 inches (10 cm) per

year (Christensen 1955). Gambel oak sprouts quickly

after burning, wood harvesting, chaining, cabling, or

other disturbances. Sprouts may appear 10 days after

removal of top growth (Harrington 1985). Rate and

degree of sprouting are greatest from clones with

smaller stems. Clones with stems greater than 12

inches (31 cm) in diameter are poor sprouters (Reynolds

and others 1970).

Sexual reproduction is by acorns. Acorns develop

and mature in 1 year (Harper and others 1985). They

usually ripen in late September and October, and can

be collected by beating them into a container. Acorns

must be stored in a cool, moist environment or they

will dry and die. Storage should not extend beyond

the first winter following collection because of rapid

loss of viability. Seeding should occur in late fall or

early spring. High pregermination mortality can result from predation by birds, mammals, insects, and

parasitism. Acorns can germinate in fall after dispersal; they may be killed by frost if not covered by

litter and snow (Christensen 1949). Seedlings are

susceptible to damage by early fall or late spring

frosts, grazing, and summer drought (Neilson and

Wullstein 1983). Gambel oak seedlings are rare in

many areas, particularly in the northern part of its

range (Christensen 1955; Reynolds and others 1970).

Plants can be established from properly planted

container stock. Establishment success has also resulted from moving partial clones with front-end

loaders. This involves moving and placing 2 to 3 ft

(61 to 90 cm) of soil, roots, rhizomes, and lignotubers.

Gambel oak wildings do not transplant well. Figure 31—Clumps of Gambel oak provide soil

protection and habitat for wildlife and livestock.
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Chapter 23 Shrubs of Other Families

Uses and Management—Gambel oak and associated species provide valuable food and cover for livestock and many wildlife species. This species is generally not highly palatable, but its abundance and

availability, particularly on fall, winter, and spring

ranges, make it an important browse species. Gambel

oak is regarded as fair forage for all classes of livestock

(Harrington 1985). Summer cattle use has been identified in northern Utah (Julander and Robinette 1950).

Sheep use can be light (Dayton 1931). Gambel oak is

used extensively by mule deer throughout the year

(Bowns 1985; Harper and others 1985; Kufeld and

others 1973). Elk browse oak during winter and spring

(Kufeld 1973; Reynolds and others 1970). Bighorn

sheep make use of the plant during summer months

(Rominger and others 1988).

Young shoots and new growth are the most preferred portions of this shrub. Preference of most browsing animals for Gambel oak varies with availability of

other species. Use increases as other species become

less available. Gambel oak can be poisonous to livestock when used exclusively, but when consumed in

combinations with other species, it is generally harmless (James and others 1980).

Quantity and quality of associated forage production can be very high in Gambel oak communities.

Understory production may reach 3,000 lb per acre

(3,360 kg per ha). Understory production within oak

clones can be twice as high as between clones. Understory species within clones also green up 2 to 3 weeks

earlier in the spring and stay green later in the season

than do the same species between clones (Bowns 1985;

Stevens and Davis 1985).

In most areas the Gambel oak type has the highest

forage productivity potential of any vegetative type in

the Intermountain West. It has been severely abused

by livestock at many locations. Productive understory

species have been grazed out and replaced by less

productive plants. In most areas the understory is

almost nonexistent. As a result oak density and clone

size have increased substantially.

The Gambel oak type is important for livestock,

wildlife, and watershed protection. These areas can be

critical spring and fall ranges for cattle, sheep, deer,

and elk, and winter range for elk. Floods produced by

high-intensity July and August rainstorms are common on depleted Gambel oak stands. Heavy snow pack

and spring snowmelt have the potential for producing

destructive spring runoff from these depleated communities. It is important that the Gambel oak type be

well managed and restored to its productive potential.

Gambel oak also provides valuable escape, thermal,

and travel cover for deer, elk, livestock, and small

mammals. A large number of birds use Gambel oak for

nesting, hiding, and resting (Marti 1977). Acorns are

highly preferred by deer, elk, hogs, javelina, turkey,

band tailed pigeons, squirrels, and numerous small

mammals and birds (Christensen 1949; Harper and

others 1985; Reynolds and others 1970; Steinhoff

1980). Many small birds and mammals cache acorns

for later use.

Gambel oak is a valuable browse and cover plant in

many areas, but it can grow too dense and exclude

understory vegetation, big game, and livestock. A

mosaic of clones or stands interspersed with open

areas provides the best habitat for big game and

livestock (Stevens and Davis 1985). Management objectives should be directed toward maintaining a mosaic of clones and interspaces. Fire, herbicides, and

mechanical treatment can be used successfully to open

stands and increase accessibility to clones (Bowns

1985; Stevens and Davis 1985). When Gambel oak is

disturbed, understory species should be seeded to

improve composition and production and to retard oak

suckering and regrowth (Stevens and Davis 1985).

Wood value of Gambel oak can be high (Clary and

Tiedemann 1986). Average energy content is estimated at 340,000 Btu per ft2 (4.77 kcal per g) (Harper

and others 1985); it can be an economically important

fuel wood with high heat-yielding qualities. Gambel

oak has been used successfully for long-term rehabilitation of disturbed sites (Dittberner and Olsen 1983).

Gambel oak resprouts rapidly following fire. Rate of

recovery varies with fire severity, climatic factors, and

site characteristics (Harper and others 1985). Fire

generally does not harm or kill Gambel oak. Spring

and fall burns have little detrimental effect. However,

summer burns can inhibit regrowth and may even kill

Gambel oak (Harrington 1985). Following fire, the

rate and amount of resprouting can be reduced by

seeding competitive herbs (Stevens and Davis 1985).

Gambel oak generally sprouts as early as 10 days after

being burned (Harrington 1987). When Gambel oak is

burned, chained, cabled, or otherwise disturbed, deer

and elk browsing increase (Stevens and Davis 1985).

Deer, elk, and livestock seem to prefer burned over oak

communities (Harper and others 1983).

Improved Varieties—None. Considerable variation exists with this species throughout its geographical

range.

Family Oleaceae ________________

Fraxinus anomala

Singleleaf ash

Description—Singleleaf ash is one of only a few

native ashes to occur in the Intermountain region. It is

a small, deciduous shrub or tree, commonly 8 to 13 ft

(2.4 to 4 m) tall (Welsh and others 1987), but sometimes attaining heights of 26 ft (8 m) (Goodrich and

Neese 1986; Harrington 1964). It usually develops
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Chapter 23 Shrubs of Other Families

multiple stems, and branchlets that are characteristically four angled. Leaves are simple to two or three

foliate, ovate, crenate-serrate to subentire, glabrous,

0.6 to 2.6 inches (1.5 to 6.5 cm) long, and 0.4 to 2.4

inches (1 to 6 cm) wide. Flowers are polygamous and

occur in panicles with a minute calyx and no corolla.

Fruit is a one-seeded samara, 0.5 to 1 inch (12 to 25

mm) long, 0.31 to 0.4 inch (8 to 10 mm) wide, and

winged almost to the base (Goodrich and Neese 1986;

Harrington 1964; Kearney and Peebles 1942; Munz

and Keck 1959; Welsh and others 1987).

Ecological Relationships and Distribution—

Singleleaf ash occurs from Colorado west to California

and south to New Mexico and Arizona (Harrington

1964). It is common in west-central and southwestern

Colorado at 4,500 to 6,000 ft (1,400 to 1,800 m)

(Harrington 1964). It occurs on rocky, sandy canyons

along the Green River in Utah (Goodrich and Neese

1986), but is more widespread throughout eastern and

southern Utah. It exists in pinyon-juniper woodlands

and mixed desert shrub communities mainly on rimrock or along drainages at elevations between 2,900

and 8,500 ft (890 and 2,600 m) (Welsh and others

1987). It occupies dry canyons and gulches between

3,000 and 11,000 ft (910 and 3,400 m) in California

(Munz and Keck 1959).

The plant is important on upland sites where it

grows intermixed with salt desert shrubs, pinyonjuniper, and basin big sagebrush. It is particularly

abundant in the Four Corners region and throughout

much of the Colorado River drainage where it occurs

on sandstone outcrops and shallow soils (fig. 32). It

does not form dense stands, but is normally found

scattered among rocky bluffs and ledges. It also occurs

on limestone soils, but usually on well-drained sites

(Stark 1966).

Plant Culture—Small, rather inconspicuous flowers appear from April to May at about the same time

the leaves develop. Flowers are grouped in terminal or

axillary clusters. Fruits are long, single-seeded samaras with distinct wings that are borne in clusters.

They ripen from mid-July to mid-August (Plummer

and others 1968). As fruits mature they change from

green to yellow or light brown. Fruits often remain on

the bush for a number of weeks before dispersing.

Ripened fruits can easily be collected by hand picking

or dislodging them onto canvas spread on the ground.

Fruits are dried before cleaning. They are separated

from stems and leaves by fanning or with air screen

cleaners. The wings can be removed from the fruit by

grinding or maceration. Removing the wings aids in

planting, but does not affect germination. Seeds of

most ash species must be stored in sealed containers at

4l ∞F (5.0 ∞C) and 7 to 10 percent moisture content.

Singleleaf ash produces good seed crops infrequently.

Abundant crops may be produced once every 3 to 5

years. Insects damage a number of seeds as they ripen.

Usually, less than 50 percent of the seeds from wildland collections are viable. A number of freshly collected seeds will germinate, but most are dormant and

require wet prechilling. Seeds sown in fall usually

receive an adequate wet prechilling treatment and

germinate uniformly in spring. Artificial wet prechilling of ash seeds is accomplished by diurnal alternating

temperatures of 86 and 68 ∞F (30 and 20 ∞C) (Bonner

1974b).

Uncleaned or dewinged fruits are difficult to seed

with conventional drills. The fruits are large and

irregular in shape and do not flow through seedboxes

and seeding equipment. Fruits should be planted in

the soil at a depth of about 0.5 inch (1.3 cm).

Trial seedings, primarily in central Utah, have produced irregular stands. This has been attributed to

seeding poor quality seed. Seedlings and young plants

grow quite slowly. Even under irrigation, nurserygrown plants develop slowly. Wildland planted seedlings often increase only 6 inches (15.2 cm) in height

during a single season. As plants reach 3 to 5 years of

age, the growth rate increases. A dense, well-branched

root system is formed the first year; once established

the plants are very persistent.

Singleleaf ash can be grown as transplant stock for

field plantings, although transplanting success has

been quite erratic. One-year-old transplants are not

very large, usually less than 5 to 6 inches (12.7 to

15.2 cm) tall, but with healthy root systems. Survival

of transplanted stock has often been 25 to 40 percent,

but losses are often attributed to extreme conditions of

the planting sites. Growth response of singleleaf ash

has varied among collections. Plant materials obtained from eastern Utah have grown quite slowly, but

plants establish and persist very well.

Figure 32—Singleleaf ash occurs on shallow soils and

sandstone outcrops in the Four Corners area.
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Chapter 23 Shrubs of Other Families

Uses and Management—Singleleaf ash is one of

only a few shrubs or trees adapted to rocky and arid

sites in the sagebrush, salt desert shrub, and pinyonjuniper communities. It develops an upright growth

form that provides food and cover for wild animals and

domestic stock. The plants are very persistent, and are

not eliminated by extended periods of drought or

heavy browsing. Singleleaf ash is also well adapted to

infertile soils, and performs well in such situations.

The plant serves as an overstory species with sagebrush and fourwing saltbush, providing cover and

protection for wildlife in barren areas. When intermixed with pinyon and juniper, it provides forage

diversity for game and livestock.

Singleleaf ash competes well with pinyon and juniper, and if not heavily grazed, it is able to remain

vigorous and healthy amid dense stands of these trees.

The species is an important plant for watershed protection; it grows in areas where high-intensity rains

occur infrequently, but can cause considerable erosion. This species is also able to persist and provide

cover on sites where few other species may occur.

Singleleaf ash has been used to a limited extent on

mine sites, roadways, and recreational disturbances.

A slow growth rate, however, limits its value in areas

where rapid development of cover is desired. It is

suited to rocky soils, but is not well adapted to mine

disturbances if planted on mine wastes or substrates.

It has considerable potential for landscape plantings,

particularly in highly scenic areas of eastern and

southern Utah. It can be used to restore disturbances

near recreational areas, parks, and roadways. Additional information is needed to improve the culture of

this important species.

Varieties and Ecotypes—None.

Family Polygconaceae ___________

Eriogonum heracleoides, Wyeth or

 whorled eriogonum, buckwheat

Eriogonum umbellatum, Sulfur

 eriogonum, buckwheat

Eriogonum wrightii, Wright eriogonum,

 buckwheat

Introduction—Eriogonum or buckwheat, as most

species are commonly called, occurs only in North

America. There are a considerable number of

Eriogonumspecies in the Western United States (Dayton 1931, 1960; Harrington 1964; Hitchcock and others 1964; Judd 1962; Welsh and others 1987). The

genus includes annual and perennial herbs, subshrubs, and woody shrubs. Most species have taproots; some of the subshrubs have spreading or prostrate stems that root at the joints (Dayton 1960). The

shrubby species generally produce semierect stems.

Flowering stocks are herbaceous, erect, and leafless.

Flowers are small and arranged in umbels, cymes, or

racemes. The fruit is a three-angled or three-winged

achene (Dayton 1931, 1960; Hitchcock and others

1964; Welsh and others 1987). Sulfur eriogonum,

Wyeth eriogonum, and Wright erigonum have been

the most successful species used in restoration projects.

They are easily cultured and establish well without

extensive seedbed preparation or protection of young

seedlings. They exist on dry situations, and prefer

rocky, sandy, well-drained soils in areas of moderate

to low rainfall. Consequently, they are useful shrubs

in restoration of harsh sites.

Varieties and Ecotypes—None.

Family Polygconaceae ___________

Eriogonum heracleoides

Wyeth or whorled eriogonum, buckwheat

Description—Wyeth eriogonum is a native perennial, mat-forming subshrub with woody branches and

persistent vegetative stems. Stems arise from

rosettelike bases and produce distinctive whorled

leaves. Flowering stems are densely branched and

grow to 2.5 ft (76 cm) in height. Cream-colored to yellow

flowers develop on umbellate inflorescences (fig. 33)

(Hitchcock and others 1964; Welsh and others 1987).

Abundant seeds are generally produced each year.

There are about 141,000 seeds per lb (310,800 per kg)

at 100 percent purity.

Ecological Relationships and Distribution—

Wyeth eriogonum is widely distributed in British

Columbia, Montana, Utah, Wyoming, Nevada, and

California (Dayton 1960). It is most prevalent in mountain brush and pinyon-juniper communities, but it

also occurs in sagebrush-grass, aspen, and spruce-fir

types. It invades disturbed sites but is less competitive

with annual weeds, particularly cheatgrass, than sulfur buckwheat. This species does well on disturbed

sites and openings in various communities and will

spread naturally if not heavily used (Plummer 1970).

It does increase following burns.

Plant Culture—Wyeth eriogonum, sulfur erigonum,

and Wright eriogonum produce good seed crops most

years. Even during dry years, some seeds develop.

Seeds are quite easily collected by beating them into a

container. Seeds are not difficult to clean; they are

simply separated from the debris with a fan and screen

separator.

Seeds of all three species can be drilled or broadcast

seeded, but they should not be planted more than 0.1

inch (3 mm) deep. Fall and winter seeding are preferred. Seeds of all three species germinate readily,
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Chapter 23 Shrubs of Other Families

reduced. Birds and small mammals seek out and

consume the seeds. This species has considerable

ornamental potential, especially on low-maintenance

areas.

Improved Varieties—None.

Family Polygonaceae ____________

Eriogonum umbellatum

Sulfur eriogonum

Description—Sulfur eriogonum or buckwheat is a

perennial subshrub with a strong taproot and freely

branching crown. The branches can be prostrate and

mat forming, but are usually upright, growing to a

height of 1 ft (30.5 cm). Leaves are oval to oblanceolate,

green above and grayish beneath, arising from

rosettelike stem tips. The inflorescence is a freely

branching umbel on a stem about 1 ft (30.5 cm) long.

Leaflike bracts develop near the middle of the flowering stem. Flowers are cream colored to deep yellow or

greenish yellow. Flowering stems and floral tissue

often persist throughout the winter (Dayton 1931,

1960; Hitchcock and others 1964; Welsh and others

1987).

Ecological Relationships and Distribution—

Sulfur eriogonum occurs widely throughout Western

North America in big sagebrush, pinyon-juniper, mountain brush, ponderosa pine, spruce-fir, aspen, and

subalpine communities (Dayton 1960). It is especially

well adapted to rocky and exposed sites, and quickly

invades and occupies disturbed areas (fig. 34). Seedlings

are persistent and competitive. They grow rapidly and

Figure 33—Wyeth eriogonum flowers in early summer and generally produces a reliable seed crop.

resulting in fairly uniform emergence over a 2-week

period in early spring. Seedlings are fairly frost and

drought tolerant. All three species establish fairly well

when planted in combination with grasses, shrubs,

and forbs. Aggressive perennial grasses can, however,

become dominant.

Sulfur, Wyeth, and Wright eriogonum establish well

from direct seeding and spread naturally on severely

disturbed sites such as mines, road cuts and fills,

construction sites, and gravel and sand pits. These

species are generally able to persist and increase in

density following wildfires. Buckwheats most often

increase in importance.

Uses and Management—Livestock and big game

graze Wyeth eriogonum (Dayton 1960). Most use occurs in late fall and winter when other species are

dormant, but some use occurs during summer. This

species recovers well from heavy use and will increase

in density and productivity when grazing pressure is

Figure 34—Sulfur eriogonum directly seeded on

shallow soils persists with competitive annual grasses.
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Chapter 23 Shrubs of Other Families

will spread into cheatgrass and perennial bunchgrass

communities.

Plant Culture—Seeds of sulfur eriogonum can be

harvested, processed, and seeded as described for

Wyeth eriogonum.

Uses and Management—Sulfur buckwheat can

be an important browse species for sheep, goats, cattle,

mule deer, elk, and antelope (Dayton 1931, 1960;

Kufeld 1973; Kufeld and others 1973; Monsen 1975;

Smith and Beale 1980). This shrub is semievergreen

and produces useful forage. Plants receive some summer use, but most use occurs in late fall and winter.

Even when heavily grazed, sulfur eriogonum survives

and spreads on sites where other important forage

plants have succumbed (Monson 1975). Birds and

small mammals consume the seeds.

The species has been seeded successfully on severely

disturbed sites. It provides rapid soil stabilization and

will spread fairly aggressively from seeds to fill openings in the community. It is able to survive on unstable

soils, but is not able to persist if buried by soil sloughing. Like other species of eriogonum, sulfur eriogonum

has considerable ornamental value, especially in lowmaintenance, low-precipitation situations.

Varieties and Ecotypes—“Sierra” sulfur flower

buckwheat (E. umbellatum var. polyanthum) was released for use in California.

Family Polygconaceae ___________

Eriogonum wrightii

Wright eriogonum

Description—Wright eriogonum is a low, white,

woolly, perennial shrub usually less than 2 ft (61.0 cm)

tall. It commonly forms a basal crown from which

many branched stems emerge. The leaves are oblanceolate, sharp pointed at the tips, and covered with

fine, white, woolly hairs. The white or pinkish flowers

are borne in umbels at the apex of rather short leafless

flower stalks (Judd 1962; Welsh and others 1987).

Ecological Relationships and Distribution—

Wright eriogonum is found primarily in pinyon-juniper,

oak woodlands, and chaparral types in southern Utah,

California, Arizona, New Mexico, and Texas (Welsh

and others 1987).

Plant Culture—This species is easily established

by direct seeding. Seedlings are vigorous, drought

tolerant, and persistent. Seeds are usually hand harvested and processed with an air separator.

Uses and Management—Wright eriogonum is considered fairly good forage for sheep and goats and fair

for cattle (Judd 1962; USDA Forest Service 1937).

Under heavy grazing, this species can be replaced by

burroweed and snakeweed (Judd 1962). Birds and

small mammals seek out and consume the seeds.

Wright eriogonum persists well during drought periods and will increase following burns. This species has

been seeded successfully on disturbed sites. It can

provide rapid soil stabilization and will spread fairly

aggressively from seed.

Varieties and Ecotypes—None.

Family Ranunculaceae ___________

Clematis ligusticifolia

Western virginsbower

Description—Western virginsbower is a woody,

climbing vine. The vigorous, glabrous to densely strigose, or villous stems grow to 33 ft (10 m) in length,

forming masses of vegetative growth that clamber

over neighboring trees and shrubs. Root systems are

shallow and fibrous. The opposite deciduous leaves

are pinnately compound with 3, 5, or 7 lanceolate to

broadly ovate to chordate leaflets. Each leaflet is 0.8 to

3.0 inches (2 to 7.6 cm) long and coarsely few toothed,

or sometimes lobed or nearly entire. The twisting petioles develop from enlarged stem nodes and curl around

supporting vegetation (USDA Forest Service 1937).

The sweet-smelling dioecious flowers are clustered in

few-to-many-flowered bracteate cymes. The four oblonglanceolate sepals are white to cream colored, 0.2 to

0.6 inch (5 to 15 mm) long. There are no petals. Male

flowers have numerous stamens and no pistils. Stamens of the female flowers are numerous, but sterile.

Pistils are numerous in a capitate cluster. The plumose styles of the villous achenes elongate to 1.0 to 2

inches (2.5 to 5.1 cm) at maturity (Clebsch 1979;

Harrington 1964; Hitchcock and others 1961; Welsh

and others 1987). Chromosome number is 2n = 16.

Geographic races are sometimes recognized (Vines

1960).

Ecological Relationships and Distribution—

There are more than 200 species of Clematis, occurring primarily in the Temperate Zone of the Northern

Hemisphere (Hitchcock and others 1964; Rudolf 1974).

Western virginsbower is distributed from British Columbia west of the Cascades to the Columbia River,

south on both sides of the Cascades to southern California, and east from Saskatchewan to New Mexico

(Hitchcock and others 1961; Rudolf 1974; Welsh and

others 1987). It commonly occurs along drainages, in

riparian communities, and on steep talus slopes and

canyon walls where it trails over boulders, banks, and

vegetation.

Western virginsbower is common on roadsides and

other disturbed sites. It frequently occurs with other

woody species such as cottonwood, willow, common
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Chapter 23 Shrubs of Other Families

chokecherry, Rocky Mountain maple, bigtooth maple,

rubber rabbitbrush, ponderosa pine, and Douglas-fir.

It grows on well-drained sandy to rocky soils, ranging

from weakly acidic to moderately basic, and may

occur on somewhat saline soils (Wasser 1982). It

usually grows in areas receiving 12 to 20 inches (30.5

to 50.8 cm) of annual precipitation, but it is most

abundant in areas receiving supplementary water.

Drought tolerance of established plants varies among

populations. Most are weakly to moderately drought

tolerant (Wasser 1982). Seeds and vegetative production are markedly reduced in dry years.

Plant Culture—Flowering occurs from March to

August and achenes ripen from May to December,

depending on geographic location (Mirov and Kraebel

1939; Plummer and others 1968; Swingle 1939). Moderate to heavy seed crops are produced almost every

year. Mature fruits are dry and brown with feathery

white styles. They are dispersed by wind and gravity;

rate of dispersal depends on weather conditions. Fruits

are gathered by hand or with a vacuum seed harvester

(Plummer and others 1968). Dried fruits are hammermilled with a 3⁄16-inch (0.5-cm) screen operated at

1,120 rpm to detach the styles from the achenes

(Wasser 1982). Debris is removed by fanning. Seeds

are not removed from the achenes.

Estimates of the number of seeds per lb range from

93,000 to 328,000 (205,000 to 723,000 per kg) (Mirov

and Kraebel 1939; Rudolf 1974; Swingle 1939). A

purity of 20 to 50 percent and germination of 70

percent are recommended for seed purchases. Greater

purities are difficult and expensive to achieve. Viability has been maintained for 2 years in dry storage

(Plummer and others 1968).

Wet prechilling at 33 to 40 ∞F (0.6 to 4.4 ∞C) for 2 to

180 days is required to release embryo dormancy

(Heit 1968a; Rudolf 1974; Young and Young 1986).

Germination of wet prechilled seeds after 40 to 60

days at 68 to 86 ∞F (20 to 30 ∞C) ranged from 11 to 84

percent (Mirov and Kraebel 1939; Plummer and others 1968; Swingle 1939).

Western virginsbower may be direct seeded in mixtures with other shrubs by drilling or broadcasting.

Untreated seeds may be fall planted (Rudolf 1974).

Seed should be wet prechilled prior to spring planting

(Swingle 1939). Seeds should be covered with about

0.3 to 0.5 inch (7 to 13 mm) of soil. Seedlings are

vigorous, but growth is reduced by herbaceous competition. Consequently, no more than 50 percent of the

mixture should consist of grass seeds. Western

virginsbower normally should be seeded in mixtures

at a rate of 1 lb per acre (1.1 kg per ha) or less (Wasser

1982). Holmgren (1954) reported good growth rates,

but low survival of seedlings emerging from seedings

on southern Idaho big game winter range. Seedlings

were quite sensitive to competition from cheatgrass

and broadleaf herbs.

Bareroot and container stock establish readily and

grow rapidly. Seeds should be cleaned carefully to

improve stand uniformity in nursery plantings. Container seedlings may be established from seeds or from

cuttings of partially matured wood harvested from

late spring to late summer. Young wood with short

internodes gives less satisfactory results. Leaf bud

cuttings taken in summer may be rooted under mist

(Hartmann and others 1990). Layering is useful if only

a few plants are needed.

Uses and Management—Western virginsbower

is used in seeding or transplanting projects to improve cover and forage for many birds and small

mammals. Smith (1953) found that captive mule deer

preference for western virginsbower was greatest in

early summer and decreased considerably by late

summer relative to other native shrub forages provided. The plant receives little use by big game on

mule deer winter ranges in southern Idaho (Holmgren

1954). Dixon (1934) reported that its palatability to

mule deer in California was low. It receives little use

by livestock.

Because of its rapid growth rate, generally low

palatability, and ability to naturally invade disturbed

sites, western virginsbower is a potentially valuable

species for restoring disturbed sites with native vegetation (Bailey 1947; Plummer and others 1968; Rehder

1940; Van Dersal 1938). It has been used for soil

stabilization on riparian areas and mined lands

(Thornburg 1982), and is particularly useful on steep

slopes that would otherwise be revegetated only slowly

by herbaceous species (fig. 35). Wider use of the species

Figure 35—Western virginsbower growing on a

road fill provides excellent soil stabilization.
656 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

Figure 36—Capsules of wedgeleaf ceanothus must

be harvested when green as they dehisce explosively

when dry.

is precluded by the difficulty of harvesting and processing large quantities of seed. Seedlings established

from seeds or transplant stock develop rapidly, often

flowering the first year, and provide cover while other

planted species develop more slowly. Seeds and seedlings are sometimes consumed or damaged by small

mammals. Leaf spots and rusts are often noted on the

foliage, and downy mildew is occasionally a problem

(Holmgren 1954).

Plants are moderately shade and frost tolerant, but

thermal tolerance varies among ecotypes (Wasser

1982). Shoots may be destroyed by wildfires. Some

plants survive by resprouting from rootstocks. The

species also reestablishes burned areas by wind dispersal of seeds produced offsite (Plummer and others

1968; Thornburg 1982).

Western virginsbowers’ cascade of yellowish foliage

and clusters of feathery styles make it a unique and

attractive ornamental for use in dry areas (Plummer

and others 1968). The plant requires a large shrub or

small tree for support. It may be trained to grow over

a fence or trellis. Late winter pruning confines the

plant to smaller areas (Clebsch 1979). Plants are most

productive if planted well away from the supporting

plant in slightly acid to moderately alkaline soil. Rich,

moist soil and mulching provide needed thermal protection for the root system.

Varieties and Ecotypes—None. Selection trials

are being conducted by the USDA Soil Conservation

Service, Pullman Plant Materials Center, Pullman,

WA, for a cultivar adapted to the northern Intermountain region.

Family Rhamnaceae _____________

Ceanothus species

Introduction—Numerous species of Ceanothus occur in the Western United States. Munz (1974) described over 75 species in California. Considerably

fewer species occur in the Intermountain region

(Harrington 1964; Welsh and others 1987). However,

many species common to the Southwest, northern

California, Oregon, and southern Washington have

been tested in the Intermountain area. Species that

have been entered into adaptation trials were selected

from sites with somewhat similar climatic and edaphic

conditions.

Different species of Ceanothus possess important

traits and have been planted in an attempt to enhance wildlife, range, watershed, or horticultural

resources in areas outside their natural range. Although a number of species have been initially

screened for adaptability to the Intermountain region, the species discussed in this chapter have been

the most promising.

In general, species of Ceanothus have a number of

attributes that are important for revegetation. Seeds

are easily cleaned and planted. Small seedlings are

vigorous and young plants grow rapidly; only a few

other shrubs grow as quickly as Ceanothus. Species of

Ceanothus are capable of nitrogen fixation and most

are well adapted to infertile soils. Most are excellent

forage plants, yet recover well from grazing.

Family Rhamnaceae _____________

Ceanothus cuneatus

Wedgeleaf ceanothus

Description—Wedgeleaf ceanothus (fig. 36) is an

evergreen shrub (Munz 1974) distributed from Mexico

to California, Nevada, Oregon, and southern Washington (Munz 1974; Peck 1941). It is a highly palatable

evergreen species, and is heavily browsed as a winter

forage (Conard and others 1985; Gibbens and Pieper

1962; Gibbens and Schultz 1963). It is an upright

shrub ranging in height from 3 to 23 ft (0.9 to 7 m)

(Munz 1974) and forming dense thickets (Stubbendieck

and others 1986).

Ecological Relationships and Distribution—

Collections from Butte, Shasta, and Siskiyou Counties

in California, and Jackson County, OR, have been

planted throughout the Intermountain area. Selections from approximately 2,000 ft (610 m) elevation,

growing on granitic and basalt soils, have been planted

in Utah, Idaho, Nevada, Wyoming, and Colorado. This

species occurs as an understory in a number of ponderosa pine communities; outplantings have primarily

been confined to ponderosa pine or mountain brush

sites in the Intermountain area. Various collections

have also been used to treat disturbed sites in southern California (Fessenden 1979).
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 657

Chapter 23 Shrubs of Other Families

Plant Culture—Seeds of the approximately 12

collections tested in the Intermountain area express

similar features. They are small with 49,000 to 54,000

cleaned seeds per lb (108,000 to 119,000 per kg) (Reed

1974; Van Dersal 1938). Plants established in the

Intermountain area usually produce good seed crops

annually from healthy bushes. Few seedlings have

appeared from natural spread in any study site, although some sites are frequently cultivated, which

would result in seed burial.

Seeds germinate and establish well from late fall

and early winter seedings. Rodents gather ripened

seeds from trial plantings and native stands (Conard

and others 1985). Consequently, late fall plantings are

advised to reduce rodent damage to new plantings.

Wedgeleaf ceanothus seeds germinate abundantly

after fires (Reed 1974; Sampson and Jespersen 1963);

seeds also benefit after being treated with boiling

water (Grisez and Hardin 1967). Seeds placed in

boiling water for 1 to 3 minutes germinated rapidly

and more uniformly than untreated seeds. This response has been recorded for numerous collections.

Seeds that are fall seeded following hot water treatment are generally adequately wet prechilled and

germinate well in the spring.

New seedlings are quite vigorous and grow rapidly.

Growth exceeds that of big sagebrush, antelope bitterbrush, and other commonly seeded shrubs. Young

plants are able to recover following some browsing,

but Medin and Ferguson (1980) reported that developing stands established by seeding and transplanting

on winter game ranges in Idaho ultimately succumb to

continued heavy grazing by big game, small mammals, grasshoppers, and to winter injury. Plants are

very palatable (Gibbens and Schultz 1963; Scrivner

and others 1988) and are heavily used by game and

livestock.

Plantings conducted within the Intermountain area

exhibit adaptation to a wide array of sites. The species

does express broad amplitude (Detling 1961). Of the

selections being tested, few exhibit adaptation to calcareous soils common in Utah and Nevada. Plants

have established on these basic soils, but normally

succumb in 4 to 6 years. Plantings established in

central Idaho on granitic neutral or slightly acidic soils

have survived quite well. However, collections from

central California and Oregon have suffered winter

damage. Normally portions of the crown are killed in

a single season; however, some plants have persisted

for over 30 years. This species cannot be recommended

for use on wildlands outside its natural range. However, within its natural range it could be used for

watershed protection and wildlife. It can be used for

low-maintainance landscaping in areas within its natural range and on similar sites elsewhere.

Uses and Management—The species demonstrates

usefulness for seeding disturbed watersheds; it grows

rapidly and furnishes excellent ground cover. It also

provides cover on open, wind-blown game ranges. This

species’ ability to fix nitrogen also contributes to its

usefulness in range, watershed, and disturbed land

plantings.

Varieties and Ecotypes—None.

Family Rhamnaceae _____________

Ceanothus greggii

Desert ceanothus

Description—Desert ceanothus is a low-growing

to erect shrub 1 to 6.5 ft (0.3 to 2 m) tall, with thick,

persistent, entire leaves (Welsh and others 1987).

Desert ceanothus grows in mixed desert shrub communities including pinyon-juniper, ponderosa pine,

chaparral, and mountain brush sites in Nevada, California, Arizona, New Mexico, Texas, and Mexico

(Kearney and Peebles 1960; Thornburg 1982; Welsh

and others 1987). It occurs at elevations between 4,000

and 9,400 ft (1,220 to 2,870 m) in southern Utah

(Welsh and others 1987).

Ecological Relationships and Distribution—

Only a limited number of collections, primarily from

southern Utah, have been used in wildland seedings.

Collections acquired from scattered stands growing on

rocky slopes, and from nearby pure stands, have been

tested. Although this species grows on a variety of soil

types (Carmichael and others 1978; Plummer 1977;

Stark 1966; Thornburg 1982), collections obtained

from basic to coarse-textured soils have been the only

ones tested. This species is well suited to arid sites and

provides an opportunity to enhance plantings where

few shrubs are adapted and most species are difficult

to establish. Desert ceanothus has been incorporated

into a few range and wildlife improvement projects in

central and southern Utah.

Plant Culture—Seedlings establish best from late

fall and winter seedings. Seeds are small, yet larger

than most other species of Ceanothus. There are approximately 23,000 cleaned seeds per lb (50,700 per

kg) (Reed 1974). Seeds are borne in capsules that split

and disperse the seeds from the bush. Hot water

treatments effectively improve germination. Untreated

seeds require approximately 42 days of wet prechilling

to initiate germination (Stark 1966).

Good seed crops are not produced each year, even

from bushes grown under cultivation. Generally, commercially harvestable crops are produced once every 3

to 4 years. Only about one-half the seeds acquired from

arid native stands and wildlife plantings in Utah are

viable. Keeley (1977) reported similar numbers of
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Chapter 23 Shrubs of Other Families

Figure 37—Deerbrush ceanothus in south-central

Idaho plantings provide excellent wildlife habitat and

watershed protection.

viable seeds from wildland collections at other locations. Plants flower in April and May; seeds ripen in

July (Reed 1974; Van Dersal 1938) and can be hand

collected by stripping the capsules. Like other species

of Ceanothus, desert ceanothus seeds disseminate as

they ripen. Seeds must be collected when slightly

green and stored to allow drying.

Uses and Management—Desert ceanothus is a

valuable browse species for big game (Bradley 1965;

Gullion 1964; Leach 1956; Pase and Pond 1964; Powell

1988), and is a particularly important winter forage

(Conard 1987; Van Dersal 1938) and summer browse

for livestock (Boles 1987).

This shrub establishes quickly and is well adapted to

fresh disturbances. Horton (1949) reported that plants

survive well on road disturbances on coarse-textured

soils at elevations near 5,350 ft (1,630 m) in California,

and recommended planting container stock or direct

seeding in spots to control erosion. Desert ceanothus

recovers following fire because seeds stored in the soil

are fire activated. Resprouting from root crowns does

occur, but only infrequently.

Varieties and Ecotypes—None.

Family Rhamnaceae _____________

Ceanothus integerrimus

Deerbrush ceanothus

Description—Deerbrush ceanothus is a spreading, erect, deciduous shrub that grows rapidly

(Cronemiller 1959; Hitchcock and Cronquist 1973;

Munz 1974; Van Dersal 1938). Leaves are large and

oblong-ovate in shape (Cronemiller 1959); some remain on the bush throughout the winter (Cronemiller

1959; Munz 1974; Van Dersal 1938). Like other species of Ceanothus, it is able to fix nitrogen (Biswell

1960; Harvey and others 1980).

Plant Culture—Seeds ripen in midsummer and are

dispersed as the capsule dries and dehisces (Keeley

1987). The seeds are smaller than those of most other

Ceanothus species with about 70,000 cleaned seeds

per lb (154,300 per kg) (Reed 1974). Seeds are longlived and may persist in the soil for decades

(Cronemiller 1959; Pase and Brown 1982; Quick and

Quick 1961). Seedcoats are impermeable to water, and

embryo dormancy also exists (Heit 1967a). Seed germination can be enhanced by heat pretreaments

(Shmida and Barbour 1982), and is best induced by

boiling the seeds for 1 to 3 minutes followed by wet

prechilling (Quick and Quick 1961). Viability of

deerbrush ceanothus seed lots usually is higher than

for seed lots of other species of this genus. Over 85

percent of cleaned seeds are often viable (Van Dersal

1938). Empty or immature seeds are difficult to

separate from filled seeds using a gravity table or by

flotation.

Ecological Relationships and Distribution—

Deerbrush ceanothus is widely distributed in the Cascade Mountains of Oregon and Washington southward to southern California, Arizona, and New Mexico

(Conard and others 1985; Kearney and Peebles 1960;

Munz 1974; Peck 1941). It grows as the dominant

understory in a number of tree and shrub communities

(Conard 1987; Conard and others 1985; Gratkowski

1961a; Hartesveldt and others 1975). It is also well

suited to open, dry slopes with and without shade

(USDA Forest Service 1937). Based on its range of

adaptation and rapid growth, this species has been

evaluated for watershed and wildlife habitat improvement projects in central Idaho (fig. 37). Plantings have

established well from fall and winter seeding or

transplantings of bareroot stock. Young plants grow

rapidly; yearly growth rates have exceeded 30 inches

(76 cm). In addition, the shrub exhibits an ability to

persist with understory grasses and herbs. The species sprouts aggressively following clipping. Although

stem layering was reported by Biswell and Gilman

(1961), this attribute has not been observed from

plantings in Idaho. Deerbrush ceanothus grows well

on fertile soils, but collections currently under study

are not especially drought tolerant.

Uses and Management—Collections from California

and Oregon have been evaluated for their ability to

control erosion and restore wildlife habitat on logging roads and related disturbances where ground

cover is essential. The plants form extensive, fibrous

root systems when nursery grown. The root system

provides excellent soil stability, and plants effectively

stabilize erodible slopes and unstable banks. Plantings

established at numerous sites in Idaho have been
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 659

Chapter 23 Shrubs of Other Families

seriously weakened and thinned by cold winter temperatures. The plants are not cold tolerant and serious

dieback occurs each year. Less than 20 percent have

survived on these plantings. Winter losses have also

been noted from 1-year-old stock growing in nursery

beds. The shrub has failed when planted on mine

disturbances and infertile sites in Idaho, Utah, Nevada, and Wyoming, although Biswell (1960) and

Harvey and others (1980) reported the plant is able to

fix nitrogen and has been successfully planted on

disturbed sites in the Southwest. It has not done well

throughout the Intermountain region on disturbances

lacking topsoil. It does well on road fills created in the

Idaho Batholith, but not on mine wastes.

Although game animals utilize the shrub on research plantings and the plant provides considerable

cover, it is not native to the Intermountain region, and

replacement of other shrubs with this species is not

advisable. It is not sufficiently winter tolerant to

persist within the Intermountain region.

Varieties and Ecotypes—None.

Family Rhamnaceae _____________

Ceanothus lemmonii

Lemmon ceanothus

Description—Lemmon ceanothus is a low, evergreen, spreading shrub that attains a height of 1.5 to

3 ft (46 to 91 cm); leaves are alternate, elliptical, 0.5 to

1.2 inches (13 to 30 mm) long, bright green and waxy

above. Flowers are pale blue and abundant; seeds

form within capsules (Munz and Keck 1959; Sampson

and Jespersen 1963).

Ecological Relationships and Distribution—

Lemmon ceanothus is a less widely distributed shrub

than other species of Ceanothus. It is primarily confined to ponderosa pine communities from the base of

the Sierra Nevada Mountains from Tuolumne and

Eldorado Counties northward, and in the inner Coast

Ranges from Lake and Yuba Counties north to

Humboldt and Shasta Counties, California (Munz and

Keck 1959). Although regionally adapted, it is an

important deciduous shrub for wildlife (Sampson and

Jespersen 1963).

Plant Culture—Seeds of lemmon ceanothus have

an impermeable seedcoat and embryo dormancy. Heat

scarification reduces seedcoat impermeability and

hastens germination. Following heat treatment, wet

prechilling for approximately 2 months is required to

relieve embryo dormancy. Young seedlings are vigorous and can survive some herbaceous competition.

Uses and Management—Research with this species for revegetation plantings in the Intermountain

region has been limited. Plantings have been established on wildlife and watershed sites in the Interior

Western States. Plant materials acquired from northern California have been the primary source for most

plantings.

The species is best adapted to sites receiving 15 to

25 inches (380 to 640 mm) of annual precipitation. It

has not proven adapted to basic soils, and it is not

winter hardy. Less than 10 percent of all plants have

survived for more than 5 years in plantings in the

Intermountain region.

This species has received considerable use by big

game. Its rapid rate of growth is an impressive feature.

It is an attractive shrub, particularly when in bloom

(fig. 38). The flowers are abundant and cover the bush

with a dark blue blanket. Seed production is erratic.

Plants are easily cultivated and have considerable

potential for horticultural uses. They can also be used

in low maintenance plantings and on recreational

sites. This species cannot be recommended for use

outside its natural range. However, within its area of

adaptation the shrub can and should be used to restore

disturbed communities and provide wildlife forage

and watershed protection.

Varieties and Ecotypes—None.

Family Rhamnaceae _____________

Ceanothus prostratus

Prostrate ceanothus

Description—Prostrate ceanothus is a low decumbent or prostrate, evergreen, layering shrub most

prevalent in California (fig. 39). It grows 2 to 6 inches

(5.1 to 15.2 cm) in height; stems root at the nodes,

Figure 38—Lemmon ceanothus is covered by

masses of tiny blue flowers in spring.
660 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

spreading to form dense mats 2 to 8 ft (0.6 to 2.4 m)

across. Leaves are opposite, dark green, smooth above

and sometimes gray green beneath. The margins usually have coarse teeth near the tip. Flowers are blue

and borne in umbels. Fruits are roundish capsules

(Sampson and Jespersen 1963).

Ecological Relationships and Distribution—

Prostrate ceanothus occur in California on open flats

and in pine forests at elevations from 3,000 to 6,500 ft

(910 to 2,000 m) from Calaveras and Alpine Counties

northward to Modoc County, and west to Siskiyou and

Trinity Counties. It also occurs in Washington, Oregon, and western Nevada (Hitchcock and Cronquist

1973). It grows in mountain shrub, ponderosa pine,

and red fir forests of higher mountain regions, usually

forming carpets (Sampson and Jespersen 1963). A

disjunct population occurs near Council, ID.

Plant Culture—Seeds are large, ranging from

37,500 to 44,500 per lb (82,700 to 98,100 per kg).

However, they are difficult to collect from the lowgrowing plants. Seeds must be hand collected by

picking small groups of fruits before the capsules are

fully ripe. When dry, the capsules dehisce and scatter

the seeds.

Seeds persist for many years in the soil, and can be

stored under warehouse conditions for long periods

without loss of viability. Seed germination is fire

dependent. Treatment of the seeds with boiling water

for 0.5 to 3 minutes hastens germination (Reed 1974).

Seedlings develop well from fall and late winter

seeding.

Nursery-grown bareroot stock establishes well as do

rooted cuttings. Container stock can be propagated

from seed or from stem cuttings (USDA Forest Service

1976). However, transplants must be dormant when

field planted. If plants begin growth before field planting, they are susceptible to frost and drought. Seedlings and young plants are vigorous but grow slower

than most other ceanothus species. New seedlings and

transplants survive well on disturbed soils (Brown

and others 1971; Tiedemann and others 1976).

Uses and Management—Prostrate ceanothus has

been widely used for soil stabilization (Brown and

others 1971; Tiedemann and others 1976; USDA Forest Service 1976), conservation (Plummer 1977), and

horticultural plantings (Van Rensselaer and McMinn

1942), particularly in central California. Extensive

transplanting projects have been employed to treat

harsh, erodible sites. Prostrate ceanothus has attractive evergreen leaves and has been used to provide

ground cover and serve as background plants to complement floral plantings and mixed shrub beds.

Prostrate ceanothus has been successfully used to

control erosion and stabilize roadways, and logging

and mine disturbances in the Intermountain area. Its

ability to fix nitrogen (Delwiche and others 1965;

Stewart 1967) possibly contributes to its success on

infertile soils. Collections from Oregon and California

have been widely used and are well adapted to mountainous areas dominated by forested communities.

Most of these collections are tolerant of both shaded

and open planting sites. The planting stock is widely

adapted to infertile soils, exposed substrata, and mine

wastes. Townsend (1966) reported this species recovers quickly following fires, and is one of the first

species to reoccupy disturbances.

Prostrate ceanothus does not persist well with other

plants. It spreads to provide a dense ground cover

intermixed with few other plants. Prostrate ceanothus

grows very slowly, and it spreads by stem layering.

Numerous shallow roots are formed along the spreading stems. Prostrate ceanothus is able to persist and

spread when buried by unstable soil. It is particularly

useful for stabilizing steep banks, roadcuts, and unstable surfaces; however, it does grow slowly. Wildlife

have made little use of the shrub on roadway plantings

in Idaho, which allows it to be used on steep sites

where animals may cause damage. However, Sampson

and Jespersen (1963) reported that the shrub is browsed

heavily by deer in California. Kufeld and others (1973)

reported, from a summarization of studies in California, that the species is moderately used in all seasons.

This species provides attractive cover and has been

used to landscape recreational sites and wildland

areas where low maintenance is provided.

Varieties and Ecotypes—Various ecotypes and

collections of prostrate ceanothus have been developed for horticultural uses. Stock can be purchased

from wildland nurseries. A collection acquired near

Council, ID, is particularly adapted to more arid sites,

Figure 39—Plants of prostrate ceanothus form a

dense, low ground cover on an unstable granitic

slope in central Idaho.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 661

Chapter 23 Shrubs of Other Families

surviving in areas receiving 15 to 20 inches (38 to 51

cm) of annual precipitation. It does well in open areas,

and is better adapted to sites supporting other shrubs

and some herbs than most other collections.

Family Rhamnaceae _____________

Ceanothus martinii

Martin ceanothus

Description—Martin ceanothus is a low, rounded

to spreading, unarmed shrub, between 8 and 32 inches

(20 to 81cm) tall (Welsh and others 1987). The leaves

are alternate and deciduous with short petioles. Blades

are green on both sides, 0.3 to 1.2 inches (7.6 to 30.5

mm) long, 0.2 to 0.8 inch (5.1 to 20.3 mm) wide, elliptic

to oval, ovate, or obovate, and entire or serrulate

except at the base. The inflorescence is corymbose

with numerous white flowers (fig. 40). The fruit is a

three-lobed capsule 0.16 to 0.2 inch (4 to 5 mm) thick.

Seeds are shiny brown and 0.1 inch (3 mm) long.

Sutton and Johnson (1974) reported the plant is evergreen, although plants may retain their leaves until

late into winter when most are shed.

Ecological Relationships and Distribution—

Martin ceanothus is much more limited in distribution

than most other species of ceanothus in the Intermountain region. It occurs in Utah, Nevada, Colorado,

and Wyoming, at elevations between 6,000 and 9,500

ft (1,800 and 2,900 m) (Welsh and others 1987). Kearney

and Peebles (1942) reported the plant grows in the

Kaibab Plateau and Grand Canyon areas at about

7,500 ft (2,300 m).

Martin ceanothus grows in small patches or as

scattered plants in pinyon-juniper, mountain brush,

big sagebrush, ponderosa pine, Douglas-fir, aspen,

and bristlecone pine communities in Utah. Youngblood

and Mauk (1985) reported that Martin ceanothus

occurs in limited areas within the conifer forest types

of central and southern Utah. The plant is not highly

shade tolerant, but is able to persist beneath Gambel

oak, mountain maple thickets, and ponderosa pine

overstories, and it responds quickly to burning or

other methods of clearing. It is noticeably absent from

the aspen communities in Utah described by Mueggler

and Campbell (1986), possibly due to the dense overstory provided by this tree species. Where it grows

with other shrubs, particularly Gambel oak, it is most

prevalent in small, open areas where overstory species

do not occur.

At lower elevations Martin ceanothus normally grows

as scattered plants with big sagebrush, skunkbush

sumac, antelope bitterbrush, and rubber rabbitbrush.

Populations occur as somewhat isolated stands on

very harsh sites that also support green ephedra,

Wyeth eriogonum, mountain ash, and bristlecone pine.

It exists as a minor component in Gambel oak, creeping barberry, and gray horsebrush communities. It

seldom forms thickets. It often occupies ridgetops and

exposed south and west slopes where few other species

exist. It is well adapted to rocky, well-drained soils

(Van Dersal 1938), and frequently occurs as the dominant plant in these situations.

This shrub is fire tolerant, but would not be considered a seral species in most situations. Although it

recovers quickly following fire, it does not spread

rapidly to occupy extensive areas. Other mountain

brush species also recover well and are able to maintain their postfire position in these communities following burning.

Plant Culture—Plants begin growth early in spring,

and leaves normally appear by early April. Flowers

are formed in clusters on short stalks that often

overtop the leaves and foliage. Under wildland conditions not all plants flower profusely each year. Good

seed crops normally occur about once every 3 years,

but even during periods of low production, some bushes

produce considerable seed. Grazing by big game and

livestock normally restricts seed production on many

wildland sites. Plants established at protected nursery sites in Utah and Idaho have developed good seed

crops nearly every year. Plants reach seed-bearing age

in about 4 years if protected from grazing. Thereafter,

annual seed production increases for about 3 years or

until plants attain full stature.

When plants are in full bloom, the bushes are very

attractive and colorful. Flowering occurs in May and

June, and seeds mature from mid-July to mid-August

(Plummer and others 1968). Flowering generally occurs

late enough in spring that seeds are not damaged.

Flowers are insect pollinated and attract large

numbers of insects. Developing seed crops are frequently

Figure 40—Martin ceanothus receives extensive use

by wildlife and livestock.
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Chapter 23 Shrubs of Other Families

damaged by insects, causing some to abort. Seeds are

formed in three-lobed capsules that are gummy and

sticky when green. As the fruit ripens, the capsules

split open and seeds are dispersed 1 to 6 ft (0.3 to 1.8 m)

from the plant. Rodents eagerly gather the ripening

fruits and cache the seeds. The seeds or fruits are

difficult to harvest for commercial sale. Hand collection, a slow process, is required to remove the fruits

from the bush. The fruits must be collected when

slightly green before the seeds are dispersed. When

green, the fruits adhere tightly to the plant and are not

easily dislodged by beating or flailing. Once the fruit is

collected, it is air dried to allow the capsules to dehisce.

During drying, fruits should be covered with a light

screen to prevent the seeds from being thrown off the

drying tables. Seeds can be separated from the debris

using fanning mills or screens. After screening, empty

seeds can be removed on a gravity table or by floating

the seeds in water. Cleaned seeds can be stored in open

warehouses for 10 to 15 years without loss of viability

(Plummer and others 1968).

Fresh collected seeds are dormant. Germination can

be aided by immersion in hot water. The most effective

treatment has been to place the seeds in boiling water

for 1 to 2 minutes. The water is then allowed to cool to

room temperature, and the seeds are removed and

dried. Seeds are then fall seeded or wet prechilled for

approximately 30 to 90 days before germination begins.

Martin ceanothus seeds should be fall seeded in a

firm, weed-free seedbed. Seeds are relatively small,

round, and easily planted. They should be planted

0.25 to 1 inch (0.6 to 2.5 cm) deep. Rodents will gather

planted seed; consequently, late fall seeding is recommended. Seedlings appear early in the spring and are

tolerant of frost. Young seedlings grow moderately

well, but are not able to withstand competition from

herbaceous plants. Martin ceanothus should not be

seeded directly with herbs, but in separate spots or

strips. Seedlings are well adapted to harsh sites, and

survive well from plantings on well-drained, infertile

soils free of herbaceous competition.

Transplants are easily cultured as bareroot nursery

stock or container material. Bareroot and container

stock must be dormant when field planted. If plants

are lifted from the nursery bed and field planted after

vegetative growth begins, serious losses will occur.

Uses and Management—Martin ceanothus is one

of the most promising and useful shrubs for rehabilitation of big game winter ranges. It is well adapted to

big sagebrush and mountain brush communities. It

occupies some of the most critical wintering areas in

Utah, including open south and west aspects, and it

has established well when planted in these situations.

Seedlings are sensitive to drought and must be planted

in areas free of weeds. They grow at a moderate rate

and can withstand heavy grazing even as young plants.

The shrub survives well on mine wastes and associated disturbances. It spreads slowly by natural recruitment, but it is persistent and long lived. It is

adaptable to open slopes where big game may concentrate throughout the winter. The shrub has a low,

rounded growth habitat; its forage is usually available

to grazing animals.

Herbage quality is quite high, particularly for native

stands in central Utah. Deer, elk, and livestock browse

this species heavily (Diebert 1968). Tueller (1979),

however, reported only minimal use by big game in

Nevada. When planted on mine sites and on big game

habitat improvement projects in Utah, the plants are

heavily browsed by big game and livestock. Use can

often be severe enough to reduce stand establishment.

Big game animals seek areas occupied by the shrub in

late fall and winter. Both leaves and stems are grazed

during that season.

Seed production and harvesting are problems preventing the wide use of this species. Plants grown

under protection and in nursery conditions produce

abundant seed crops, but fruits are located amid the

twigs in small clusters that are difficult to reach and

detach. Plants are low growing and relatively uniform

in size and shape. Consequently, mechanical harvesters could be developed to dislodge and collect the

fruits. If such equipment were developed, use of this

species would be greatly enhanced.

Varieties and Ecotypes—Collections of Martin

ceanothus from Sanpete County in central Utah have

proven well adapted to wildlife habitats throughout

the State. This ecotype is a good seed producer, and is

heavily browsed by big game. It appears well suited to

infertile sites, including mine disturbances. Small

seed production fields have been established, but seed

yields are not sufficient to meet current demands.

Commercial production should be encouraged.

Family Rhamnaceae _____________

Ceanothus sanguineus

Redstem ceanothus

Description—Redstem ceanothus is an erect, deciduous, nonrhizomatous shrub, 5 to 10 ft (1.5 to 3.0 m)

tall (fig. 41); with purple or reddish, flexible, glabrous

twigs (Hitchcock and Cronquist 1973; Munz and Keck

1959). The leaves are thin, broadly elliptical to ovate

or obovate, rounded or subcordate at the base, 0.4 to

3.9 inches (1.0 to 10 cm) long, subglabrous, and glandular-serrulate (Munz and Keck 1959). The flowers

are numerous, white, and borne in showy compound

clusters 2 to 4 inches (5 to 10 cm) long from previous

year’s wood (Kartesz and Kartesz 1980; Munz and

Keck 1959). The fruit is a slightly lobed, 3-celled capsule

about 0.2 inch (4 mm) long with one seed per cell.
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Chapter 23 Shrubs of Other Families

Ecological Relationships and Distribution—

Redstem ceanothus is distributed from British Columbia and Montana south to Washington, Oregon, Idaho,

and California (Hitchcock and Cronquist 1973; Munz

and Keck 1959; Reed 1974). This species is best adapted

as an understory species in open or partially shaded

areas (USDA Forest Service 1937). It occurs at mid to

low elevations to 2,400 ft (730 m) in Montana (Dittberner

and Olsen 1983), and 4,000 ft (1,200 m) in California

(Munz and Keck 1959). It is most abundant throughout much of the Pacific Northwest at mid elevations in

burned forest communities (Mueggler 1965). It is a

principal component of extensive brush fields that

develop after fires, logging, or clearing (Halpern 1988).

The shrub is most prevalent in early seral stages of

grand fir, Douglas-fir, western hemlock, and mixed

conifer forests (Armour and others 1984; Conard and

others 1985; Dahlgreen 1984; Franklin and Dyrness

1973; Schoonmaker and McKee 1988). It also grows as

a scattered understory in mesic, open sites with ponderosa pine in central Idaho and the Northwest

(Franklin and Dyrness 1973). Miles and Meikle

(1984) reported that within the dry submontane

Douglas-fir subzone, redstem ceanothus favors sites

with low evapotranspiration.

Redstem ceanothus is less shade tolerant than other

species that coexist in brush fields of the Northwest

(Hooker and Tisdale 1974). It is often replaced by

oceanspray and chokecherry as tree canopies close

(Conard and others 1985), yet its presence with other

shrubs provides extensive brush fields for 25 to 50

years (Morgan and Neuenschwander 1988). Miles and

Meikle (1984) reported successional dieoff as a result

of combined factors including drought stress, severe

frost, utilization, and shade. Although the density of

this species may decline rapidly as shade increases

(Antos and Shearer 1980), it reestablishes quickly

following disturbances and becomes a dominant cover

(Zamora 1975). Nearly closed stands may develop

within 3 to 5 years (Morgan and Neuenschwander

1988).

Like other species of Ceanothus, redstem ceanothus

is primarily fire dependent for natural regeneration

(Antos and Shearer 1980; Mitchell 1983; Wittinger

and others 1977). Heat scarification stimulates seed

germination (Lyon and Stickney 1976), and new seedlings flourish following wildfires or controlled burns.

The species is capable of resprouting following burning and crown damage by deer (Miles and Meikle

1984). However, mature plants can disappear as shade

increases. Periodic reburning maintains the species as

an “obligate pioneer” (Laursen 1984). The shrub survives with 10- to 15-year intervals between burning

(Lyon and Stickney 1976).

Plant Culture—Redstem ceanothus shrubs are

normally robust and healthy plants, but abundant

seed crops are infrequent. Individual plants may develop heavy seed crops, and the better bushes are

normally “high graded” during seed collection. Most

plants are tall, and hand stripping is difficult. Fruits

are not easily dislodged from the plant by beating or

shaking the stems. Seeds are dispersed as the capsule

ripens; thus, capsules must be collected when still

green and allowed to dry. Seeds are fully developed

before the capsule dries; thus, seed quality or viability

is not adversely affected by harvesting green capsules

shortly before they dry. Rodents actively gather the

ripening seeds (Conard and others 1985) and will

collect entire seed crops as the fruits ripen.

Seedbanks develop through annual accumulation

of seeds in the soil (Kramer 1984; Morgan and

Neuenschwander 1985). Although seeds are eagerly

eaten by rodents (Conard and others 1985), numerous

seeds may accumulate in the soil and may remain

dormant for decades (Furniss and others 1978).

Seed densities ranging from 1.9 to 107 per ft2 (20 to

1,150 per m2) have been reported in Idaho (Kramer 1984).

Large amounts of redstem ceanothus seeds have not

been collected for commercial sales. The shrub usually

recovers satisfactorily by natural seedings so artificial

planting is generally not necessary. However, seeds

are easily cleaned for storage or seeding. Seeds are

approximately 0.08 inch (2 mm) in diameter; smaller

than those of most Ceanothus species. There are about

130,000 cleaned seeds per lb (287,000 per kg). Seed

cleaning is quite simple. The capsules are allowed to

air dry, causing them to break apart and propel the

seed. Consequently, the capsules should be covered

with a screen to prevent loss of seeds as fruits ripen.

Seeds and debris can be separated using an air screen

fanning mill.

Seeds have a hard, impermeable seedcoat and a

dormant embryo (Heit 1967a; Kramer 1984). They

Figure 41—Redstem ceanothus resprouts rapidly following logging in central Idaho.
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Chapter 23 Shrubs of Other Families

require a heat treatment to open the seedcoat or hilar

fissure (Furniss and others 1978). Immersing the seed

in boiling water at temperatures between 190 and

212 ∞F (88 and 100 ∞C) (Heit 1967a; Niering 1981) for

1 to 3 minutes, followed by cooling in the standing

water, reduces seedcoat imposed dormancy. Heattreated seeds require wet prechilling for approximately 60 days to enhance germination (Heit 1967a;

Reed 1974). Radwan and Crouch (1977) reported germination after 1 month of wet prechilling, but 4 months

was the optimum period. Miles and Meikle (1984)

suggested seeds collected from different aspects and

elevations have different optimum wet prechilling

periods. Seeds can be stored for many years in an

open warehouse, but may be damaged by chalcids.

Gratkowski (1973) and Radwan and Crouch (1977)

recommended storage at 37 to 69 ∞F (3 to 21 ∞C) to

reduce insect infestations.

Heat-treated seeds should be planted in fall in a

firm, well-prepared seedbed at depths between 0.25

and 0.5 inch (6.4 and 12.7 mm). Broadcast seeding on

a loose, rough seedbed or surfaces with considerable

litter also is successful. When seeding on a bare mineral surface, seeds should be incorporated into the soil

and not left on the surface. Lyon (1971), and Youngberg

and Wollum (1976) found that seedling survival by

natural recruitment was much higher following fall

burning than spring burning. Most seedlings developing from spring burning emerged a year following

the burn; thus, they were subjected to considerably

more herbaceous competition. Leege and Hickey (1975)

found that the amount of precipitation received from

May through August had a significant effect on seedling survival.

Natural recovery following wildfire is often dramatic,

as seedlings are able to establish with little site or surface

preparation. A considerable number of seedlings naturally succumb the first or second year after disturbance

(Lyon 1971; Youngberg and Wollum 1976). However,

seedlings of this species are quite vigorous and can

persist on exposed sites. High seedling losses from

natural seeding is to be expected as seedlings compete

for limited resources. Although less than 4 percent

may remain after two growing seasons (Orme and

Leege 1976), sufficient plants often survive to fully

occupy the site.

Natural thinning is not as evident where controlled

seedings are conducted. If sites are not overseeded,

little dieoff or thinning will occur. Once seedlings

attain 1 to 2 years of age, they are often of sufficient

size and vigor to persist with herbaceous competition.

Redstem ceanothus is capable of nitrogen fixation

(Torrey 1978), which enriches the biomass and density

of associated species. Thus, this shrub serves as an

important nurse plant in many situations (Rose and

Youngberg 1981; Scott 1970). It has been particularly

useful in seeding fresh disturbances where few other

species are initially able to establish. Under many

situations where little or no vegetation exists, redstem

ceanothus can be seeded with herbs to provide a mixed

composition of plants. The shrub competes with other

herbs without serious loss of young seedlings.

Transplants are easily grown as bareroot or container stock. Bareroot plantings survive as well as

container seedlings and are much cheaper and easier to

transplant. Transplanting is a useful method of establishing the shrub on selected sites and locations. Transplants grow rapidly and furnish a rapid-developing

cover, particularly if plants are inoculated with nitrogen-fixing organisms. Even when planted on mine or

roadway disturbances, redstem ceanothus is an

aggressive early developer.

Uses and Management—Redstem ceanothus is

an important forage and cover plant for big game and

other wild animals. In the Okanogan Valley of Washington and British Columbia, redstem ceanothus is

one of two browse plants that provide 60 to 70 percent

of the winter forage consumed by deer (Miles and

Meikle 1984). Although it provides 11 percent of the

available winter forage, it supplies 52 percent of the

herbage consumed by big game.

In many burned or cleared sites, redstem ceanothus

is often heavily grazed, and all annual growth is

utilized. This species is particularly important for use

by elk during the winter months (Kufeld 1973; Leege

1972; Leege and Hickey 1975). It is grazed in summer

by both deer and elk, although the amount and period

of use may vary (Conard and others 1985; Keay 1977;

Keay and Peek 1980; Thilenius 1960). Game animals

are frequently attracted to burned sites where this

species is abundant. Important wintering ranges are

dependent on burning and logging to maintain this

shrub (Leege and Hickey 1975).

Redstem ceanothus is particularly palatable to wildlife. It furnishes considerable herbage in areas where

the plant is not abundant or the dominant species.

Grazing animals seek and utilize the new growth

(Gaffney 1941; Leege 1968; McCulloch 1955; Thilenius

1960). Scattered plants growing in riparian areas and

forest openings are selectively used at all seasons.

The age of the plant and seasonal period of grazing

affect its palatability. Leege (1968) reported that new

growth is much more palatable than older twigs, and

following burning, the nutritional value decreases

with age (Asherin 1973; Leege 1969; Leege and Hickey

1971). However, Miles and Miekle (1984) did not

recommend burning to regulate or enhance the forage

value of this shrub.

The protein content of this plant during winter is

about 10.1 percent; this provides the protein level

necessary for deer maintenance (Miles and Miekle
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Chapter 23 Shrubs of Other Families

1984). In addition, the average fiber content is a

relatively low 34.7 percent. Livestock also are attracted to plants with high nutritive quality and make

considerable use of this species.

Redstem ceanothus provides important cover for

wildlife including big game, small mammals, and

birds. It is particularly important in winter. It is also

an important ground cover, and provides soil stability

on burned slopes where erosion potential is high. It

provides protection for streambanks and moist areas.

Following fires it recovers quickly and provides ground

cover and soil stability on areas where slumping and

surface runoff are common.

Seedling establishment is very good when redstem

ceanothus is planted on disturbances; consequently,

this species is utilized to seed abandoned roadway and

logging disturbances. Both direct seedings and

transplantings have been successful in conservation

plantings to control erosion. The species, however, is

not well adapted to mine wastes, and has been slow to

invade road and logging disturbances where vehicle

activity has occurred and soils are compacted. It has

been very useful for restoring riparian disturbances,

and withstands livestock use. However, grazing of

young plants can weaken developing stands.

This shrub is able to establish with seeded herbs. It

can be seeded with herbs or transplanted into new

grass/forb seedings. It can be used as a pioneer species

to restore harsh sites, and provides favorable areas for

other plants to establish.

Varieties and Ecotypes—None.

Family Rhamnaceae _____________

Ceanothus velutinus

Snowbrush ceanothus

Description—Snowbrush ceanothus is a lowspreading, many-stemmed, evergreen shrub (fig. 42)

(Countryman 1982; Welsh and others 1987; Zamora

1982) with olive-green to reddish brown bark. It attains heights of 3 to 9 ft (0.9 to 2.7 m) when growing in

thickets or closed stands (Sutton and Johnson 1974).

On open, windswept sites it may grow as low, rounded

plants usually about 3 ft (0.9 m) tall; however, on

fertile sites individual shrubs may be 12 ft (3.7 m) tall

(Stanton 1974).

Leaves are 1.2 to 2.0 inches (3 to 5 cm) long with

three main veins and a glossy dark-green surface

(Miles and Miekle 1984; Stark 1966). They are alternate, thick, pubescent to glabrous, ovate to elliptic,

and very fragrant (Sutton and Johnson 1974; Welsh

and others 1987). The leaves often curl in drought

conditions; this conserves water by reducing evapotranspiration (Miles and Miekle 1984).

The flowers are white, heavily scented, very abundant, and borne in more-or-less corymbose clusters on

short branches (Mozingo 1987; Sutton and Johnson

1974). The fruit is a three-lobed capsule, with a single

seed per locule. The seeds are small and dark brown or

tan with shiny coats (Welsh and others 1987).

The plant forms a deep spreading root system, and

nitrogen-fixing actinomycetes occur on the roots

(Hickey and Leege 1970; Stanton 1974). Snowbrush

ceanothus does not spread by layering or root suckering,

but by resprouting from the crown. The plants have a

distinct cinnamon odor if the leaves or stems are

crushed (Welsh and others 1987).

Ecological Relationships and Distribution—

Snowbrush ceanothus grows from the coastal ranges

of California north to British Columbia and eastward

to Alberta, Montana, South Dakota, and Colorado

(Mozingo 1987; Reed 1974). Hitchchock and Cronquist

(1973) reported that C. v. var. hookeri occurs on the

west side of the Cascade Mountains from northern

California to British Columbia. East of the Cascades,

var. velutinus occurs in California, Nevada, Utah,

Colorado, and South Dakota. In Utah, Wyoming, and

Colorado, the species is less abundant and restricted

to more specific communities. Goodrich and Neese

(1986) reported the plant occurs primarily at elevations between 7,000 and 9,000 ft (2,100 and 2,700 m)

in the Uinta and Blue Mountains and the west portion of the Tavaputs Plateau, where it is mostly

confined to mountain brush, ponderosa pine, and

aspen communities.

This shrub has a broad ecological amplitude and is

considered a late seral species in some communities,

and an early seral species in others. It is most often

codominant with ponderosa pine and Douglas-fir. It

Figure 42—Snowbrush ceanothus is a common understory shrub in many ponderosa pine communities.
666 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

is also commonly encountered in various mountain

brush, antelope bitterbrush, and forested shrub communities. It is a principal species in forested communities in Oregon and Washington (Franklin and

Dyrness 1973) and Idaho (Steele and Geier-Hayes

1987, 1992). As an early seral species, it occupies

Douglas-fir, white fir, grand fir, ponderosa pine, and

lodgepole pine forests (Dyrness 1973; Franklin and

Dyrness 1973; Halpern 1989; Schoonmaker and McKee

1988).

Although snowbrush ceanothus commonly grows on

dry, open hillsides (Weber 1987), and is considered to

be shade intolerant (Stanton 1974), it will persist as an

understory in forested communities for considerable

periods. Increasing closure of the canopy and competition affect removal of the shrub. The rate of succession

is regulated by soil moisture, and snowbrush ceanothus

can maintain a presence on xeric sites as an edaphic

climax community (Miles and Miekle 1984). The shrub

may persist as a midseral species with Douglas-fir, but

if tree stands are somewhat open, it may remain as a

permanent understory (Conard and others 1985).

The species responds well to disturbances, particularly fires (Halpern 1988). High-intensity fires are often

followed by the rapid recovery and postfire dominance

of this plant, usually within 1 year, if an adequate

seedbank exists and climatic conditions favor establishment (Halpern 1989; Halvorson 1982). Seed germination is activated by burning (Halpern 1988; Marshall

and Waring 1984; Mozingo 1987). This species is able to

compete with pioneering herbs. Snowbrush ceanothus

is, therefore, a major component of brush fields and

shrubby communities that develop after wildfires

(Gratkowski 1961b, 1978; Morris 1958). It is often a

primary understory component on ponderosa pine sites.

It may exist as a long-lived seral species or a climax

dominant. Along with antelope bitterbrush, it forms

important associations as a dominant climax shrub

(Franklin and Dyrness 1973).

Snowbrush ceanothus grows across a wide elevational range from 3,500 to over 10,000 ft (1,100 to

3,000 m) (Mozingo 1987; USDA 1937). Aspect affects

the distribution of the species due to increasing evapotranspiration demands on south and west slopes,

particularly in pioneering systems (Miles and Miekle

1984). The shrub is unable to establish on south,

southwest, or west aspects at elevations below 3,250 ft

(990 m) in British Columbia because of xeric conditions. At high elevations it is not normally found on

steeper east, northeast, or north aspects because of

poor shade tolerance and competition (Miles and Miekle

1984).

Although snowbrush ceanothus is a fire-dependent

pioneer species, successional dieoff is caused primarily by drought stress due to competition. Shading,

frost damage, and utilization may hasten the process

(Miles and Miekle 1984). Severe frosts frequently

damage the shrub, particularly during years of low

snow cover when the shrubs are exposed. Although

plants may be killed to ground level, regrowth from the

crowns can usually occur. Fires can damage and kill

mature plants, but regrowth does occur, and stand

regeneration is often accomplished with controlled

burns.

Snowbrush ceanothus exists on a wide number of

soil types and parent materials. It is able to grow on

well-drained, coarse-textured soils (Stephens 1973),

but water-holding capacity and nutrient availability

greatly affect the presence of this shrub (Miles and

Meikle 1984). Soils that contain sufficient moisture to

sustain growth into July and August are important to

its survival. Snowbrush ceanothus occurs on slightly

acidic and neutral soils, and is particularly adapted to

soils derived from granitic parent materials (Sutton

and Johnson 1974; Watson and others 1980;

Zavitkovski and Newton 1968).

Plant Culture—Plant growth and flowering are

regulated by elevation. Vegetative growth normally

begins by mid-April at lower elevations and as late as

early June at higher sites (Noste and Bushey 1987).

Flower development begins early in the season, often

by mid-April. At higher elevations flowering may

occur in June or July. Seed maturation occurs from

July to August (Plummer and others 1968; Reed 1974;

Stark 1966; Swingle 1939).

Plants usually produce a number of flowers each

year, but viable seeds may not always mature. Seed

production varies annually among wildland sites. Few

areas produce abundant crops each year, but some

shrubs will generally produce seed. Young plants that

occur on open burn sites are not always better seed

producers than those existing in mixed shrub/conifer

forests. Normally, some sites will bear commercially

harvestable crops each year, but sites must be located

and inspected regularly.

Seed is harvested by hand stripping the fruits before

they mature or the capsules dry. As capsules dry,

seeds shatter immediately. Capsules ripen irregularly

on the bush and fruits must be collected when still

green. However, seeds are fully ripe before the capsules are dry, and harvesting of slightly green fruits

does not diminish seed quality. Seed harvesting is

slow and expensive, and is a principal deterrent to the

wide use of this species.

Yields of 50 to 75 lb per acre (56 to 84 kg per ha) of

cleaned seed have been obtained from sites in Idaho.

Rodents actively gather the fruits and seeds, and will

remove much of the seed as it matures. Nearly complete removal of the seed by rodents has been reported

by Conard and others (1985).

Mature seeds have a hard impermeable seedcoat

and a dormant embryo. Heat and wet prechilling are

necessary for germination (Mozingo 1987). Soaking
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Chapter 23 Shrubs of Other Families

the seed in boiling water for 1 to 3 minutes normally

enhances germination. Seeds are then germinated in

a cool, moist environment at 34 to 41 ∞F (1 to 5 ∞C) for

63 to 84 days (Reed 1974). Germination is usually

completed in about 14 days (Quick 1935). Seed germination is usually very high, exceeding 70 to 80 percent

(Reed 1974). Mozingo (1987) suggested that the optimum period of exposure to heat varies geographically

among ecotypes, but differences have not been noted

in separate collections obtained from central Idaho.

Seeds can be stored for long periods without loss of

viability. Good stands have developed from planting in

the fall at a depth of 0.5 to 0.75 inch (1.3 to 1.9 cm).

Seedings also establish well by broadcasting seeds on

a rough seedbed in the fall or early winter. Plantings

on bare mineral soil or amid surface litter are both

successful if the soil surface remains wet throughout

the germination period. Miles and Miekle (1984) reported that natural seedings that germinate in the

autumn, following late summer or fall burning, survived much better than those that germinate in spring

after spring burning.

Snowbrush ceanothus seedlings establish well on

fresh, open disturbances and compete favorably with

pioneering herbs. The species establishes very well

from artificial seedings and is particularly adapted to

unstable sites. Sutton and Johnson (1974) and Miles

and Meikle (1984) reported young plants grow slowly,

but Scott (1970) found that nodulated seedlings produced 2.5 times more dry weight than non-nodulated

plants. Results from plantings conducted in Idaho and

Utah do not support reports that seedling growth is

slow. In fact, plantings on forested roads and study

sites are vigorous and aggressive. Seedlings and young

plants develop well and are able to establish and

persist with herbaceous competition. However, considerable differences between nodulated and non-nodulated plants were also noted on these sites.

A large number of natural seedlings can be expected

to emerge on burn sites. Natural thinning normally

occurs soon after emergence (Lyon 1971), but sufficient numbers usually survive to provide an adequate

stand. Winter frost, grazing, and lack of soil water

contribute to seedling losses.

Snowbrush ceanothus is easily grown as bareroot or

container stock, and transplanting is an excellent

method for stabilizing disturbances. Snowbrush

ceanothus forms a symbiotic relationship with Streptomyces ceanothii to fix nitrogen (Hickey and Leege

1970). These organisms form clusters of modified

short roots, not typical nodules (Furman 1959; Torrey

1978). Seedlings grown for field plantings should be

inoculated with these nitrogen-fixing organisms.

Uses and Management—Snowbrush ceanothus is

a particularly important species for restoring and

enhancing soil fertility. Annual fixation of nitrogen

ranges from 21 to 62 lb per acre (23.5 to 69.5 kg per ha)

(Rose and Youngberg 1981; Tiedemann 1981;

Zavitkovski and Newton 1968). Nodulated plants can

improve growth of associated species. Scott (1970)

reported Douglas-fir trees growing in the open were

half the size of seedlings nourished by snowbrush

ceanothus.

Snowbrush ceanothus has been particularly valuable as a pioneer species for improving poor soils and

disturbed sites. It has been extensively used to revegetate logging roads, mine wastes, and other disturbances. It is particularly adaptable as a pioneer species capable of stabilizing and improving soil

conditions that can then be used to enhance the

establishment of associated species (Conard and others 1985; Geier-Hayes 1987; Steele and Geier-Hayes

1992). The shrubs improve soil fertility, which promotes early establishment and growth of other plants

(Franklin and Dyrness 1973; Steele and Geier-Hayes

1987, 1992).

This shrub has been one of the most successful

species used to revegetate roadways and unstable

slopes in the Idaho Batholith, where erosion control is

needed during the year of disturbance. Plants establish quickly, and small seedlings or transplants grow

rapidly to furnish excellent ground cover. The plant

can be seeded in mixtures with fast-developing herbs

including western yarrow, fireweed, and Canada goldenrod, or it can be transplanted at various spacings to

reduce wind erosion and runoff.

Ecotypes with different growth habits have been

selected and used for erosion control. Plants obtained

from windswept sites have dense, low-growing growth

forms, and have demonstrated adaptability to a wide

range of sites. These collections maintain a low, dense,

leafy growth habit that provides excellent ground

cover. One-year-old bareroot stock normally develops

a crown spread of 18 to 30 inches (46 to 76 cm) during

the year of establishment.

Plantings established along roadways have persisted following wildfires, and their extensive root

system stabilizes roadfills. Regrowth occurs quickly

and new shoots protect the soil surface. The evergreen

growth habit of this shrub also aids in furnishing

yearlong soil cover. The plant has been widely planted

for erosion control in the West (Dietz and others 1980;

Fessenden 1979; Monsen and Christensen 1975;

Plummer 1977; Stark 1966).

Snowbrush ceanothus provides forage and cover

for game and wildlife. It is regarded as a primary

winter forage for big game, but use varies considerably

among geographic regions (Martinka 1976; Stanton

1974; USDA Forest Service 1976). Undoubtedly,

differences occur among ecotypes, and collections from

some regions are much more heavily grazed than

others. Miles and Miekle (1984) reported that
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Chapter 23 Shrubs of Other Families

snowbrush and redstem ceanothus are the most important winter browse species for mule deer in the

Okanogan Valley in British Columbia, and that 60 to

70 percent of the forage consumed in winter is supplied

by these two shrubs. Plants in Idaho (Leach 1956),

portions of Montana (Klebenow 1965; Youds and

Herbert 1988), and some collections from central California received moderate to heavy use. Within the

Great Basin this species receives much less use by big

game (Tueller 1979). Although plants are browsed in

winter, use is not excessive. Collections of this species

have been assembled and field planted at various

locations in Idaho. Collections from California, northern Idaho, and Oregon have received more use than

most Intermountain region accessions. Use also varies

by season (Martinka 1976; McCulloch 1955) and among

big game animals (Kelbenow 1965; Peek 1974; Watson

and others 1980).

Snowbrush ceanothus withstands considerable

winter browsing and is able to recover without

apparent loss in vigor. Snow cover and winter frost

damage reduce both forage availability and animal

use. Many sites are often covered with deep snow,

and plants are unavailable for long periods (Miles

and Miekle 1984).

Cattle generally make little use of this shrub (Curtis

1952). However, Miles and Miekle (1984) noted that

they selectively used it in late fall and winter, thus

competing with big game for winter browse. Sheep and

goats browse new growth after fire (Stanton 1974).

Nutrient value varies among sites. In British Columbia, Miles and Miekle (1984) found snowbrush and

redstem ceanothus were the only two native shrub

species that provided maintenance level protein for

deer in the winter months. Digestible energy was

higher for snowbrush ceanothus than for all other

native shrubs. The winter fiber content of snowbrush

ceanothus was 26 percent, lower than any other shrub

tested (Miles and Miekle 1984). This is due to the

retention of leaves on the bush. This provided a much

higher amount of forage with higher nutritive content

than that of deciduous shrubs (Miles and Miekle

1984). In contrast, Dittberner and Olsen (1983) described protein and energy values for this shrub as

poor. Blank (1984) and DeByle and others (1989)

suggested that fire increases nutritive content, but

Miles and Miekle (1984) concluded that the changes do

not support the use of fire for nutrient enhancement.

The plant has considerable value in landscape and

conservation plantings. Its growth habit, color, and

evergreen features provide attractive cover. The plant

is suited for extensive cultivation and will survive in

organized plantings. In addition, it can be used in

low-maintenance situations, and is particularly useful for restoring disturbances where native species

are desired.

Varieties and Ecotypes—None. However, ecotypes

with low growth forms, and selections that receive

heavy browsing have been identified. These selections

could quickly be advanced with more extensive seeding projects.

Family Salicaceae _______________

Salix species

Willow

Introduction—There are about 300 species of willows worldwide; about 70 species are native to North

America (Brinkman 1974i). Willows are constituents

of many riparian zones in the Western United States.

Their root systems stabilize streambanks, while stems

and leaves dissipate flood energy and catch sediments.

Willows contribute to the formation of overhanging

banks that provide shade, cover, food, and travel

corridors for a large number of vertebrates and invertebrates and microsites for establishment of many

plant species (Clary and McArthur 1992; Kovalchik

and Elmore 1992; Thomas and others 1979b).

Damage to many riparian ecosystems has resulted

from improper grazing practices, logging, road construction, recreation, water impoundments, and other

human activities. Land management efforts have

recently centered on restoration and maintenance of

these dynamic ecosystems. Research and riparian

classification studies are rapidly providing knowledge

of willow ecology and guidelines for management

(Hansen and others 1988a,b, 1995; Kovalchik 1987;

Manning and Padgett 1995; Padgett and others 1989;

Szaro and Patten 1986; Youngblood and others 1985).

Natural regeneration processes and techniques for

restoration of riparian communities have also received recent attention. Improvement of degraded

riparian areas requires that factors causing degradation be addressed (Briggs 1995; Briggs and others

1994). Restoration efforts are unlikely to be successful

unless the causes of degradation are first mitigated.

Most willow species are easily propagated; but little

information is available for many individual species.

General techniques for vegetative propagation and

production from seed are provided below. Specific

requirements, where known, are described in the

plant culture chapter for each species.

Vegetative Propagation—Most willows can be

propagated from hardwood cuttings. Techniques for

harvesting and preparing cuttings are provided in

Chapter 29 and in many books on plant propagation.

Stems of most willows contain dormant, preformed

root primordia. These are formed during the first year

and quickly develop into roots when conditions are

favorable (Haissig 1970, 1974). Upland species such as
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Scouler willow do not produce root primordia. They

root more slowly, and root initials appear only at the

base of the cutting (Densmore and Zasada 1978).

Cuttings should be collected on or near the planting

site whenever possible. Released varieties of some

willow species are available from commercial nurseries, and can be used if restoration using local material

is not an objective or if local material is unavailable

(Carlson 1992; Darris and Lambert 1993; USDA Soil

Conservation Service 1993a,b).

Unrooted cuttings may be used on sites with favorable planting conditions and long growing seasons.

Dormant cuttings may be taken from fall through

early spring. Densmore and Zasada (1978) found that

cuttings taken in early spring rooted more readily

than cuttings harvested in late fall. Rooted cuttings

may be required for species that do not produce preformed root primordia and for plantings on sites with

short growing seasons, rapidly declining water tables,

or other factors that might slow establishment.

Rooted and unrooted cuttings should be planted in

areas appropriate for each species. Cuttings or rooted

stock should be planted at depths permitting the

developing root system to remain in contact with the

water table throughout the summer (Busch and others

1992). Plantings may be conducted in fall when water

levels are low and appropriate planting areas exposed.

This aids planters in placing the cutting base or root

system at an adequate depth and permits early-spring

root growth, but it also exposes cuttings or plants to

spring runoff and flooding. Some plantings must be

completed after spring runoff. Streams with artificially manipulated flows present special problems, as

root growth may not be rapid enough to advance with

a rapidly declining water table.

Survival and growth of willow plantings may require

that vegetative competition be removed or reduced at

the time of planting. Neiland and others (1981) found

that competition with tall, dense grass stands adversely

impacted survival. Svejcar and others (1992) and

Conroy and Svecar (1991) obtained greater secondyear survival of Geyer willow cuttings planted on bare

ground than on Nebraska sedge/Baltic rush sites even

though depth to the water table in midsummer was

less on the Nebraska sedge/Baltic rush sites. They

interpreted these results to represent an interaction of

biotic and abiotic factors on willow establishment.

Large planting stock may be used when depressed

water tables or browsing are problems (York 1985).

Planting poles or dormant stubs permit the base of the

cutting to be placed in the water table; foliage is

produced beyond the reach of browsing animals, and

cover and soil protection are obtained quickly. However, pole plantings are costly, and natural regeneration is generally unlikely to occur unless the water

table is restored or browsing curtailed.

Willows may also be used to stabilize eroding

streambanks and other disturbances. Such practices

as wattling, brush matting, and branch packing are

used to provide physical stability as vegetative cover

develops (Gray and Leiser 1982; USDA Soil Conservation Service 1992). Live and dead plant materials including stems and branches are buried, planted, or

placed on the soil to reduce soil movement, protect

erosion control structures, and enhance vegetation

development. Live willow cuttings used in these procedures often root, providing dense root systems and cover.

Propagation From Seed—Willows are dioecious;

male and female flowers are borne on separate plants.

Inflorescences are catkins or aments that, depending

on the species, appear before, with, or after the leaves

(Brunsfeld and Johnson 1985). The flowers are pollinated by bees. Fruits are capsules; each contains

numerous tiny, hairy seeds. Seeds of most species

ripen and are rapidly dispersed in early to midsummer

(Brinkman 1974i). Dispersal may be timed to coincide

with low water levels. Seeds are carried over long

distances by wind and water. Hairs on the seeds may

aid in dispersal; they may also function in the germination process (Martens and Young 1992). Summerdispersed seeds may be viable for as little as 1 week

(Densmore and Zasada 1983). They are nondormant

and germinate rapidly, often in 24 to 48 hours, if

deposited on wet, exposed substrates. Germination

occurs rapidly over a wide range of temperatures. A

requirement for exposure to light may ensure that

germination occurs in areas free of vegetative competition. The ability to germinate over a wide range of

temperatures may compensate for the short period of

viability (Densmore and Zaszada 1983). Seed coats are

transparent, and embryos contain chlorophyll. Consequently, germination and seedling development are

rapidly initiated under favorable circumstances.

Wide dispersal enables willows to act as pioneer or

early successional species following flooding, burns, or

other disturbances that create suitable microsites for

germination. Continued rapid growth enables seedlings to develop root systems that remain within reach

of the declining water table in summer, compensate

for the late period of germination, and withstand

flooding during the succeeding spring.

Some arctic and high-elevation willows such as

grayleaf willow mature seed in summer, but delay

dispersal until fall (Densmore and Zasada 1983; Zasada

and Viereck 1975). These seeds exhibit conditional

dormancy; they are capable of germinating at high

temperatures, but not at the lower ambient temperatures of the season. Embryos of late-dispersing species

generally contain low amounts of chlorophyll. Wet

overwinter chilling of these seeds releases dormancy,

permitting seeds to germinate soon after snowmelt
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Chapter 23 Shrubs of Other Families

and maximizing the period for seedling development

during the short growing season.

Willow seeds may be hand harvested once the capsules begin opening, or recently dispersed seed may be

collected from drifts (Brinkman 1974i). Use of a commercial shaker has been suggested for harvesting

yellow poplar seed; the feasibility of using this system

for collecting willow seed has not been assessed (Cech

and Keys 1987). However, because ripening within

stands, plants, and catkins is uneven, it is often advantageous to harvest entire catkins just as the capsules

turn from green to yellowish or as the first capsules

begin opening (Brinkman 1974i; Martens and Young

1992). Catkins can then be spread to dry in an enclosed

area until the capsules open. With either method, it is

imperative that maturation be monitored closely.

Seeds may be planted without cleaning, other than

removal of twigs, leaves, and catkin branches. Martens and Young (1992) suggested that hairs not be

removed from the seeds; their function is not fully

understood. However, the hairs do cause the seeds to

cling together. They may be removed by gentle carding

(Atchley 1989) or by forcing air through a series of

sieves containing the seeds (Fung and Hamel 1993;

Harder 1970). Both techniques are rather slow if large

quantities of seed are to be cleaned.

Brinkman (1974i) reported willow seed could be

stored for only 10 days at room temperature, but

viability could be maintained for 1 month if seed is

stored wet in sealed, refrigerated containers. Zasada

and Densmore (1977, 1980) found that seed of several

summer and fall-dispersing willows could be stored

for up to 3 years if dry seed was placed in doubled 3-mil

polyethylene bags and stored at 14 ∞F (–10 ∞C).

Seeds may be sown directly into containers or onto

wet soil surfaces. In the bareroot nursery, seed may be

covered lightly using a roller (Brinkman 1974i). It is

essential that the soil surface be kept wet and shaded,

if possible. Seeds may be broadcast on appropriate

sites if surfaces remain wet. Little information is

available on production of planting stock from seed or

direct seeding. Use of seed may be advantageous if

local cutting material is not available. The ability to

store seed for up to 3 years provides some flexibility in

planning propagation and in timing direct seeding.

Family Salicaceae _______________

Salix bebbiana

Bebb willow, beaked willow

Description—Bebb willow is a thicket-forming

shrub or small tree growing to 33 ft (10 m) in height.

Roots are shallow and dense. Twigs are reddish or

grayish brown and hairy when young. Leaves are

elliptic, obovate or oblanceolate, mostly 1.2 to 2.4

inches (3 to 6 cm) long and 0.4 to 1.2 inches (1 to 3 cm)

wide with entire to shallowly toothed margins. Upper

surfaces are green, and lower surfaces are glaucous

with prominently raised veins. Catkins emerge and

mature with the leaves. Staminate catkins are 0.4 to

0.8 inch (1 to 2 cm) long. Pistillate catkins are whitish

yellow and 0.6 to 2.4 inches (1.5 to 6 cm) long. Floral

bracts are narrow, pale yellow to light brown, and

sparsely to densely hairy. Capsules are very loosely

arranged and borne on stipes 0.08 to 0.16 inch (2 to 4

mm) long. They are ovoid conic, 0.2 to 0.3 inch (5 to 8

mm) long, short hairy, and long beaked. Stigmas are

two lobed (Brunsfeld and Johnson 1985; Hitchcock

and others 1964; Welsh and others 1987).

Plants begin flowering when 2 to 10 years old;

optimum seed producing age is 10 to 30 years (Hansen

and others 1988a,b; Rawson 1974). Bebb willow flowers from April to July or August (Gill and Healy 1974;

ISTA 1966; Viereck and Little 1972). Fruits ripen and

seeds are dispersed from May to August (Densmore

and Zasada 1983; ISTA 1966).

Ecological Relationships and Distribution—

Bebb willow is widespread in Canada and the United

States, occurring from low to upper-middle elevations

(Brunsfeld and Johnson 1985; Hitchcock and others

1964). It does not occur west of the Cascade Mountains

or in California. At low elevations Bebb willow grows

in broad meadows and on alluvial terraces and

subirrigated slopes. It is most common on organic or

mineral soils with textures ranging from silty to sandy

or gravelly (Marchant and Sherlock 1984), and it will

tolerate moderately alkaline to moderately acidic soils

(pH 5.5 to 7.5) (Haeussler and Coates 1986). Water

tables are usually within 3.3 ft (1 m) of the soil surface

throughout the summer. Mottled or gleyed soils are

common in the first 3.3 ft (1 m) (Hansen and others

1995). At upper elevations Bebb willow is found in

drier riparian areas.

Uplands associated with Bebb willow may be dominated by Wyoming big sagebrush, mountain big sagebrush, Douglas-fir, or Engelmann spruce communities. Aspen, black cottonwood, and water birch are

common overstory associates (Brunsfeld and Johnson

1985; Padgett and others 1989).

Bebb willow may dominate early seral willow communities along rivers and streams, overflow channels,

or seeps (Boggs and others 1990). Seeds germinate on

wet mineral substrates with full exposure to sunlight.

Suitable seedbed microenvironments include recent

alluvial deposits along streams and abandoned, siltfilled beaver ponds (Hansen and others 1988a). Seedlings colonize wet, exposed mineral soils following

fires and have been noted to invade mine spoils in

southwestern Canada (Marchant and Sherlock 1984;

Watson and others 1980). Burned or otherwise damaged trees resprout readily. Plants may also develop
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Chapter 23 Shrubs of Other Families

from detached branch segments (Watson and others

1980).

Bebb willow communities are relatively stable once

established. They may persist for long periods due to

the presence of high water tables or short periods of

flooding (Hansen and others 1988a,b). Bebb willow

requires full sun and lacks shade tolerance (Watson

and others 1980); it may be replaced by taller species.

Plant Culture—Bebb willow may be propagated as

rooted cuttings, but reports of rooting capability vary.

In Alaska, cuttings harvested in autumn after leaf fall

or in spring following leaf expansion failed to root

(Densmore and Zasada 1978). In contrast, rooting

success of current year’s softwood, also harvested in

Alaska, was 42 percent (Halloway and Zasada 1979).

In Montana, Atchley (1989) obtained 40 percent rooting when cuttings of 4-year-old wood were harvested

in May. Platts and others (1987) reported dormant

cuttings rooted readily. Growth was initiated in about

10 days and shoot growth in 10 to 20 days. Roots were

formed along the entire length of the stem.

For production of rooted cuttings in Saskatchewan,

Cram (1976) recommended that 6-inch (15-cm) cuttings be harvested in early fall and immediately placed

in cold storage at 34 ∞F (1 ∞C) for spring planting in a

bareroot nursery. Rooted cuttings were subsequently

outplanted as 1-0 stock. Watson and others (1980)

reported 30 to 70 percent survival of 1-0 rooted hardwood cuttings on mine overburden in northern Alberta;

most mortality resulted from rodent damage. Unrooted

cuttings are sometimes used. Hansen and others

(1988a,b, 1995) recommended use of early springharvested dormant cuttings 12 to 20 inches (30 to 50

cm) long with diameters greater than 0.4 inch (1 cm).

Bebb willow seeds must be harvested immediately

when mature; seeds are dispersed in a period of 1 or 2

days (Marchant and Sherlock 1984). Fruits are green

when preripe and yellowish when ripe (Watson and

others 1980). Seeds may be hand harvested from trees

or from recently dispersed drifts. Seeds may be used

with or without cleaning (Brinkman 1974i). Atchley

(1989) found that carding was more effective than the

air-blowing technique described by Harder (1970) for

removing the dense hairs from the seeds. Seeds vary in

size and shape. Atchley (1989) found that viability was

greater for plump, green seeds compared to small

dark-brown seeds or wrinkled seeds. There are about

2,500,000 cleaned seeds per lb (5,500,000 per kg)

(Brinkman 1974i). Bebb willow seeds collected in

Alaska can be stored dry for at least 3 years in doubled

3-mil polyethylene bags at 14 ∞F (–10 ∞C) (Zasada and

Densmore 1980).

Fresh seeds collected in Alaska are nondormant.

Germination at temperatures from 41 to 77 ∞F (5 to

25 ∞C) exceeded 90 percent (Densmore and Zasada 1983;

Zasada and Viereck 1975). This adaptation permits

the short-lived seeds to germinate in highly variable

environments during the short Alaskan growing season. By contrast, freshly harvested Montana seeds

were highly germinable at 59 to 77 ∞F (15 to 25 ∞C) and

less germinable at 41 to 50 ∞F (5 to 10 ∞C) (Atchley

1989). Thus, germination may be inhibited by cool

seedbed microenvironments. Bebb willow seedlings

can tolerate some shade (Atchley 1989).

Uses and Management—Watson and others

(1980) described Bebb willow as being a rapid grower,

capable of recovering rapidly after browsing, moderately aggressive on suitable sites, a relatively good soil

stabilizer, and exhibiting low drought tolerance. Stands

stabilize streambanks, providing protection from

flooding. Bebb willow often occurs in clumps rather

than continuous canopies (fig. 43). Thus, it provides

shading for fisheries while permitting good access to

recreational fishing sites. Campgrounds cannot be

established within Bebb willow sites because the soils

are generally wet early in the season.

Figure 43—Thickets of Bebb willow dominate many

riparian communities from low to upper mid elevations in western mountain ranges.
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Chapter 23 Shrubs of Other Families

Figure 44—Shrubby Booth willow is overshadowed by

treelike whiplash willow.

Bebb willow provides a major source of highly productive and palatable browse for moose, deer, rabbits,

elk, beaver, and bighorn sheep (Marchant and Sherlock

1984; Watson and others 1980). It is more tolerant of

browsing than Geyer willow, Booth willow, yellow

willow, and Drummond willow, and may increase at

their expense following heavy browsing (Hansen and

others 1995). Use of Bebb willow increases in late

winter. It remains accessible in deep snow as branches

are bent down and become accessible to browsing

animals (Dorn 1970; Viereck and Little 1972). Song

birds and game birds use the species for nesting and

food.

Bebb willow communities are often adjacent to

meadows grazed heavily by livestock (Boggs and others 1990; Dorn 1970). Trees are usually widely spaced,

allowing for easy access (Tesky 1992). Consequently,

deferring grazing until sites are dry may be required

to reduce livestock trampling and soil compaction

(Boggs and others 1990). Heavy livestock use may

restrict Bebb willow seedling establishment, resulting

in stand degeneration.

Prescribed burns can be used to rejuvenate degraded Bebb willow stands (Hansen and others 1988a,b;

Tester and Marshall 1962). Burned plants sprout

rapidly from basal stems. The small seeds are dispersed over considerable distances (Haeussler and

Coates 1986; Kovalchik and others 1988). Establishment from seed depends on timing of the fire and

availability of wet mineral soils to support germination and seedling establishment (Chrosciewicz 1988;

Viereck and Little 1972; Viereck and Schandelmeirer

1980).

Bebb willow is used as a “diamond willow” for carving

canes, lampposts, and furniture (Watson and others

1980). Wood of this species has also been used for

baseball bats, charcoal, and gunpowder (Viereck and

Little 1972) .

Varieties and Ecotypes—“Wilson” Bebb willow

was released in 1985 by the Alaska Plant Materials

Center (Wright 1989). It exhibits wide adaptability

and a dense growth habit; it is useful for windbreaks, living fences, and screens.

Family Salicaceae _______________

Salix boothii

Booth willow

Description—Booth willow is a highly branched,

rounded shrub with numerous basal stems 10 to 20 ft

(3 to 6 m) tall. Young twigs are yellow to dark brown

and hairy, becoming glabrous at maturity. Mature

leaves are elliptical to broadly lanceolate, thick, firm,

and about 0.8 to 3 inches (2 to 8 cm) long. Margins are

entire to toothed with fine, gland-tipped teeth. Both

leaf surfaces are moderately pubescent when young.

Upper surfaces of mature leaves are green; lower

surfaces are paler, but not glaucous. Catkins expand

with or slightly before the leaves. Staminate catkins

are 0.4 to 1 inch (1 to 2.5 cm) long; pistillate catkins are

0.8 to 1.6 inches (2 to 4 cm) long. Floral bracts are

brown to black and pubescent with long, curly, tangled

hairs. Capsules are borne on stipes less than 0.08 inch

(2 mm) long. They are 0.12 to 0.24 inch (3 to 6 mm)

long, oblong ovate, and glabrous. Stigmas are nearly

entire (Brunsfeld and Johnson 1985; Hitchcock and

others 1964; Welsh and others 1987).

Ecological Relationships and Distribution—

Booth willow occurs at midelevations from British

Columbia to Alberta and south to northeastern California and Colorado (Brunsfeld and Johnson 1985). It

may dominate or codominate early seral willow communities in fens, bogs, abandoned beaver ponds, wet

meadows, and riparian sites, ranging from recently

exposed rocky or gravelly deposits with high water

tables to drier benches with deep sandy to clay-loam

soils and water tables to 3.3 ft (1 m) below the soil

surface (Brunsfeld and Johnson 1985; Manning and

Padgett 1995; Padgett and others 1989; Youngblood

and others 1985). Booth willow will tolerate moderately alkaline, but not strongly acidic or basic soils

(Haeussler and others 1990).

Booth willow often grows in association with Geyer

willow and whiplash willow in open bottom lands

(fig. 44) and with Drummond willow in forested

stream bottoms. Understory species range from

beaked sedge and Nebraska sedge in wetter communities to various forbs and shrubs in drier areas.

Kentucky bluegrass may be a dominant understory
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Chapter 23 Shrubs of Other Families

species in areas with histories of grazing disturbances and dry surface soils (Hansen and others

1988a,b). Uplands associated with Booth willow

support sagebrush, juniper, western spruce-fir,

lodgepole pine, Douglas-fir, and alpine meadow communities (Esser 1992).

Booth willow resprouts following burning or mechanical damage. Resprouting is more vigorous if stem

removal occurs during the dormant season (Kovalchik

and others 1988). Booth willow is also tolerant of frost

and flooding. Efficiency of gas exchange and root

regeneration are increased by the presence of aerenchyma; adventitious rooting may occur above the level

of flooded soil (Kovalchik 1992). However, growth is

reduced by prolonged flooding above the level of the

root crown.

Flowering occurs from May to June. Fruits ripen

and seed is dispersed in late July and early August

(Haeussler and others 1990; Zasada and Viereck 1975).

Greatest quantities of seeds are produced by plants

that are 2 to 10 years old (Brunsfeld and Johnson 1985;

Haeussler and others 1990). Booth willow regenerates

from short-lived seeds that are dispersed by wind or

water. Suitable microsites for germination include

wet sand or gravel surfaces receiving full sunlight

(Hudak and Ketcheson 1992).

Plant Culture—Booth willow may be used to revegetate degraded riparian areas. Cuttings root readily

and abundantly along the lower one-third of the stem

(Platts and others 1987). New roots and stems are

initiated in 10 to 15 days. Seedlings must be protected

from browsing for at least 3 years to ensure establishment (Kay and Chadde 1992).

Fresh seeds harvested in Alaska germinated rapidly

and nearly completely at constant temperatures

ranging from 41 to 77 ∞F (5 to 25 ∞C) (Densmore and

Zasada 1983). This behavior may be an adaptation to

a short period of viability, the short growing season,

and extreme variability in microsite conditions.

Uses and Management—Booth willow provides

browse for moose, elk, deer, small mammals, birds,

and other wildlife (Chadde and Kay 1988; Manning

and Padgett 1995; Youngblood and others 1985). Excessive browsing can reduce Booth willow regeneration and seed production (Kay and Chadde 1992).

Booth willow communities provide cover for many

terrestrial species and shade and overhanging banks

for fish (Brunsfeld and Johnson 1985; Manning and

Padgett 1995). Streams in Booth willow stands may

provide recreational fishing, but access is restricted by

the dense canopies.

Palatability of Booth willow is fair for livestock and

sheep; use increases late in the season (Manning and

Padgett 1995; Padgett and others 1989; Youngblood and

others 1985). Geyer willow, which is often abundant or

dominant in Booth willow communities, is generally

more palatable. Livestock use of both species should be

restricted until soils are dry to avoid compaction

(Hansen and others 1988a,b). Production of Booth

willow stands is often high, while understory production is variable, depending on the amount of willow

canopy present. Heavy use on sites with fine soils and

palatable understory species must be avoided to preclude soil erosion and bank sloughing.

Prescribed burning may be used to rejuvenate decadent stands and increase sprouting for use by big game

(Haeussler and others 1990; Manning and Padgett

1995). Plants also reestablish from offsite seed sources

if burning occurs prior to seed dispersal and fires are

severe enough to expose mineral seedbeds (Densmore

and Zasada 1983; Zasada and Viereck 1975).

Varieties and Ecotypes—None.

Family Salicaceae _______________

Salix drummondiana

Drummond willow, beautiful willow, blue

willow

Description—Drummond willow is a multiplestemmed shrub usually 10 to 13 ft (3 to 4 m) tall. Young

twigs are finely hairy and yellow or green, later becoming glabrous and purple brown to yellow with a bluish

glaucous bloom. Mature leaves are elliptic to lanceolate or oblanceolate, 1.6 to 3.5 inches (4 to 9 cm) long

and 0.5 to 0.8 inch (1.3 to 2 cm) wide. Margins are

entire and somewhat revolute. Leaf surfaces are dark

green above and glaucous to strongly silvery pubescent beneath. Catkins expand before or with the leaves;

flowering generally occurs in May (Dittberner and

Olson 1983; Munz 1973). Staminate catkins are 0.4 to

1 inch (1 to 2.5 cm) long. Pistillate catkins are 0.6 to 1.6

inches (1.5 to 4 cm) long, densely flowered, and silvery

pubescent. Floral bracts are black or dark brown, long

hairy, and persistent. Capsules are 0.12 to 0.16 inch

(3 to 4 mm) long and densely short hairy. Stigmas are

lobed (Brunsfeld and Johnson 1985; Hitchcock and

others 1964; Sutton and Johnson 1974; Welsh and

others 1987).

Ecological Relationships and Distribution—

Drummond willow occurs from Yukon and

Saskatchewan south to California and New Mexico

and east across southern Canada and the Northern

United States. It occurs at mid to high elevations from

the upper sage to the spruce-fir zone (Brunsfeld and

Johnson 1985; Hansen and others 1995; Manning and

Padgett 1995). It is found around springs, seeps,

beaver ponds, lakes, ponds, and on flood plains and

benches adjacent to swift, rocky, high gradient streams

(fig. 45). At the lower edge of its range it generally
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Chapter 23 Shrubs of Other Families

grows in wet, sandy or gravelly soils adjacent to

streams. In cooler habitats at higher elevations it is

more abundant and occurs across broad valley bottoms. In such areas it grows on moist, well-aerated

mineral soils ranging from sandy to clayey loams. The

water table may be near the surface in spring, but it

drops below 3.3 ft (1 m) by late summer. Common

associates are Booth willow and Geyer willow. Due to

the similarity of their ecological requirements, community types in Utah, western Wyoming, and eastern

Idaho dominated by either Drummond willow or Booth

willow are classified as Booth willow types (Padgett

and others 1989; Youngblood and others 1985). However, Goodrich (1992) noted that the dominance of this

species and absence of Booth willow along rocky, highelevation streams of the Uinta Mountains indicates a

need to describe one or more riparian community

types in which Drummond willow is the dominant

woody species.

Plant Culture—Drummond willow is useful for

stabilization of disturbed sites adjacent to streams

and ponds. The species may be established from

unrooted or rooted cuttings. Unrooted cuttings are

most successful if planted on sites that remain wet

throughout the growing season. Two to 4-year-old

hardwood cuttings harvested in early spring root

readily with root initials appearing along the length of

the stem. Roots and shoots appear about 10 days after

planting (Platts and others 1987). Seed biology of

Drummond willow has not been studied. Seeds ripen

in summer and remain viable for about 1 week (Uchytil

1991a).

Uses and Management—Drummond willow communities are long lived, but shade intolerant. Productivity is generally high. Stands provide stream stabilization, dense shade for fish, and habitat for beaver,

birds, and other wildlife.

Moose and elk make heavy use of Drummond willow, particularly in winter (Peek 1974; Singer 1979;

Stevens 1970). Plants in Yellowstone National Park

have been stunted by moose and elk use (Chadde and

Kay 1988; Patten 1968). In Oregon, Drummond willow

is considered highly palatable to livestock, big game,

and beaver (Kovalchik and others 1988). Heavy browsing by big game may lead to degradation or loss of this

species. Loss of willows resulting from heavy use by

beaver may result in an increase in graminoids, loss of

beaver populations, and lowering of the water table.

Drummond willow communities stabilize streambanks and limit livestock access (Hansen and others

1988a,b). Heavy livestock use or trailing at streamside, particularly when soils are wet, however, may

lead to highlining, dead clumps, stand loss, and bank

compaction and sloughing. Decadent willows can recover rapidly if browsing is reduced. Excessive livestock grazing may also lead to a loss of palatable

graminoids and their replacement by less palatable

weedy species. Drummond willow stands may act as

natural firebreaks in wet years, but during dry seasons they may burn (Crane 1982). Decadent Drummond

willow stands may be rejuvenated using prescribed

burning in late summer or early fall when sites have

dried. Burning near streambanks should be carefully

regulated. Livestock grazing must be curtailed for at

least 2 or 3 years following burning. Plants recover by

resprouting from the root crown following fire (Boggs

and others 1990; Kovalchik and others 1988). Seed

dispersed from unburned areas may germinate on

exposed, wet mineral soils (Viereck 1982; Viereck and

Schandelmeier 1980). Recovery from seed depends on

seed availability, season of burning, and the presence

of open, wet mineral soils.

Varieties and Ecotypes—“Curlew” Drummond

willow was cooperatively released by the USDA Soil

Conservation Service Pullman Plant Materials Center

Figure 45—Drummond willow borders a high

gradient stream in eastern Utah.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 675

Chapter 23 Shrubs of Other Families

in 1993 (USDA Soil Conservation Service 1993a).

Curlew is valuable for inclusion in conservation

plantings for streambank stabilization, erosion control, improvement of wildlife habitat, naturalized landscaping, and shelterbelts. It grows on wet, sandy to

gravelly sites in areas receiving 20 to 25 inches (51 to

64 cm) of precipitation. The release originates from a

native population growing near the Curlew River,

Ferry County, Washington. It is propagated from seed

or cuttings. Hardwood cuttings taken in late winter

are most commonly used.

Family Salicaceae _______________

Salix exigua

Coyote willow, dusky willow, narrowleaf

willow, sandbar willow

Description—Coyote willow is a colonial shrub

that spreads underground to form dense thickets.

Stems develop from shoot buds on lateral roots. Stems

are numerous, slender, and up to 16 ft (5 m) tall. Young

twigs are thinly to densely hairy; older ones are glabrous and brown to reddish brown. Mature leaves are

linear to oblong, entire to minutely toothed, 0.8 to 4.3

inches (2 to 11 cm) long, and 18 to 30 times longer than

wide. Leaf surfaces are gray green to silvery white,

and glabrous to densely pubescent. Catkins terminate

the twigs and emerge and mature with or after the

leaves. Staminate catkins are yellowish and 0.6 to 1.8

inches (1.5 to 4.5 cm) long. Pistillate catkins are 0.6 to

2.4 inches (1.5 to 6 cm) long. Floral bracts are yellowish

and hairy. Capsules are 1.6 to 2.8 inches (4 to 7 cm)

long, sessile, and glabrous to hairy. Stigmas are deeply

lobed (Brunsfeld and Johnson 1985; Hitchcock and

others 1964; Welsh and others 1987).

Flowering occurs from April to July (Dittberner and

Olsen 1983). Fruits mature and seeds are dispersed

from May to July (Brinkman 1974i; Noble 1979; Ware

and Penfound 1949). Seed dispersal may correspond to

low water levels (Youngblood and others 1985).

Ecological Relationships and Distribution—

Coyote willow is widespread in North America, but

occurs entirely east of the Cascade Mountains at low,

or occasionally midelevations from the sagebrush to

the spruce-fir zone. Coyote willow is one of the earliest

pioneer species to colonize freshly deposited sand and

gravel bars, which may be below the high water mark

where it is subject to annual flooding, ice jams, and

associated scouring and deposition (Hansen and others 1988a,b; Wasser and Hess 1982). It frequently

occurs along streams and rivers in bottom lands,

forming a narrow zone between the water’s edge and

adjacent cottonwood, water birch, thinleaf alder, or

other willow communities (Kovalchik 1987; Szaro and

Patton 1987; Youngblood and others 1985). It also

grows on rocky, gravelly, and sandy lake and pond

edges; wet, well-drained alluvial terraces and bottom

lands that may have fine-textured soils (fig. 46); along

irrigation ditches; and in wet areas along roadways

(Brunsfeld and Johnson 1985). Soil surfaces in communities at the water’s edge are often characterized by

high percentages of exposed areas and rocky surfaces,

and provide seedbeds for establishment of a wide

array of annual and perennial herbaceous species

(Youngblood and others 1985). Sandbar willow may

continue to dominate repeatedly disturbed areas, regenerating by rootsprouting or establishing from seed.

Stabilized communities of this shade-sensitive species

may eventually be replaced by cottonwoods or other

willow species.

Two subspecies of coyote willow are widespread in

the Intermountain area; considerable variability occurs within each subspecies. Salix exigua ssp. exigua

occurs primarily in the Wyoming big sagebrush zone of

foothill areas and is distributed from southern British

Columbia and Alberta to northern Mexico. This subspecies is characterized by entire or few-toothed,

gray-green to silvery pubescent leaves; late-developing catkins; and narrow, acute floral bracts. S. e. ssp.

melanopsis is distributed from Alberta to California

and south to northern Colorado, and occurs at higher

elevations than ssp. exigua (Hitchcock and others

1964). Subspecies melanopsis is distinguished by its

toothed to subentire leaves that are glaucous beneath,

glabrous capsules, and blunt floral bracts (Brunsfeld

and Johnson 1985).

In addition to spreading by suckering, coyote willow

produces numerous small seeds that are dispersed by

wind and water (Arno and Hammerly 1977; Mozingo

Figure 46—Coyote willow forms a dense thicket

along a dry streambed.
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Chapter 23 Shrubs of Other Families

Figure 47—Geyer willow forms clumps or corridors

that provide cover and travel routes for big game.

1987). Seeds germinate within 24 hours if exposed to

freshly deposited, wet alluvium in full sunlight

(Densmore and Zasada 1983). In the field, seeds generally do not remain viable for more than 1 week (Ware

and Penfound 1949). Branches are flexible and resprout

if buried in sediment. Plants may also regenerate

vegetatively from broken stems and roots deposited on

exposed, wet soils.

Plant Culture—Coyote willow is an excellent species for restoration of disturbed riparian areas and

recently deposited sediments because of its early successional status, rapid growth, and ability to root

sprout (Uchytil 1989a). Unrooted cuttings may be

used successfully on low-elevation sites where the

water table remains high throughout the growing

season. Rooted stock should be used on sites with

fluctuating water tables, high flooding potentials, or

short growing seasons. Coyote willow cuttings develop

roots along the entire length of the stem (Platts and

others 1987). Roots and shoots appear in about 10

days.

There are about 10,000,000 cleaned seeds per lb

(22,000,000 per kg) (Brinkman 1974i). At 72 ∞F (22 ∞C),

Brinkman (1974i) obtained 83 percent germination in

4 days.

Uses and Management—Maintenance of coyote

willow stands is important because of their ability to

stabilize streambanks (Uchytil 1989a). Loss of this

species as a result of browsing and human-caused

disturbances can lead to serious erosion problems

(Hansen and others 1988a,b).

Coyote willow is browsed by moose, elk, and to a

limited extent, mule deer (Patten 1968; Van Dersal

1938). It is heavily used by beaver (Mozingo 1987).

Thickets of sandbar willow provide valuable cover for

birds and other wildlife species (Lindauer 1983). However, because of their upright growth habit, they

provide only moderate amounts of shade for fish

(Hansen and others 1988a,b). Dense stands of coyote

willow bordering streams may limit access by fishermen. As such stands are seasonally flooded, they

should not be used for campgrounds or roadways

(Hansen and others 1988a,b). Coyote willow is used for

forage and especially for cover by livestock (Van Dersal

1938). Forage production is low to moderate (Hansen

and others 1988a,b).

Coyote willow communities often act as firebreaks

due to their occurrence on sites with high water tables

or near streams, but they may burn during unusually

dry years. Burned plants may resprout from roots and

spread from offsite seeds dispersed to wet mineral

soils (Conrad 1987; Rowe and Scotter 1973; Zasada

1986).

Varieties and Ecotypes—“Silvar” coyote willow

was cooperatively released by the USDA Soil

Conservation Service, Pullman Plant Materials Center in 1993 (USDA Soil Conservation Service 1993b).

The release originated from material collected on the

Tucannon River, near Starbuck, Washington, at an

elevation of 560 ft (170 m). It is recommended for use

in conservation plantings for riparian area stabilization and restoration, wildlife habitat improvement,

and shelterbelts. It is normally propagated from cuttings, but seed can be used.

Family Salicaceae _______________

Salix geyeriana

Geyer willow

Description—Geyer willow is a large shrub or

small tree with numerous slender, ascending stems 10

to 13 ft (3 to 4 m) in height arising from a tight basal

cluster (fig. 47). The root system is deep, fibrous, and

spreading (Sutton and Johnson 1974). Young twigs

are densely hairy, but become glaucous; older twigs
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Chapter 23 Shrubs of Other Families

are brownish purple and glabrous. Early leaves of the

season are glabrous above with long, silky hairs beneath. Mature leaves are lanceolate to elliptical, entire to inconspicuously toothed, 1.2 to 3 inches (3 to

8 cm) long, and 0.3 to 0.5 inch (8 to 12 mm) wide.

Surfaces are silvery to gray green above, glaucous

below, and hairy on both sides. Catkins expand with

the leaves on short, leafy branchlets. Staminate catkins are yellow to reddish and 0.3 to 0.6 inch (0.7 to 1.5

cm) long. Pistillate catkins are globose, reddish, 0.4 to

0.6 inch (1 to 1.5 cm) long, and loosely flowered. The

persistent floral bracts are sparsely hairy and yellow

to brown or blackish. Capsules are hairy, ovoid oblong,

and 0.1 to 0.2 inch (3 to 6 mm) long. Stigmas are nearly

entire (Brunsfeld and Johnson 1985; Hitchcock and

others 1964).

Ecological Relationships and Distribution—

Geyer willow occurs at low to upper elevations from

British Columbia to Montana and south from California

to Colorado. It grows in wet meadows and marshes,

beaver ponds, valley bottoms along meandering

streams, and on dry stream benches from the sagebrush to the spruce-fir zone (Brunsfeld and Johnson

1985; Uchytil 1991b). It most commonly grows on finetextured silty to clay-loam soils that may contain some

cobbles or gravels and have considerable organic material and mottling near the surface (Padgett and

others 1989; Youngblood and others 1985). Plants

usually grow in clumps, but may form a more or less

continuous corridor near streamside. Communities in

broad valleys are relatively stable and maintained by

seasonal flooding and high water tables within 3.3 ft

(1 m) of the surface. Associated willow species often

include Bebb willow, Booth willow, yellow willow,

plainleaf willow, Drummond willow, and Lemmon

willow. The understory often includes a moderate to

dense cover of graminoids and forbs.

Geyer willow flowers from May to August (Dittberner

and Olson 1983; Munz 1973). In eastern Oregon seed

dispersal begins in early July (Padgett 1981). Seedlings colonize wet, exposed, well-aerated mineral soils,

but can withstand anaerobic conditions (Manning and

Padgett 1995).

Plant Culture—Either unrooted or rooted Geyer

cuttings may be used to revegetate disturbed sites.

Cuttings form roots in about 10 days and shoots in

about 10 to 15 days (Platts and others 1987). Conroy and

Svejar (1991) and Svejcar and others (1992) found that

establishment of Geyer willow was best if unrooted

cuttings were planted with the base within 12 inches

(30 cm) of the midsummer water table. Waterlogged

conditions had no effect on growth of the cuttings.

Uses and Management—Geyer willow is used by

moose and elk, especially in winter (Chadde and Kay

1988; Gaffney 1941; Padgett and others 1989). It is

more palatable than Drummond willow, Wolf willow,

and Booth willow (Boggs and others 1990). Geyer

willow clumps and corridors provide excellent cover

and travel routes for big game. Beaver use it for food

and building material (Allen 1983). It also provides

food and cover for birds and small mammals (Argus

1957). Roots of Geyer willow growing along streams

contribute to formation of overhanging banks; the

canopy provides shade for fish.

In eastern Oregon palatability of Geyer willow to

livestock was rated as moderately high (Kovalchik

and others 1988). Willows and understory species in

these communities are often highly productive and

may receive heavy use (Hansen and others 1988).

Geyer willow communities provide easy access to livestock. Early season use by wild ungulates, livestock, or

vehicles may adversely impact wet organic soils (Manning and Padgett 1995). Excessive use may lead to

decreased willow vigor or stand loss, replacement of

understory graminoids with less desirable species,

lowering of the water table, erosion, and conversion to

drier community types (Kovalchik 1987). Decadent

Geyer willow can recover if released from browsing

pressure.

Plants recover from burning or mechanical damage

by sprouting or possibly by establishment of seedlings

from offsite seed sources (Kovalchik and others 1988).

Fires are infrequent in these areas, but may occur in

dry years. Prescribed burning may be used to rejuvenate decadent stands. Areas must be protected from

browsing until the stands recover (Boggs and others

1990; Kovalchik 1987).

Varieties and Ecotypes—None.

Family Salicaceae _______________

Salix glauca

Grayleaf willow, glaucous willow

Description—Grayleaf willow is a low shrub 1 to 4

ft (0.3 to 1.2 m) tall. Young twigs are grayish and hairy,

becoming reddish brown and glabrous as they mature.

Bark of older plants may be rough and furrowed.

Mature leaves are lanceolate to elliptical, entire to

minutely serrulate, and up to 2.2 inches (5.5 cm) long

and 0.3 to 0.9 inch (0.7 to 2.2 cm) wide. Mature leaf

surfaces are pale green above and glaucous beneath.

They are grayish hairy when young, becoming glabrous as they mature. Catkins expand with the leaves

and remain on the plant through summer. Staminate

catkins are 0.6 to 1.6 inches (1.5 to 4 cm) long. Pistillate catkins are 0.6 to 2 inches (1.5 to 5 cm) long and

densely flowered. Floral bracts are persistent, hairy,

and pale brown to black. Capsules are 0.2 to 0.3 inch
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Chapter 23 Shrubs of Other Families

Figure 48—Grayleaf willow occurs in alpine areas

and exhibits a low, spreading growth habit.

(5 to 7 mm) long, long beaked, and densely hairy,

becoming nearly glabrous as they mature. Stigmas are

bilobed (Brayshaw 1976; Brunsfeld and Johnson 1985;

Hitchcock and others 1964; Viereck and Little 1972).

In Alaska and Yukon, grayleaf willow flowers in

June. Fruits ripen in July and August and seeds are

dispersed in late August and September (Densmore

and Zasada 1983; Viereck and Little 1972).

Ecological Relationships and Distribution—

Grayleaf willow is a circumboreal species, extending

south in North America to alpine and subalpine sites

in eastern Idaho, Utah, Montana, Wyoming, Colorado,

and northern New Mexico. In the Intermountain area

it occurs along streams and in other wet places, on

talus slopes, and in snow concentration areas in alpine

meadows and forest openings (Padgett and others

1989; Welsh and others 1987). It is found on soils

ranging from well drained to poorly drained and usually waterlogged (Alaska Rural Development Council

1977). A low-growing form occurs in open, alpine

situations (fig. 48); plants in somewhat protected,

subalpine environments are more erect and shrublike

(Dorn 1970). An early successional species, grayleaf

willow pioneers fresh alluvial deposits, burns, and

other disturbed areas with exposed mineral soils

(Viereck and Little 1972).

Densmore and Zasada (1983) and Zasada and

Viereck (1975) reported that grayleaf willow produces large numbers of small, lightweight seeds that

remain on the plant through summer and are dispersed in fall. These seeds remain viable through

winter and germinate in spring following snowmelt;

this permits seedling development to occur over the

entire growing season. Spring ripening populations,

however, have been reported (Watson and others

1980). Grayleaf willow often becomes highly abundant following fire, establishing from seed sources in

unburned areas (Foote 1983). Seedlings have been

noted to invade disturbed sites in Arctic areas

(Densmore and Zasada 1983; Viereck and Little 1972).

Grayleaf willow sprouts following mechanical damage or burning (Haeussler and Coates 1986). It also

spreads vegetatively through rooting of horizontal

stems (Sampson and Jones 1977).

Plant Culture—Densmore and Zasada (1978) found

that untreated cuttings of 2-year-old wood, taken in

autumn following leaf fall or in spring following leaf

expansion, produced few roots. Cuttings treated with

rooting hormones produced few to moderate numbers of

roots along the entire length of the cutting (Platts and

others 1987). Roots and stems appeared in about 10

days. Cuttings have been used to plant unstable sand

dunes in northern Alberta (Hardy BBT Limited 1989).

Seed may be used to propagate planting stock. Direct

seeding may also be practical (Uchytil 1992).

Densmore and Zasada (1983) and Zasada and Viereck

(1975) found that fresh seeds were conditionally

dormant. Germination increased with constant incubation temperatures from 41 to 77 ∞F (5 to 25 ∞C). Wet

chilling for 90 days released dormancy; germination

was initiated during wet chilling, and nearly all seeds

germinated when incubated at constant temperatures

in the 41 to 77 ∞F (5 to 25 ∞C) range. Environmental

conditions may influence the level of seed dormancy;

this was indicated by variability in germination of

seeds by year of collection and geographic origin.

Germination also varied among seeds from different

catkins on the same plant.

Seeds may be stored at 14 ∞F (–10 ∞C) in doubled 3-

mil polyethylene bags for up to 3 years without loss of

viability (Zasada and Densmore 1980; Zasada and

Densmore 1977). The initial dormancy of some seed

lots is lost in storage.

Use and Management—Grayleaf willow is browsed

by moose, but it is generally less palatable than other

co-occurring willows (Milke 1969). Caribou make use

of the species, primarily in summer; snowshoe hare

use is concentrated in winter (Cody 1965; Smith and

others 1988). Nutrient quality of grayleaf willow for

ungulates is good in winter (Risenhoover 1987; Scotter

1972), but lower growing forms may be covered by

snow. Plants are tolerant of heavy browsing. The cover

value of grayleaf willow for large ungulates is also

limited by its low-growing habit, but dense stands

provide good cover for small animals (Uchytil 1992).

Erosion control value of grayleaf willow is considered moderate due to its intermediate growth rate

(Alaska Rural Development Council 1977). Established plants, however, provide good soil stabilization.

Varieties and Ecotypes—There are no releases.
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Chapter 23 Shrubs of Other Families

Family Salicaceae _______________

Salix lasiandra var. caudata

Whiplash willow

Description—Salix lasiandra plants growing in

the Intermountain region belong to S. l. var. caudata,

whiplash willow. Whiplash willow ranges from 10 to

20 ft (3 to 6 m) in height (fig. 49). It is treelike at

lower elevations, becoming shrubby at higher elevations. Young twigs are finely hairy, becoming glabrous

and yellowish with age. Large branches are yellow to

red or brown in winter, while small branches are

orangish (Sutton and Johnson 1974). Mature leaves

are lanceolate to elliptic, widest at or below the middle,

and serrulate. Leaf surfaces are green and glabrous.

The lower surface is paler than the upper, but not

glaucous. Petioles often bear glands on the upper side

near the junction with the leaf blade. Catkins expand

with the leaves. Staminate catkins are white to yellow

and 0.6 to 1.8 inches (1.5 to 4.5 cm) long, with three to

five stamens. Pistillate catkins are pale green to yellow or white and 0.8 to 2.8 inches (2 to 7 cm) long.

Floral bracts are deciduous, yellow green, hairy on the

lower part, and glabrous distally. Capsules are narrowly ovate, glabrous, and 0.16 to 0.3 inch (4 to 8 mm)

long. Stigmas are lobed. Flowering occurs in April or

May, and fruiting and seed dispersal from June to

August (Brinkman 1974i; Hitchcock and others 1964).

Pacific willow (S. l. var. lasiandra) occurs mostly

west of the Cascade Mountains, but also in wetter

portions of eastern Washington, northern Idaho, and

northwestern Montana. It is distinguished by the

glaucous lower surface of its leaves (Brunsfeld and

Johnson 1985).

Ecological Relationships and Distribution—

Whiplash willow is distributed from Alaska to

Saskatchewan and south from southern California to

New Mexico at low to mid elevations (Welsh and

others 1987). Associated uplands support communities ranging from Wyoming big sagebrush to pinyonjuniper, mountain big sagebrush, and Douglas-fir

(Uchytil 1989b). Whiplash willow is commonly a pioneer or early seral species on alluvial sands and

gravels adjacent to fluctuating streams and rivers or

abandoned river channels (Brunsfeld and Johnson

1985). Stands are maintained by periodic flooding.

Whiplash willow may maintain its dominance on stabilized sandbars, as they develop into benches within

the flood plain, but it is eventually replaced by later

seral willow or cottonwood species (Manning and

Padgett 1995). It often occurs with coyote willow,

another pioneer species; they form intermittent linear

communities between the high water level and later

successional woody species growing further from water. Whiplash willow exhibits high tolerance to deposition and flooding and low tolerance of drought and

salty soils (USDA Soil Conservation Service 1992). Its

productivity is generally low due to repeated flooding.

Whiplash willow spreads primarily from seeds that

are dispersed by wind and water (Zasada 1986). Broken twigs or branches of whiplash willow transported

downstream and deposited on wet alluvial surfaces

may sprout, developing new plants vegetatively (Argus

1973).

Plant Culture—Whiplash willow is widely used for

stabilizing streambanks due to its early seral status.

It may also be used on appropriate sites to provide

naturalized landscaping, screens, or windbreaks.

Unrooted or rooted cuttings may be used. Unrooted

cuttings are most successful in areas with high water

tables throughout the summer and a long growing

season. Whiplash willow roots readily along the entire

length of the stem. Roots initiate growth in about 10

days and shoots in 10 to 15 days (Platts and others

1987).

Figure 49—Pistillate catkins of whiplash willow are

conspicuous, white to green, and appear with the

leaves.
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Chapter 23 Shrubs of Other Families

Figure 50—Arroyo willow grows along washes, ditches,

and well-drained sites along ephemeral streams.

Seeds are green when preripe and yellowish when

ripe (Brinkman 1974i). There are about 11,500,000

cleaned seeds per lb (25,300,000 per kg). Brinkman

(1974i) reported 25 percent germination at 72 ∞F

(22 ∞C) in 3 days. Densmore and Zasada (1983) found

that fresh seed collected in Alaska germinated rapidly

and nearly completely at constant incubation temperatures ranging from 41 to 77 ∞F (5 to 25 ∞C).

Uses and Management—Whiplash willow colonizes and stabilizes recent alluvial deposits. Roots

provide streambank stability; the canopy provides

shade for fish. Consequently, management to maintain these stands is critical. Whiplash willow provides

valuable habitat for deer and nongame birds (Argus

1973; Arno and Hammerly 1977; Bernard and Brown

1977; Gray and Greaves 1984). Although palatability

of whiplash willow for wildlife is generally considered

poor or fair, it is an important browse for mule deer

(Sampson and Jespersen 1963; USDA Forest Service

1937). Beaver use it heavily in winter (Kindschy 1985).

Palatability of whiplash willow is greater for sheep

than for cattle. On wet sites, healthy whiplash willow

stands are not easily accessed by livestock because of

the dense overstory. Therefore, these stands are only

moderately susceptible to overgrazing, trampling,

and compaction (Manning and Padgett 1995).

Communities with coarse, dry surface soils and limited herbaceous cover are more vulnerable to heavy

grazing. With prolonged heavy use, willow recruitment is decreased, mature plants are highlined, and

plants become decadent or die. Willows may be replaced by more mesic shrubs, introduced grasses such

as Kentucky bluegrass, or weeds (Hansen and others

1995). With reduction or elimination of browsing,

existing plants often recover quickly.

The response of whiplash willow to burning has been

poorly documented, and its resprouting capability is

not known. Windblown seeds play an important role in

reestablishing the species in burned or disturbed areas

(Miller and Miller 1976; Shaw and Clary 1996; Zasada

1986).

Varieties and Ecotypes—“Nehalem” Pacific willow is a male clone released by the USDA Soil Conservation Service Corvallis Plant Materials Center in

1978 (Darris and Lambert 1993). It originated from a

native population growing near the Nehalem River in

northwestern Oregon. Nehalem was selected for its

high basal stem density and attractive foliage. It may

be used to stabilize streambanks, improve riparian

and aquatic habitat for wildlife and fish, and provide

natural area landscaping and screens. Nehalem is

adapted to planting sites on sandbars, lakeshores, and

riverbanks at elevations below 4,900 ft (1,500 m) on

the west side of the Cascade Mountains from Washington to northern California. It may be planted on

soils ranging from sandy loams to gravelly or rich,

rocky soils where soil water is adequate. Nehalem is

tolerant of flooding and may be planted in reservoir

drawdown areas. Stands may be established from

cuttings; suitable materials may be used for wattling,

brush matting, and branch packing to provide slope

protection.

“Roland” whiplash willow was released in 1985 by

the Alaska Plant Materials Center (Wright 1989). It is

used for landscaping and stream revegetation and

protection throughout Alaska.

Family Salicaceae _______________

Salix lasiolepis

Arroyo willow

Description—Arroyo willow is a shrub or small

tree 13 to 20 ft (4 to 6 m) tall (fig. 50) with multiple

stems, a large root crown, and a fibrous root system.

Young twigs are hairy and yellowish olive to reddish.

Mature leaves are coriaceous, oblong or oblanceolate,

and entire or rarely minutely serrate with somewhat

revolute margins. They are 0.6 to 1.6 inches (1.5 to 4.1

cm) long and 0.2 to 0.5 inch (5 to 13 mm) wide. Leaf

surfaces are dark green, glabrous above, and glaucous

beneath. Catkins expand before to slightly after the

leaves. Staminate catkins are 0.9 to 1.8 inches (2.2 to

4.5 cm) long. Pistillate catkins are densely flowered

and 0.9 to 1.8 inches (2.2 to 4.5 cm) long. The persistent

floral bracts are purplish black and densely hairy.

Capsules are ovate, glabrous, and 0.1 to 0.2 inch (3 to

5 mm) long (Hitchcock and others 1964; Welsh and

others 1987).

In northern Arizona, arroyo willow flowers from

mid-March to mid-April (Sacchi and Price 1992). Seeds

are dispersed over a 3-week period from late April to

mid-May.
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Chapter 23 Shrubs of Other Families

Ecological Relationships and Distribution—

Arroyo willow is distributed at low elevations from

British Columbia to Idaho and south from Baja California to west Texas and northern Mexico. It occurs

entirely east of the Cascade Mountains. Arroyo willow

may be found growing along streams, ditches, and

washes (Welsh and others 1987). It is adapted to welldrained, ephemerally wet riparian sites (Manning and

Padgett 1995). In Nevada, arroyo willow occurs on

streambenches with coarse-textured or dry surface

soils, along incised or ephemeral streams, and near

seeps (Manning and Padgett 1995).

Plant Culture—Arroyo willow may be propagated

from cuttings. Rooting ability ranges from erratic

(Platts and others 1987) to very good (USDA Soil

Conservation Service 1992). Roots form from the callus and over the lower one-third of the stem (Platts and

others 1987). Stem and root growth are initiated in

about 10 days.

Seed biology of arroyo willow has received little

study. Seeds are nondormant and remain viable in

nature for only 1 to 3 weeks (Sacchi and Price 1992).

Uses and Management—Arroyo willow has good

habitat value, high tolerance to flooding and sediment

deposition, moderate drought tolerance, and low salt

tolerance (USDA Soil Conservation Service 1992).

Management to preserve arroyo willow communities

in otherwise arid areas is critical; they provide important habitat for a wide range of invertebrate and

vertebrate species.

Native seedlings of arroyo willow establish on fresh

fluvial deposits and open or partially vegetated

streambanks (Sacchi and Price 1992). High seedling

mortality is associated with drying of the soil surface;

competition and herbivory play minor roles. Seedling

growth is reduced by shading, and small seedlings

have low overwinter survival. Seedlings surviving for

3 growing seasons may be considered established.

Varieties and Ecotypes—“Rogue” arroyo willow

was cooperatively released by the Corvallis USDASCS Plant Materials Center in 1990 for use in stabilizing streambanks of low velocity meandering streams,

improving freshwater fish and wildlife habitat, naturalized landscaping, and windbreaks or screens in

riparian or moist upland situations (Darris and Lambert 1993).

Rogue is a male clone derived from a population

growing along the Rogue River in Curry County, OR

(Darris and Lambert 1993). Plants are large, multistemmed or occasionally single trunked with spreading crowns. Rogue was selected for its rapid, early

growth rate and low incidence of insect and disease

pests. It is widely adapted to riparian areas and moist

upland sites receiving greater than 35 inches (890 mm)

of annual precipitation from Washington to northern

California on the west side of the Cascade Mountains.

The release is propagated from cuttings and can be

used for wattling, brush matting, and branch packing

in combination with mechanical site treatments (Darris

and Lambert 1993).

Family Salicaceae _______________

Salix lutea

Yellow willow, shining willow

Description—Yellow willow is a rounded shrub or

rarely a small, usually multi-stemmed tree 10 to 20 ft

(3 to 6 m) tall (fig. 51). Young twigs are sparsely hairy

and yellowish to reddish, becoming glabrous and yellowish white, gray, or brownish with age. Mature

leaves are lanceolate to elliptical, entire to serrulate,

0.8 to 2.2 inches (2 to 5.5 cm) long, and 0.4 to 0.8 inch

(10 to 21 mm) wide. They are dark green to yellow

green above and pale green and glaucous below. However, a nonglaucous form has been reported (Manning

Figure 51—A low- to mid-elevation species, yellow

willow frequently occurs as an early seral species on

periodically flooded streambars.
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Chapter 23 Shrubs of Other Families

and Padgett 1995). Catkins expand slightly before or

with the leaves. Staminate catkins are 0.8 to 2 inches

(2 to 5 cm) long. Pistillate catkins are 0.8 to 2.8 inches

(2 to 7 cm) long and densely flowered. The persistent

floral bracts are hairy and brown to blackish red.

Capsules are glabrous and 0.1 to 0.2 inch (3 to 6 mm)

long, produced on stipes less than 0.1 inch (2.5 mm)

long. Stigmas are often scarcely lobed (Brunsfeld and

Johnson 1985; Hitchcock and others 1964; Welsh and

others 1987).

Flowering and fruiting dates vary within and among

populations (Martens and Young 1992). Catkins appear in late March or early April on stands near Reno,

NV. Flowering has been reported to occur from May

to June in California (Munz 1973). Capsules ripen

about a month following flowering (Martens and

Young 1992). Catkins on a single branch vary in

phenological development, and flowering within each

catkin is indeterminate. Consequently, fruit ripening is not uniform. Seeds are dispersed by wind and

are buoyant in water. Hairs on the seed may aid in

dispersal, but they may also be important in germination and seedling establishment (Martens and

Young 1992).

Ecological Relationships and Distribution—

Yellow willow is distributed from Washington to

Manitoba and south from California to Nebraska

(Brunsfeld and Johnson 1985). A common species

within its range, it grows along streams and ditches in

valleys and canyon bottoms. It generally occurs at low

to mid elevations, but may be found at higher elevations, particularly in the southern portion of its range.

Associated upland communities include mountain big

sagebrush, pinyon-juniper, mountain mahogany,

Jeffrey pine, or lodgepole pine (Uchytil 1989c).

Yellow willow is capable of withstanding prolonged

inundation and often occurs as a pioneer or early seral

species on periodically flooded streambars. On such

sites it may occur with, or replace, whiplash willow or

coyote willow. Understory cover on such sites is often

sparse. Yellow willow stands also occur on wet alluvial

benches or terraces with well-developed, usually finetextured soils. Water tables are generally high, but

may fall below 3.3 ft (1 m) in summer. Understory

species include beaked sedge, Nebraska sedge, bluejoint

reedgrass, and other graminoids (Hansen and others

1995; Manning and Padgett 1995; Youngblood and

others 1985). Understory productivity and accessibility to livestock depend on the extent of willow canopy.

Yellow willow communities may occur in mosaics with

other riparian community types dominated by cottonwoods, Booth willow, or Drummond willow.

With excessive livestock grazing in yellow willow

communities, the water table falls, surface soils dry,

and streams may become incised (Hansen and others

1995; Manning and Padgett 1995). Associated species

may be shifted to shrubs such as Woods rose, mesic

forbs, or with prolonged excessive grazing, Kentucky

bluegrass and other weedy and exotic species.

Plant Culture—Yellow willow may be planted as

unrooted or rooted cuttings. Unrooted cuttings are

better able to establish on sites with a high water table

that persists throughout the summer and allows roots

to develop. Roots are initiated along the entire length

of the stem, but are most abundant along the lower

one-third. Under greenhouse conditions, new roots

and stems are initiated in about 10 days (Platts and

others 1987). Although cuttings root rapidly, only

moderate numbers of roots are produced.

Seed of yellow willow may be collected by harvesting the inflorescences as the capsules begin to open

(Martens and Young 1992). Developing fruits should

be monitored carefully because seed is dispersed

quickly when mature. Harvested catkins can be spread

to dry in a warm room until capsules open, releasing

the seed.

Fresh yellow willow seed harvested near Reno, NV,

exhibited greatest germination in the dark at 50 to

77 ∞F (10 to 25 ∞C) (Martens and Young 1992). Germination at 36 ∞F (2 ∞C) exceeded 10 percent; no seeds

germinated at 95 to 104 ∞F (35 to 40 ∞C). When

germinated in the dark at 55 alternating temperature

regimes, utilizing temperature combinations between

32 and 104 ∞F (0 and 40 ∞C) (8 hrs/16 hrs), greatest

germination (52 percent) occurred at 50/36 ∞F (10/2 ∞C).

Temperatures corresponding to very cold, cold, and

moderate seedbed conditions enhanced germination;

alterations corresponding to warm and warm/fluctuating seedbed conditions reduced germination.

Yellow willow seeds can be stored for short periods

of time (Martens and Young 1992). Seeds exhibited

some viability after 7 weeks when stored in water at

36 ∞F (2 ∞C), in paper bags at 32 ∞F (0 ∞C), or at room

temperature (Martens and Young 1992). Viability of

seeds stored in a desiccator over calcium chloride at

32 ∞F (0 ∞C)declined from 25 to 2 percent after 2 weeks.

Uses and Management—The USDA Soil Conservation Service (1992) described yellow willow as having good habitat value, good rooting ability, moderate

tolerance of flooding, moderate drought tolerance, and

low salt tolerance. Yellow willow often forms a thick

corridor that provides streambank stabilization. The

fibrous root system contributes to the development of

overhanging banks; the canopy provides shade and

litter, enhancing fish habitat.

A dense canopy cover of yellow willow provides food

and cover for big game, beaver, birds, and other wildlife. Moose and elk use yellow willow in summer and

winter. Excessive browsing, however, can result in

stand opening or loss of willows (Gaffney 1941; Hansen

and others 1995; Van Dersal 1938).
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Chapter 23 Shrubs of Other Families

Livestock may make heavy summer and fall use of

palatable understory species associated with some

yellow willow stands. Heavy livestock use of willows

may reduce their vigor, or if prolonged, result in stand

loss. Wet soils are easily damaged by livestock trampling early in the season, resulting in soil compaction,

early drying of the soil surface, and bank sloughing.

Decadent stands resulting from excessive browsing by

livestock or big game will regenerate if browsing

pressure is reduced. Fire may also be used to rejuvenate decadent stands (Hansen and others 1995). Recreation and travel within these communities require

careful consideration due to the dense willow canopy

and early season wet soils (Hansen and others 1995).

Campsites and trail or road routes should skirt these

areas when possible.

Varieties and Ecotypes—None.

Family Salicaceae _______________

Salix planifolia

Plainleaf willow

Description—Plainleaf willow is a shrub growing

to 13 ft (4 m) in height. Twigs are glabrous and black

to purplish black, exfoliating in translucent flakes.

Mature leaves are elliptical, entire, dark green, shiny

above, strongly glaucous beneath, and not permanently pubescent on both sides (fig. 52). Veins are

prominent and partially parallel. Catkins expand before or with the leaves. Staminate catkins are 0.4 to

1 inch (1 to 2.5 cm) long. Pistillate catkins are densely

flowered and 0.8 to 1.6 inches (2 to 4 cm) long. Floral

bracts are persistent, black, and long hairy. Capsules

are short hairy and 0.1 to 0.3 inch (3 to 7 mm) long.

Stigmas are lobed (Hitchcock and others 1964).

Three varieties are recognized. Salix planifolia var.

planifolia ranges from 6.6 to 13 ft (2 to 4 m) in height

with elongate leaves 1.4 to 3.1 inches (3.5 to 8 cm) long.

The leaves are usually red tinged and sparsely hairy.

The other two varieties are generally less than 6.6 ft

(2 m) in height. S. p. var. monica is usually 3.3 ft (1 m)

or less in height with leaves that are 1 to 1.4 inches (2.5

to 3.5 cm) long. S. p. var. pennata is 3.3 to 6.6 ft (1 to

2 m) tall with leaves 1.8 to 2.6 inches (4.5 to 6.5 cm)

long (Brunsfeld and Johnson 1985; Hitchcock and

others 1964).

Ecological Relationships and Distribution—

Plainleaf willow is circumboreal and extends southward in North America from California to New England

(Welsh and others 1987). It is found along streams,

around the margins of lakes and ponds, and in other

wet areas.

Salix planifolia var. planifolia occurs east of the

mountains in Western Canada and the Northwestern

United States at low to moderate elevations. In the

United States, it is distributed from the upper sagebrush to the Douglas-fir zone in northern Idaho, Montana, and Wyoming (Uchytil 1991c). It grows on mineral soils with textures ranging from gravelly to sandy

or clayey. Water tables on these sites may fall below

3.3 ft (1 m) by midsummer. Associated species often

include other tall willows such as Bebb willow, Booth

willow, or Geyer willow.

Salix planifolia var. monica occurs in valley bottoms

at high elevations and on wet, open, subalpine slopes

from central Idaho to west-central Montana, and south

from the Sierra Nevada Mountains of California to

New Mexico. It often occurs in upper cirque basins

with cold air drainages (Brunsfeld and Johnson 1985).

It usually grows on wetter sites than S. p. var. planifolia. S. p. var. monica is often associated with Wolf

willow, but grows on saturated soils with finer textures. Soils on S. p. var. monica sites usually have an

organic surface horizon overlying alluvial sands, silts,

clays, or sometimes gravels. Soils are often gleyed

near the upper horizon (Youngblood and others 1985).

Soil pH is generally slightly to strongly acidic (pH 4.4

to 6.3). Stands may be flooded in spring, and the water

table generally remains within the rooting zone

throughout the growing season. Lowering the water

table may lead to an increase in Wolf willow. S. p. var.

pennata is more narrowly distributed. It occurs in the

Cascade Mountains from northern Washington to northern Oregon.

Plant Culture—Rooted or unrooted plainleaf willow cuttings may be planted. Unrooted cuttings root

rapidly, but rooted cuttings are recommended for

high-elevation sites with short growing seasons and

rapidly drying soils. Stems produce low to moderate

numbers of roots along their entire length (Platts and

Figure 52—Plainleaf willow is a small- to mediumsized shrub with dark green elliptical leaves that are

shiny above and glaucous beneath.
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Chapter 23 Shrubs of Other Families

others 1987). Root formation is initiated in about 10

days; shoots require 10 to 15 days to develop under

greenhouse conditions.

Densmore and Zasada (1983) found that seeds collected in Alaska were nondormant at maturity. Germination was rapid and nearly complete at constant

temperatures ranging from 41 to 77 ∞F (5 to 25 ∞C).

Uses and Management—Roots of plainleaf willows growing in riparian areas contribute to the formation of overhanging banks. Plants also furnish

shade for fish habitat. Beaver use the plant for food

and building materials. Stem pieces not consumed

may sprout and develop into new plants (Cottrell

1995). Big game use plainleaf willow communities for

forage, cover, and as travel corridors (Hansen and

others 1995; Manning and Padgett 1995; Padgett and

others 1989). Moose make heavy winter use of S. p.

var. planifolia, but use by elk and mule deer is low

(Dorn 1970; Mattson 1984; Van Dersal 1938). Winter

use of S. p. var. monica may be limited due to its low

stature. However, in some areas, moose browse all

branches that are accessible in winter. Plainleaf willow communities also provide valuable cover and food

for small mammals and birds (Douglas and Ratti 1984;

Finch 1987).

Open stands of S. p. var. planifolia provide moderate amounts of forage for livestock (Hansen and others

1995) compared to other willow communities. Production of these communities is moderate. Palatability of

S. p. var. planifolia to cattle is low (Dorn 1970), but

palatability of some associated understory species is

high. Early season or season-long use by livestock

should be avoided as trampling of saturated sites will

damage plants and soils (Manning and Padgett 1995).

Short growing seasons and continually wet soils usually limit livestock use to late summer and early fall

(Hansen and others 1995). The grazing value of sites

dominated by S. p. var. monica is limited due to its

small stature, low productivity, and association with

wet soils.

Subalpine and alpine areas occupied by plainleaf willow provide an array of recreational opportunities including hiking, wildlife viewing, and fishing (Hansen

and others 1995). These communities, particularly the

wetter S. p. var. monica sites, are easily damaged by

hikers, livestock, horses, and offroad vehicles. Heavy use

can cause soil compaction, rutting, loss of vegetation,

formation of multiple trails or roads, streambank erosion, and other long-term damage that cannot be easily

repaired. Travel should be restricted to existing trails or

roads, and these carefully maintained. New travel routes

should be constructed on adjacent uplands.

Response of the species and its varieties to burning

and its ability to resprout have not been examined

(Hansen and others 1995). Short-term productivity of

understory sedges can be increased by burning.

Protection from grazing by livestock during the growing season leaves a thick cover of dry vegetation that

burns well in spring before new growth is initiated.

Burns along streambanks should be conducted with

caution to avoid loss of the bank stabilization provided

by these communities. Burned sites should be protected from grazing for 2 to 3 years to prevent overuse

of recovering plants.

Varieties and Ecotypes—None.

Family Salicaceae _______________

Salix scouleriana

Scouler willow, fire willow, black willow,

mountain willow, nuttall willow

Description—Scouler willow is a shrub or

multistemmed tree arising from a massive root crown

and a deep, spreading root system. Mature trees may

be 5 to 60 ft (1.5 to 18 m) tall. Young twigs are gray and

short hairy, becoming dark reddish brown and glabrous. The crushed bark has a distinctive skunklike

odor. Mature leaves are oblanceolate to lanceolate

or ovate, entire to crenate or serrate, 0.8 to 2.4

inches (2 to 6 cm) long, and 0.4 to 1.2 inches (1 to 3 cm)

wide. Leaf surfaces are dark green and glabrous above

and sparsely reddish hairy beneath. Leaves turn yellow in autumn. Catkins usually expand before the

leaves and are soon deciduous. They are nearly sessile.

Staminate catkins are yellowish white, 0.6 to 1.4

inches (1.5 to 3.5 cm) long, and about as long as wide.

Pistillate catkins are reddish, densely flowered, 0.8 to

2.4 inches (2 to 6 cm) long, and 0.5 to 0.7 inch (1.3 to 1.7

cm) wide. Floral bracts are brown to black and long

hairy. Capsules are hairy, 0.2 to 0.3 inch (5 to 8 mm)

long, and more or less long beaked. Stigmas are entire

to bilobed (Brunsfeld and Johnson 1985; Hitchcock

and others 1964).

Scouler willow is one of the earliest willows to flower.

Flowers appear from April to July (Orme and Leege

1980); fruits ripen and seeds disperse from May to July

(Brinkman 1974i). Seeds remain viable for only about

1 week (Densmore and Zasada 1983).

Ecological Relationships and Distribution—

Scouler willow is distributed from Alaska to Manitoba

and south to California, South Dakota, and New Mexico

(Brunsfeld and Johnson 1985). At low to mid elevations, it occurs in valley bottoms, along streams, and

around springs, where it attains a treelike stature.

Scouler willow, however, is generally an upland species, growing on well-drained slopes at higher elevations, where it occurs as a minor understory species in

late seral Douglas-fir, subalpine fir, western hemlock,

and western red cedar forests (Arno and Hammerly

1977; Froiland 1962). It is slightly tolerant of shade
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Chapter 23 Shrubs of Other Families

and capable of persisting on these sites at low frequencies (Cooper and others 1987; Steele and Geier-Hayes

1989, 1992). Scouler willow may be found on mineral

soils of varying textures and on peaty soils (fig. 53). It

is tolerant of acidic, but not salty, soil conditions.

Scouler willow proliferates following disturbance. It

often becomes widespread following stand-destroying

wildfires, slash burning, clearcutting, road building,

or other disturbances that expose mineral soil and

provide suitable sites for seed germination and establishment of seedlings (Arno and others 1985; Steele

and Geier-Hayes 1989, 1992, 1995). It is particularly

abundant where soils have been disturbed in such a

manner as to trap water. Some broadcast burn operations and wildfires that leave 50 percent or more of

the preburn overstory intact, however, do not provide

suitable seedbeds for Scouler willow, and an increase

in snowbrush ceanothus may be favored (Forsythe

1975; Steele and Geier-Hayes 1995).

High seedling densities on exposed, wet mineral

soils may result from heavy seed rain originating from

unburned areas (Lyon 1971; Stickney 1986). Scouler

willow seeds sown on burns of differing severities on

upland black spruce sites in Alaska germinated only

on moderately (organic soil layers partially consumed)

and severely (ash layer present, organic material in

soil consumed or nearly so to mineral soil) burned

seedbeds (Gruell and others 1982; Lyon and Stickney

1976). Germination was greatest and seedlings survived after 3 years only on severely burned sites.

Burned Scouler willow plants resprout vigorously

from the rootcrown; multiple sprouts sometimes appear a few days after a fire (Foote 1983). Some plants

regenerate even following burns that destroy the entire canopy (Lyon and Stickney 1976). Regenerating

trees grow rapidly, particularly during the first two

growing seasons (Leege and Hickey 1971; Lyon 1971).

Vegetative spread occasionally occurs through rooting

of branch segments (Watson and others 1980).

Dense stands of Scouler willow seedlings and regenerating plants may form shrub fields following fire.

These may persist for varying periods of time with

willow coverage decreasing as conifers mature (Steele

and Geier-Hayes 1995).

Plant Culture—Scouler willow has been used to

stabilize cut-and-fill areas along logging roads and

other disturbed upland areas, but cuttings do not root

as readily as those of many other willows (Monsen

1975; Plummer 1976). Roots develop in moderate

numbers from the callus area; they do not develop

along the length of the stem. Formation of roots normally requires about 10 to 15 days under greenhouse

conditions (Platts and others 1987). Densmore (1978)

found that total percent and rate of rooting were

greater for cuttings of second-year wood harvested in

the spring following leaf expansion (10 percent in 30

days, 1 root each) than for dormant cuttings harvested

in the fall (4 percent in 60 days, 4.5 roots each). Secondyear wood was collected from a population near

Fairbanks, AK.

In a field test, 8- to 10-inch (20- to 25- cm) dormant

cuttings of 2- to 3-year-old-wood harvested near

Fairbanks, AK, were stored fully imbibed at 14 ∞F (–10 ∞C)

until planted soon after soil thaw. Cuttings were

planted vertically with 0.8 to 2 inches (2 to 5 cm) of

stem above ground. Survival following one growing

season was 17 percent. Nearly 100 percent of all stem

cuttings collected in late fall and early winter from

sites in Idaho stored in a moist cold environment, and

spring planted in nursery beds, rooted and produced

healthy plants. Plants grown under nursery conditions for one season developed large, robust root systems. When outplanted, the rooted plants established

quickly and provided cover on road cuts and fill slopes.

Scouler willow may also be propagated from seed. There

are about 6,500,000 seeds per lb (14,300,000 per kg)

(Brinkman 1974i). Germination of seeds incubated at 85/

70 ∞F (29/21 ∞C) reached 95 percent in 1 day. Densmore

and Zasada (1983) obtained rapid and nearly complete

germination of an Alaskan collection when seeds were

incubated over a range of constant temperatures from 44

to 71 ∞F (5 to 25 ∞C).

Uses and Management—Scouler willow is a

highly valued browse species for big game. Seral

brush fields of Scouler willow and other shrubs are

important winter ranges (Leege 1968). Moose, elk,

white-tailed deer, and mule deer use the species

throughout the year, but primarily in winter (Kufeld

1973; Kufeld and others 1973; Singer 1979; Smith

1953). In south-central Alaska, moose sometimes

remove the bark (Viereck and Little 1972). Recent

sprouts are most palatable; greatest production may

Figure 53—Generally an upland species, Scouler

willow grows on dry slopes as an understory in

conifer forests.
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Chapter 23 Shrubs of Other Families

Figure 54—Wolf willow forms dense patches of lowstatured shrubs in narrow valley bottoms at high

elevations.

occur on branches browsed the previous year. Much

of the browse on mature plants is unavailable because plants are quite tall. Scouler willow also provides cover for birds and small mammals (Steele and

Geier-Hayes 1992; Vories and Sims 1977). In seral

brush fields Scouler willow provides excellent cover

for big game. Isolated plants in late-seral coniferous

stands are not as effective. Scouler willow is highly

palatable to cattle and sheep (Dayton 1931). It may

receive greater use than other willow species due to

the high accessibility of the willow clumps.

Prescribed burning in fall or spring has been used to

stimulate sprouting of Scouler willow shrub fields that

have grown beyond the reach of big game (Leege 1968,

1969, 1979a). Spring burning may be preferable; fall

burning destroys winter browse. Considerable growth

may occur during the first growing season following

spring burning. Leege (1979a) found that sprouts 5 ft

(1.5 m) long were produced following a spring burn in

Idaho. Burning at 5-year intervals did not decrease

productivity (Leege 1968). Plant nutritive value increases only slightly after burning, but new shoots are

within reach and more palatable than older shoots.

Scouler willow in clearcuts may enhance shadetolerant spruce and Douglas fir establishment by protecting plants from wind and temperature extremes

(Steele and Geier-Hayes 1989, 1992). In the absence of

a conifer overstory, Scouler willow’s growth habit is

altered from a narrow, upright form to a more shrublike

form with a broad, rounded canopy. These vigorous

plants compete strongly with shade-intolerant seedlings of ponderosa pine. Because the two species have

similar growth rates for the first 6 to 8 years, the pine

seedlings may not survive to overtop the willow. Mechanical treatments to remove Scouler willow are

generally impractical due to the plants’ deep, spreading root system. Consequently, plantations of Douglasfir are more likely to be successful.

Scouler willow has been recommended for use in

watershed plantings (Monsen 1975). Rooted cuttings

establish quickly when planted on road fills and other

disturbances. They grow quickly and provide excellent ground cover on exposed unstable slopes. They

also may be used in naturalized landscaping to provide

screens or background plantings.

Varieties and Ecotypes—None.

Family Salicaceae _______________

Salix wolfii

Wolf willow

Description—Wolf willow is a low-statured shrub

2 to 5 ft (0.6 to 1.5 m) tall with a fibrous, spreading root

system. Young twigs are yellow to orange and more or

less persistently thin hairy, becoming chestnut brown

when mature. Mature leaves are entire, elliptical to

lanceolate or oblanceolate, 0.5 to 1.6 inches (1.2 to

4.2 cm) long, and 0.2 to 0.5 inch (0.5 to 1.3 cm) wide.

They are gray green to silvery and sparsely to densely

hairy on both sides. Catkins expand with the leaves.

Staminate catkins are yellowish and 0.2 to 0.6 inch

(0.5 to 1.5 cm) long. Pistillate catkins are globose,

fuzzy, white, densely flowered, and 0.4 to 0.8 inch (1 to

2 cm) long. Floral bracts are persistent, long hairy, and

blackish or pale at the base. Capsules are 0.1 to 0.2

inch (3 to 5 mm) long and glabrous or rarely hairy.

Stigmas are lobed (Hitchcock and others 1964; Sutton

and Johnson 1974; Welsh and others 1987).

Ecological Relationships and Distribution—

Wolf willow is distributed from northeastern Oregon

south to Nevada and east to Montana and Colorado

(Brunsfeld and Johnson 1985). In central Idaho it

occurs from cooler sites at mid elevations in the Douglas-fir zone to subalpine areas, where it grows along

streams and around lakes and ponds. Wolf willow

community types in southeastern Idaho, northern

Utah, western Wyoming, and Montana occur in broad

meadows, on alluvial terraces, around seeps, and in

old beaver ponds at mid to upper elevations (Hansen

and others 1995; Padgett and others 1989; Youngblood

and others 1985). Wolf willow often dominates a low

shrub overstory, although scattered taller willows

may be present. At high elevations, Wolf willow may

occur in pure stands (fig. 54). Wolf willow occurs on

soils with a relatively thick organic surface horizon

that develops from deposition of leaves and twigs.

Some sites with lower amounts of organic matter have

fine mineral soil with high water-holding capacity.

Oxygen is provided by lateral movement of water

through such soils. The water table is generally within
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Chapter 23 Shrubs of Other Families

3.3 ft (1 m) of the surface throughout the growing

season. Stream channels in Wolf willow communities

are often small and meandering. Streambanks are

stable due to the high density of willow roots in the soil.

Overland flow is common during snowmelt.

Plant Culture—Wolf willow may be used to revegetate streambanks and ponds, although growth may

be slow. Rooted cuttings are advised for use on highelevation sites with short growing seasons. Hansen

and others (1995) recommended use of dormant 2- to

4-year-old wood. Cuttings should be 12 to 20 inches (30

to 50 cm) long and 0.4 inch (1 cm) or more in diameter.

Rooting is erratic; few to a moderate number of roots

normally form along the length of the stem (Platts and

others 1987). Ten to 15 days are required for root and

stem initiation. The seed biology of Wolf willow has not

been studied.

Uses and Management—Wolf willow communities should be managed to maintain willows along

stream margins and provide vegetative cover throughout the community. Wolf willow communities provide

valuable streambank stabilization (Hansen and others 1995). The low overstory and diversity of plant

species often present in these communities provide

cover for birds, small mammals, and other small

vertebrates and invertebrates. Cover value for big

game is fair. Biomass production of Wolf willow is

moderate. The species is palatable to sheep, beaver,

and big game. Winter use is minimal when plants are

leafless and snow covered. Understory graminoids

and forbs vary in palatability. Species such as water

sedge or tufted hairgrass may receive heavy use by

cattle, but accessibility may be restricted by the dense

shrub overstory. Excessive grazing may lead to

replacement of the understory by less palatable species. Grazing and equipment use should be restricted

until soils are dry as they are easily compacted and

become erodible if the vegetative cover is lost. The

response of Wolf willow to fire has not been documented (Hansen and others 1995).

Varieties and Ecotypes—None.

Family Saxifragaceae ____________

Philadelphus lewisii

Lewis mockorange

Description—Lewis mockorange (fig. 55), known

as syringa, Lewis syringa, mockorange, or Indian

arrowwood was named for Captain Meriwether Lewis,

who first collected it on the Clark Fork River near

Missoula, MT, on July 4, 1806. Plants are long-lived

perennials, ranging from open to densely branched,

erect to rounded, fibrous rooted shrubs 3 to 12 ft (0.9

to 3.7 m) in height (Carson and Peek 1987; Dittberner

and Olsen 1983). Clusters of arching stems develop on

older specimens. Young branches are divaricate and

glabrous to pubescent with red to chestnut brown

bark. Conspicuous transverse cracks develop in the

bark during the second season; the bark later turns

gray and eventually exfoliates. The light green leaves

are opposite, sessile, or short petiolate and ovate to

oblong with acute to acuminate tips. They are 1 to 3

inches (2.5 to 7.6 cm) long and 0.4 to 1.6 inches (1.0 to

4.1 cm) wide. The thin blades are entire, 3 to 5 veined,

usually glabrous above, but pubescent around the

edges and along veins beneath.

Attractive terminal cymes of attractive white flowers develop on lateral branches, each with 3 to 11

perfect, regular flowers. Flowers arise from the axils of

gradually reduced leaves. There are four white petals,

20 to 60 unequal stamens with conspicuous yellow

anthers, and three to five persistent styles that are

more or less connate. Fruit is a woody capsule with

four cells attached to the persistent calyx. The light

brown fusiform seeds are numerous (Hitchcock and

others 1961; Welsh and others 1987). Flowering occurs from late May to July. Fruits develop from July to

September or October when the capsules dehisce and

seeds are wind and gravity dispersed (Orme and Leege

1980; Young and Young 1986).

Lewis mockorange exhibits a high degree of local

variation in floral and vegetative characteristics. This

variability has been assigned taxonomic status by

some workers (Hitchcock and others 1961).

Ecological Relationships and Distribution—

There are about 50 species of Philadelphus in North

America, Mexico, Asia, and central Europe. Only two

species, Lewis mockorange and littleleaf mockorange

(Philadelphus microphyllus), occur in the Intermountain region. Lewis mockorange is distributed from

Figure 55—Lewis mockorange survives on a low

foothill site following many years of excessive grazing.
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Chapter 23 Shrubs of Other Families

British Columbia south to northern California and

east to Montana (Hitchcock and others 1961;

Lackschewitz 1986). It grows from near sea level to

7,000 ft (2,100 m), but is most common at midelevations. It is a dominant shrub in some climax ponderosa

pine communities and a common shrub in some seral

ponderosa pine communities (Wright 1978). It is also

found in sagebrush deserts, lodgepole pine forests,

and coastal Douglas-fir and redwood forests. It commonly occurs in foothills and low mountains on all

aspects, but primarily on northern and eastern slopes

(USDA Forest Service 1937). The species grows on

soils ranging from dry, rocky, gravelly loams on open

hillsides, to deep, rich alluvial humic loams near

riparian zones (USDA Forest Service 1937). It generally occurs in small scattered clumps in moist, open, or

partially shaded transitional areas along gullies, canyon bottoms, riparian areas, and seeps (Carson and

Peek 1987; Hopkins and Kovalchik 1983). It also

grows on cliffs, talus slopes, and rocky hillsides with

other shrubs such as serviceberry and ocean-spray

(Carson and Peek 1987; Franklin and Dyrness 1973).

Plant Culture—Seeds may be collected by hand

stripping the capsules from the plants after maturity,

but before the valves begin to open. Dried capsules are

crushed to release the seeds, and trash is removed

with an aspirator or fanning mill. There are 3,500,000

to 8,000,000 seeds per lb (7,716,000 to 17,637,000 per

kg) of clean seed (Stickney 1974b). Seed fill is often

low. Seeds may be stored in airtight containers for

periods of up to 1 year. They require an 8-week wet

prechilling at 41 ∞F (5.0 ∞C) to release embryo dormancy (Stickney 1974b). Germination is tested at 72 to

79 ∞F (22 to 26 ∞C).

Seeds may be broadcast seeded on a rough seedbed

and covered lightly or spot seeded in selected, prepared areas. They may also be surface seeded using a

Brillion seeder or similar device. Best results are

obtained if seeds are planted in well-drained sites free

of herbaceous competition. Seeds may be mixed with

other shrub seeds that require shallow or surface

planting.

Bareroot stock may be produced by fall seeding or by

seeding wet prechilled seeds in spring. The tiny seeds

may be diluted by mixing them with rice hulls to

improve uniformity of seeding. Seeds should be covered very lightly. Seedlings develop rapidly and can be

transplanted as 1-year-old stock. Container stock may

also be grown from seeds.

Due to a lack of commercial seed sources and ease of

vegetative propagation, planting stock is frequently

grown from cuttings. Dormant 3 to 4 inch (7.6 to

10.2 cm) hardwood cuttings may be planted in a closed

cold frame with bottom heat for rooting. Softwood

cuttings gathered in early summer or early fall are

also easily rooted in a cold frame (Doran 1957; Marchant

and Sherlock 1984). Digging rooted suckers and dividing the crown of mature plants with a sharp spade or

axe are alternate means of vegetative propagation.

Lewis mockorange is a valuable plant for transplanting on disturbed steep, rocky, unstable slopes

where it provides soil stabilization and vegetative

cover. Seedlings or larger stock are recommended for

such sites. Lewis mockorange is also useful for planting in transitional areas of degraded riparian zones.

Uses and Management—Lewis mockorange is not

normally grazed heavily by livestock, but in some

areas it does receive fair amounts of use by cattle and

sheep (Leege 1968; USDA Forest Service 1937). It

frequently occurs with other species that are more

palatable to big game and, consequently, it may receive little use, except under severe conditions. However, in some areas it does provide good browse for deer

and elk, especially on winter ranges (Kufeld 1973;

Leege 1968; Marchant and Sherlock 1984;

Stubbendieck and others 1986). New growth is generally highly palatable to big game (Leege 1968). Leege

(1969) reported that elk use of Lewis mockorange

increased by 30 percent following a spring burn. The

plants provide cover for birds and other small animals.

Lewis mockorange is classified as a fire resistant or

survivor species (Fischer and Bradley 1987). Although

shoots may be consumed by fire, Lewis mockorange is

capable of resprouting from the root crown or caudex

during the first season following burning (Fischer and

Bradley 1987; Rowe 1983). Consequently, it is found in

early to late seral as well as in climax communities.

Lewis mockorange is not as prolific a sprouter as cooccurring shrubs (Leege 1969). Leege and Hickey

(1971) found that spring-burned Lewis mockorange in

northern Idaho seral brush fields (grand fir/pachistima

habitat type) resprouted within 1 or 2 months, while

fall-burned shrubs did not resprout until the following

spring. Sprouts were more numerous following fall

burning. The historic burn interval in climax ponderosa pine communities is thought to be 6 to 22 years;

thus, Wright (1978) prescribed burns at 10 to 15 year

intervals to enhance shrub growth.

Lewis mockorange is the state flower of Idaho and is

one of the more extensively used native shrubs for

landscaping (Marchant and Sherlock 1984). It is valued

for its showy white flowers; the fruits and leaves are less

attractive. Dense branches and foliage make it suitable

for hedges. Native Americans used the long woody

shoots for arrow shafts (USDA Forest Service 1937).

Lewis mockorange seedlings establish well, but

should be protected from competition and browsing.

Landscape plants should be pruned after flowering

because flowers are produced on twigs of the previous

year. The species is normally free of insect and disease

problems, but seedlings are sometimes susceptible to

damping off.
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Chapter 23 Shrubs of Other Families

Varieties and Ecotypes—The Colfax germplasm

of Lewis mockorange originated from the Colfax, WA,

area. St. Maries germplasm originated from St. Maries,

ID. Several commercial cultivars of Lewis mockorange

have been developed. “Waterton,” selected from the

Waterton Lakes area of Alberta, is a hardy, bushy

shrub with flowers scattered over the crown of the

plant (Marchant and Sherlock 1984). Philadelphus

coronarius, introduced from Europe, is the commonly

cultivated mockorange (Welsh 1987).

Littleleaf mockorange is a smaller, short-lived

shrub, growing from 3 to 7 ft (0.9 to 2.1 m) in height and

4 ft (1.2 m) in width. Shrubs have ascending stems and

a rounded crown with reddish-tan exfoliating bark.

Leaves are narrower than those of Lewis mockorange,

and flowers are solitary or in clusters of three. Littleleaf

mockorange occurs from Utah to Wyoming and south

to Texas in pinyon-juniper, mountain brush, aspen,

lodgepole pine, and Douglas-fir/white fir forests

(Goodrich 1985; Sutton and Johnson 1974; Welsh and

others 1987). Littleleaf mockorange receives some use

by mule deer (Patton and Ertl 1982) and other wildlife.

Littleleaf mockorange may be propagated from seed or

cuttings. It is easily transplanted and is a strikingly

attractive ornamental (Sutton and Johnson 1974).

Family Saxifragaceae ____________

Ribes aureum

Golden currant

Description—There are more than 100 species of

Ribes, primarily occurring in the temperate and colder

regions of the northern hemisphere and the Andes

mountains of South America. They are particularly

common in the Western United States (Pfister 1974;

Welsh and others 1987). Species with jointed pedicels,

several flowers per raceme, and glabrous to glandular

berries that are generally unarmed are grouped as

currants. The remaining species, usually armed with

spines, are referred to as gooseberries (Hitchcock and

others 1961).

Golden currant is an unarmed, irregularly shaped,

multiple-stemmed shrub 3 to 10 ft (0.9 to 3.0 m) tall

(Welsh and others 1987) that spreads by suckering

(fig. 56). The reddish bark of young twigs turns gray

with age. Leaves are alternate, deciduous, and petiolate. Blades are ovate with a cuneate to chordate

base, usually glabrous, and palmately three-lobed

with the lobes entire, toothed, or lobed. Ascending

bracteate racemes occur on the ends of lateral spur

branches with two to 15 flowers (Harrington 1964).

Flowers are complete and regular with a spicy odor

(Stark 1966). The hypanthum is cylindrical with five

spreading calyx lobes alternating with five shorter,

erect petal lobes. Young flowers are golden yellow, but

turn reddish with age. The edible fruit is a globose,

glabrous, many-seeded berry. Fruit color varies from

yellow to red orange or black. The minute embryo is

embedded in a large amount of endosperm (Goodrich

and Neese 1986; Hitchcock and others 1961; Munz and

Keck 1959; Pfister 1974; Welsh and others 1987).

Ecological Relationships and Distribution—

Golden currant is distributed from north-central

Washington to Saskatchewan and South Dakota and

south to California and New Mexico (Welsh and others

1987). Hitchcock and Cronquist (1973) reported that

the shrub occurs along streambanks and washes, and

from grasslands and big sagebrush deserts to ponderosa pine forests throughout the Pacific Northwest. In

California it grows along moist streambanks and in

bottom lands at elevations between 2,500 and 7,800 ft

(760 and 2,400 m) (Munz and Keck 1959). Golden

currant occurs in many Utah riparian and palustrine

habitats in greasewood-shadscale, sagebrush, pinyonjuniper, mountain brush, ponderosa pine, and Douglasfir communities at 4,395 to 8,490 ft (1,340 to 2,600 m)

(Goodrich and Neese 1986; Welsh and others 1987). It

occupies similar areas in Colorado and New Mexico,

and is common in the plains and foothills of South

Dakota and Saskatchewan (Harrington 1964).

Golden currant is generally not widely abundant

(Wasser 1982); it occurs as scattered plants, patches,

clumps, and in corridors along waterways. It does not

occur as even a minor component of aspen or conifer

forests in the Intermountain area (Mueggler 1988;

Steele and Geier-Hayes 1987; Youngblood and Mauk

1985), but it does grow in aspen/chokecherry communities. It normally grows on fertile, well-drained sites

such as moist streambanks, washes, ditches, seeps,

and springs. It has been widely cultivated as an

Figure 56—Golden currant planted as a lowmaintenance species in a recreation site.
690 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

ornamental (Goodrich 1985; Pfister 1974), and now

occupies drier sites within abandoned farmlands,

ranches, and wildland areas.

Although considered a species for moist sites

(Thornburg 1982), golden currant is found on welldrained soils receiving in excess of 16 inches (40 cm) of

annual precipitation. It usually occurs on sandy to

silty loam soils (Thornburg 1982), but is also adapted

to clay-loam soils. Plants generally grow in full sun to

partial shade on soils ranging from slightly acidic to

slightly basic (Dittberner and Olsen 1983; Wasser

1982).

Plant Culture—Golden currant flowers appear

with or slightly before the leaves in April or May.

Fruits ripen in June or July and are dispersed by birds

and mammals (USDA Forest Service 1948). In central

Utah berries may ripen as late as mid-August (Plummer

and others 1968).

Fruits or berries are hand harvested by stripping

them from branches or by flailing them into containers

as soon as they ripen; this reduces losses to birds

(Pfister 1974). Few wildland stands are large enough

to produce many pounds of seed; consequently, cultivated ecotypes are usually the sources sold for wildland plantings. Native stands are capable of producing

good seed crops, but only about 4 lb (1.8 kg) of clean

seed are extracted from 100 lb (45.4 kg) of fruits

(Pfister 1974). Seed collection costs are high, and tend

to restrict use of the shrub in large-scale seeding

projects. However, viable seed crops are usually produced each year at most locations, and local ecotypes

can be collected and propagated for specific plantings.

Most seeds are harvested from cultivated plantings,

fence-line plantings, or irrigated seed fields. Plants in

these situations generally produce more consistent

and abundant crops. In addition, the bushes are more

uniform in stature, and are closely spaced, which aids

in harvesting.

Collected fruits should not be allowed to overheat

prior to extraction. Fruits are processed by maceration

using a Dybvig cleaner and water to separate the berry

or pulp from the seeds (Plummer and others 1968).

Dried fruit should be soaked in water to aid cleaning

or maceration. Pulp and empty seeds may be separated from sound seeds by flotation. There are 200,000

to 285,000 seeds per lb (441,000 to 628,000 per kg)

(Pfister 1974).

Heit (1971) recommended that tetrazolium chloride

staining be used for testing viability, but embryo

excision is difficult. Seed viability is generally quite

high, usually in excess of 75 percent. Seeds retain good

viability for 5 to 17 years if stored dry in sealed

containers (Pfister 1974) or in open warehouses

(Plummer and others 1968; Stevens and others 1981a).

Seed may remain dormant in the soil for many years

(Moss and Wellner 1953; Quick 1954).

The degree of dormancy varies among seedlots and

among seeds within a seedlot; this provides an adaptive advantage. However, this characteristic reduces

the effectiveness of seed pretreatments. Most seeds

require a long period of wet prechilling to break embryo dormancy. Germination can be hastened and

increased by wet prechilling at low temperatures. A

wet prechilling of 60 to 90 days at 28 to 36 ∞F (–2.2 to

2.2 ∞C) is often used to release embryo dormancy

(Pfister 1974). Stidham and others (1982) recommended that seeds be prechilled on blotters moistened

with potassium nitrate.

Seeds should be fall planted to provide a cold, moist

period for wet prechilling. Spring plantings should use

only prechilled seeds. Seeds are small, round, and

easily dispensed through most seeders. Golden currant can be seeded alone or in mixtures with selected

shrubs and forbs. It should not be seeded directly with

grasses or broadleaf herbs that germinate and grow

rapidly. Seeds are usually sown at a rate of 0.25 to

2 lb per acre (0.3 to 2.2 kg per ha), depending on the

method of seeding and row spacings. Wildland

seedings are usually conducted using seed-dribblers

mounted on tractors. Seeds are often hand planted

in selected areas. Only a small portion of an area is

actually planted using these techniques.

Under nursery conditions, seeds are usually sown at

a rate of 60 to 80 seeds per ft2 (650 to 860 per m2

) or

40 viable seeds per linear ft (130 per linear m) of

row (Pfister 1974). Seeds should be planted 0.25 inch

(6 mm) deep on a firm seedbed. Planting on sites

having some surface mulch or debris aids in germination and seedling establishment. Adding mulch to the

soil surface is recommended for nursery beds subject

to rapid drying and crusting.

Seed germination is usually only moderate or fair.

However, seeds that germinate often do so uniformly,

and initial emergence is usually very good even under

range or wildland conditions. Compared with most

shrubs, seedlings of golden currant are very persistent. Seedlings grow rapidly and generally attain

heights in excess of 6 to 12 inches (15 to 30 cm) the first

growing season. Although natural thinning occurs,

seedlings are vigorous; normally enough survive to

provide a full stand. Seedlings of few other shrubs are

as vigorous as this species.

Planting sites should be cleared of competition to

improve shrub seedling success. Golden currant seedlings and young plants are able to compete and establish with some competitive understory. The shrub is

more sensitive to competition when planted along

streams or in moist situations, yet seedlings of this

species are able to compete well with aggressive herbs

that exist in riparian communities. New seedlings

appear from natural seeding throughout native ranges,

which attests to the survival attributes of this species.
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Chapter 23 Shrubs of Other Families

Bareroot or container-grown transplants can be

quickly grown, although overseeding is usually recommended to assure emergence of the desired number of

seedlings. Planting beds are normally thinned to the

proper spacing and density. One-year-old transplants

develop a dense, well-branched root system, and field

survival of 1-0 transplants is usually very high. Transplants establish very well on harsh disturbances due,

in part, to the well-developed root system of young

plants. Container stock is grown from seed or hardwood or softwood cuttings (Doran 1957).

Uses and Management—Golden currant has been

widely planted for wildlife habitat (Patton and Ertl

1982), ground cover, watershed protection, and conservation plantings (Pfister 1974). It establishes with

good success in most wildland plantings; consequently,

it is often used to assure the establishment of a desirable species.

Golden currant is a highly preferred spring and

midsummer browse for big game, and is recommended

for planting in mountain brush, pinyon-juniper, big

sagebrush, and wet meadow communities that receive

more than 20 inches (51 cm) of annual precipitation

(Plummer and others 1968; Wasser 1982). Kufeld and

others (1973) reported moderate summer use, but only

light fall use by big game animals. The plant is eagerly

grazed in early spring and summer, and can be planted

on riparian sites and upland ranges to provide seasonal grazing by birds and big game.

Golden currant is commonly planted with a combination of other species to furnish diversity, early

developing forage, fruits, and habitat for big game and

birds. It provides preferred roosting and nesting cover

for several songbirds (Johnson and Anderson 1980).

Chukar partridge feed on the ripe fruits (Gullion

1964). Dayton (1931) reported that this shrub species

has about average palatability for domestic livestock.

Although Dittberner and Olsen (1983) reported considerable variation in palatability among golden currant populations in the West, this species produces an

excellent amount of herbage annually.

Golden currant is an excellent species for stabilization of roadways (Wasser 1982) and other disturbances, particularly if transplant stock is used. Both

bareroot and container-grown transplants establish

and survive well on most disturbances. Transplants

grow rapidly and furnish excellent ground cover in 2 to

3 years. If planted at close spacings, 3 to 4 ft (0.9 to

1.2 m) apart, an effective cover can develop quickly.

Plants are reasonably adapted to disturbances where

portions of the topsoil remain or where the topsoil has

been mixed with the planting media. Disturbed areas

that lack good fertility and consist of coarse fragments

are less likely to sustain this shrub. However, golden

currant is quite drought tolerant and can persist on

harsh mine sites. Ferguson and Frischknecht (1985)

recommended use of this species to reclaim coal fields

in south-central Utah. The species prefers fertile,

well-drained, neutral, loamy soils (Dittberner and

Olsen 1983; Wasser 1982). Haeussler and Coates

(1986) reported the plant grows best on soils supplied

with humus.

Golden currant is widely used for hedges, windbreaks, and conservation and landscape plantings

(Cook 1981; Wasser 1982). It is usually planted in

combination with other woody species, particularly

evergreens or less open deciduous species. It is often

planted as a conservation or wildlife habitat species,

but other shrubs provide more acceptable forage for

birds (Miller and others 1948). This species is commonly planted in rows or hedges to provide berries for

jam and jelly.

Golden currant is widely distributed along streambanks, floodplains, and drainageways (Hitchcock and

others 1961; Wasser 1982). It can be used to reclaim

riparian disturbances, particularly moist sites where

the water table has been lowered by erosion and

downcutting of gullies.

Golden currant is moderately fire tolerant (Hopkins

and Kovalchik 1983; Rowe 1983), and recovers by

resprouting or natural seeding (Rowe 1983). Pfister

(1974) reported that golden currant seedlings establish well on mineral soils where the organic duff is

removed by fire. Currants are subject to defoliation by

the western tent caterpillar (Furniss and Barr 1975);

they are also an alternative host for white pine blister

rust (Quick 1954).

Varieties and Ecotypes—None.

Family Saxifragaceae ____________

Ribes cereum

Wax currant

Description—Wax currant (fig. 57) is a spreading,

unarmed, multicrowned, deciduous shrub, usually 2 to 5

ft (0.6 to 1.5 m) tall with puberulent and more or less

stipitate glandular young branches (Hitchcock and others 1961; Welsh and others 1987). Leaves are fragrant,

clustered on short spurlike branches (Sampson and

Jespersen 1963), usually shallowly three- to five-lobed.

The upper surface of the leaves is subglabrous to

shiny, and the lower surface is waxy glandular dotted

(Stubbendieck and others 1986). The inflorescence is

pubescent and sticky glandular. Two or three flowers

occur in short drooping clusters. Flowers are greenish

white to pink (Welsh and others 1987). Fruits are dull to

bright red, glandular to glabrous, and usually between

0.2 to 0.3 inch (6 to 8 mm) long.

Ecological Relationships and Distribution—

Wax currant is found from British Columbia to southern California on the east slopes of the Cascade and
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Chapter 23 Shrubs of Other Families

Sierra Mountains, and eastward to Montana, Nebraska, and New Mexico (Hitchcock and others 1961).

It is one of the most widely distributed of all western

currants, because it occurs from sagebrush deserts to

alpine areas (Great Plains Flora Association 1986;

USDA Forest Service 1937). Ribes cereum var. cereum

is found from British Columbia to southern Arizona

and southern California, and east to central Montana,

Idaho, and western Nevada. The R. c. var. inebrians is

more common eastward from central Idaho and Montana, south and east to Nebraska, New Mexico, Utah,

and eastern Nevada. Welsh and others (1987) reported that most plants in Utah are R. c.var. inebrians,

but intergradation of morphological features is common, and no geographical correlation is apparent in

Utah. R. c. var. colubrinum is confined to the Snake

River Canyon and related tributaries in Idaho, Washington, and Oregon (Hitchcock and others 1961).

Goodrich and Neese (1986) reported the plant is

common and widespread in mountainous communities between 6,500 and 11,000 ft (2,000 to 3,400 m) in

Utah. Sampson and Jespersen (1963) reported that

wax currant is more widely distributed in California

than sticky currant, although their distributions overlap. Wax currant is common in various plant communities, but not as a dominant species. It is often

associated with antelope bitterbrush in the Central

Rocky Mountains (USDA Forest Service 1937). It is

prevalent in various ponderosa pine/Douglas-fir forest

types in Colorado and several lodgepole pine associations in Oregon (Volland 1985b). It occurs in aspen

communities (Mueggler 1988) and conifer forests of

Utah (Youngblood and Mauk 1985) and Idaho (Steele

and Geier-Hayes 1987), but only as a minor species.

Plant Culture—Wax currant plants flower from

April to June; fruits ripen in August (Pfister 1974).

Fruits are harvested by hand picking or flailing the

bush to dislodge the berries. Fruits are macerated to

remove the seed; the material is then dried, and seeds

are separated from the dry pulp using a fanning mill.

Seeds are quite small (Vories 1981), averaging about

251,000 per lb (553,400 per kg) (Pfister 1974). Seed

production from wax currant is generally lower than

from golden currant and sticky currant. In addition,

seed germination is usually low and a longer period of

wet prechilling is required to overcome dormancy. Pfister

(1974) recommended that seeds be wet prechilled for

120 to 150 days at 28 to 32 ∞F (–2.2 to 0.0 ∞C).

Seeds should be fall sown at a depth of 0.2 to 0.5

inch (6 to 13 mm) on a firm seedbed. Haeussler and

Coates (1986) recommended seeding on moist mineral soils with high percentages of organic matter.

Plummer and others (1968) reported that seeds of this

species have low seed germination, and the initial

establishment of new seedlings is only fair. Young

plants grow slower than golden currant seedlings, but

persistence of established plants is excellent. Plants

spread well from natural seeding. It is less successfully established by transplanting than golden currant. Wax currant is easily cultured in nursery beds.

Seeds of wax currant may be spread by birds or small

mammals (Kramer 1984) or by being deposited directly below the shrub (Moss and Wellner 1953). Seeds

may remain viable for a long period (Lyon and Stickney

1976). Pfister (1974) reported seeds retained good

viability after 27 years of storage.

Although seeds are relatively thin walled and can

be destroyed by severe fires (Kramer 1984), they

benefit from fire scarification (Morgan and

Neuenschwander 1985; Young 1983). New seedlings

often appear following low-severity fires that remove

little surface litter (Rowe 1983). Plants have large

and deeply buried root crows that resprout from

belowground tissue if not destroyed by fire (Bock and

Bock 1984).

Uses and Management—Wax currant is somewhat less palatable throughout the entire year than

golden currant (Plummer and others 1968). Kufeld

and others (1973) reported that big game made only

light use of the shrub in the spring and fall, but

Plummer and others (1968) reported it was heavily

browsed during all seasons.

Palatability of wax currant is low for livestock, but

it is an important browse due to its abundance, productivity, and availability (Dittberner and Olsen 1983;

Mueggler and Stewart 1980; Sampson and Jesperson

1963). This shrub has been principally used for wildlife habitat improvement, although limited plantings

have been established for watershed and groundcover protection. It is an important shrub for use in

restoration of mountain brush, aspen, and subalpine

communities (Plummer and others 1968).

Figure 57—Wax currant growing on a rocky outcrop above timberline.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 693

Chapter 23 Shrubs of Other Families

Wax currant has not been widely used for control of

severe erosion problems or disturbed land plantings.

It has been used to stabilize watersheds, small gullies,

and moist streambanks in aspen and conifer forests if

topsoil is present. Plants are adapted to disturbances

related to fire and logging. Natural regeneration

frequently occurs after these disturbances, and this

species provides considerable ground cover and herbage. It can be seeded and transplanted on areas exposed by logging or fires with acceptable success. The

shrub grows well with understory herbs, and can be

planted with a number of species to control erosion and

protect unstable sites. It does not spread rapidly to

occupy harsh disturbances, and has not performed well

as a pioneer species on mines, roadways, or similar

disturbances. This shrub grows well with some overstory trees, and can be used to plant campgrounds and

recreation sites where a dense tree cover may be desired. This species could be more widely used to restore

native communities.

Varieties And Ecotypes—None.

Family Saxifragaceae ____________

Ribes viscosissimum

Sticky currant

Description—Sticky currant (fig. 58) is a spreading, unarmed, uneven, aromatic shrub, mostly 3.3 to

6.6 ft (1.0 to 2.0 m) tall. Stems, leaves, inflorescences,

and fruits are sparsely to thickly glandular (Welsh

and others 1987). Leaf blades are orbicular, three- to

seven-lobed with the main lobes crenate or dentate,

round in outline, heart shaped at base, and hairy or

glandular on both surfaces (Sampson and Jespersen

1963). Flowers are green, white, or pink, with three to

12 flowers occuring in a cluster. Berries are 0.4 to

0.5 inch (10 to 13 mm) long, black, rather dry, and

glandular (Welsh and others 1987).

Ecological Relationships and Distribution—

This species is common from British Columbia to

California on the east side of the Cascade and Sierra

Nevada Mountains, and extending east to Montana

and Arizona (Hitchcock and others 1961; Sampson

and Jespersen 1963; Welsh and others 1987). Hitchcock

and others (1961) divided it into two intergrading

geographic races. R. v. var. hallii is distinguished by

its glabrous ovaries and glaucous berries; it occurs

throughout the eastern Cascades south of Mt. Rainier

and the Sierras. R. v. var. viscosissimum has glandular and more or less pubescent ovaries and nonglaucous

berries; it occurs over the remainder of the species’

range. Welsh and others (1987) concluded that plants

occurring in Utah are unique; the herbage has dense

stipitate-glandular and nonglandular hairs and long,

broad hypanthia.

Sticky currant grows in shady woods and rocky

places in the Sierra Nevada Mountains at elevations

between 6,000 and 9,000 ft (1,800 and 2,700 m)

(Sampson and Jespersen 1963). Goodrich and Neese

(1986) reported the species is common throughout the

Uinta Basin in Utah, existing with aspen and conifer

woods at 7,500 to 10,000 ft (2,300 to 3,000 m). Welsh

and others (1987) reported that this shrub occurs

throughout most of north, central, and eastern Utah,

growing in the shade of aspen, fir, Douglas-fir, lodgepole pine, and spruce woodlands, but is less common in

mountain brush and open meadows. Steele and GeierHayes (1987) reported sticky currant as a principal

species with grand fir in central Idaho where it is often

an early seral shrub, and may be the first shrub to

dominate scarified sites following logging. These authors concluded that plants have a low tolerance for

shade and begin to decline as a canopy taller than their

own develops. Sticky currant plants attain mature

stature within 5 years following logging. Their canopy

is sparse, and the shrub does not seriously compete

with the establishment of tree seedlings (Steele and

Geier-Hayes 1987). However, plants of sticky currant

remain longer than some species of ceanothus as

overstory shade develops. Sticky currant is not a

principal species in any conifer forest (Youngblood and

Mauk 1985) or aspen type (Mueggler 1988) in Utah.

Plant Culture—Sticky currants flower in May or

June, and fruits ripen in August or September (Pfister

1974). Fruits are harvested by hand picking or beating

the bush to dislodge the berries onto canvas or other

collection tarps. Fruits are cleaned by maceration

using a Dybvig separator. Filled seeds will sink or

settle in water, and the pulp and empty seeds can be

Figure 58—Sticky current generally occurs above decanted (Pfister 1974). Following maceration, large timberline on windswept ridges.
694 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

seedlots can also be dried and the seeds separated

from the debris using fanning mills (Plummer and

others 1968). There are 255,000 to 349,000 cleaned

seeds in 1 lb of cleaned seed (562,000 to 769,000 per kg)

(Pfister 1974).

Most plants produce some fruits each year. New or

young stands produce the most consistent crops.

Although fruits are large, seed collection is slow and

costly. Seed costs and poor availability discourage the

use of this species in large projects. Enough seeds can

be economically harvested most years from specific

locations to supply seeds for rearing transplant stock.

Seeds can be stored for extended periods, exceeding 17

years (Pfister 1974), if kept in dry sealed containers.

Steele and Geier-Hayes (1987) reported seed buried in

soil and duff remain viable long after the parent

shrubs disappear.

Seeds require a wet prechilling period of 140 days at

28 to 32 ∞F (–2.2 to 0 ∞C) to initiate germination (Pfister

1974). Seed germination is only moderate. Seeds gathered from different collection sites can exhibit variable

germination patterns. Plummer and others (1968)

reported that initial establishment was only fair

because of poor germination and relatively slow growth

of small seedlings. Growth of young plants is somewhat slow, but improves as plants attain mature

stature. However, plants that reach 2 to 3 years of age

may succumb under adverse conditions; final establishment of seeded plots is only moderate or good.

Plants are not rhizomatous, and natural spread is

slow unless sites are cleared by logging, fires, or

related disturbances.

This species can be produced as bareroot nursery

stock or container material. Initial establishment is

poor even under controlled rearing conditions; replanting is often required to produce full stands. Once

established, nursery stock grows well, and when transplanted to range or wildland sites, survives quite well.

Uses and Management—Sticky currant is recommended for planting aspen, Douglas-fir, spruce woods,

and less commonly mountain brush communities

(Plummer and others 1968). It has been used to enhance wildlife habitat (Pfister 1974; Plummer and

others 1968). Plants have only fair or poor palatability,

but are quite tolerant of grazing. In Utah the plant is

most commonly used during spring and summer periods by big game (Plummer and others 1968); heavy

browsing may occur during these periods. Sampson

and Jespersen (1963) reported that plants in California produce abundant herbage, which is heavily

cropped by sheep and deer, particularly in fall.

Sticky currant grows well with other species, and

frequently occurs with many other shrubs and herbs.

It can be planted to stabilize disturbances following

fire and logging. It grows well on undisturbed soils, but

is less vigorous and poorly adapted on sites where the

topsoil is disrupted or removed. The species is more

site specific than other shrubs in its genus. Planting

unadapted ecotypes on disturbances reduces survival;

however, site factors affecting adaptation are not well

understood.

The plant can be used as a nurse crop to improve

seedling establishment of tree seedlings. It is also

relatively abundant in localized areas and should be

maintained for game habitat and ground cover. This

species frequently increases following logging and

becomes important for watershed protection.

Varieties and Ecotypes—None.

Family Scrophulariaceae _________

Penstemon fruticosus

Bush penstemon

Description—Bush penstemon is one of approximately 230 species of Penstemon that occur in Western

North America (Hylton 1974). Most are herbaceous

plants, but the list includes a few suffrutescent or

woody shrubs. Bush penstemon is a woody species that

normally reaches a height of 2.5 ft (76 cm). Plants form

a persistent, woody crown that is usually wider than

tall. They develop into wide, dense clumps when young

and spread by stem layering, but natural dieback

leaves the center open. Stems are erect, and clusters of

leaves form at the base of current year’s stems (Hitchock

and Cronquist 1976). Leaves are mostly lustrous,

evergreen, lanceolate or oblanceolate to elliptic, entire or serrulate or denticulate, 0.4 to 2 inches (1 to

5 cm) long, and 0.2 to 0.6 inch (5 to 15 mm) wide.

Flowers are bright lavender blue, 1 to 1.5 inches (25

to 38 mm) long, 0.3 to 0.5 inch (7 to 12 mm) wide (Davis

1952). Three varieties were described by Hitchcock

and Cronquist (1973); they differ in regard to leaf

length, width, margins, and size of flowers. Penstemon

fruticosus var. serratus is mainly confined to the Snake

River Canyon of Oregon and Idaho; P. f. var. scouleri

occurs in northeastern Washington, northern Idaho,

and adjacent British Columbia; and P. f.var. fruticosus

is more widespread and variable, but generally does

not occupy the range of the two other varieties

(Hitchcock and Cronquist 1973).

Ecological Relationships and Distribution—

Bush penstemon generally occurs at elevations over

4,000 ft (1,200 m) on rocky slopes in forest openings

from the Cascades of Washington and Oregon across

Idaho to the Rocky Mountains of Montana and

Wyoming (Davis 1952). Although widespread, this

species is not particularly abundant. It is not a principal understory with any forest community in the

Intermountain region (Mueggler 1988), northern Idaho

(Cooper and others 1987), or eastern Washington
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 695

Chapter 23 Shrubs of Other Families

(Daubenmire and Daubenmire 1968). Plants normally

occur on ridge crests, talus slopes, and forest openings.

They invade roadcuts and are noticeable on abandoned disturbances. Bush penstemon is encountered

in scattered stands of ponderosa pine and Douglas-fir.

At higher elevations it exists in openings with subalpine fir. It does not occur in dense shrub fields or mixed

grass/herb communities.

Plant Culture—Bush penstemon usually occurs as

scattered plants on rocky outcrops (fig. 59). Following

disturbances or clearing of overstory species, this

shrub flourishes. It invades openings and forms patches

dominating small rocky areas. Young plants are vigorous and robust. Disturbances often become excellent

seed production centers.

Plants are evergreen; a high percentage of leaves

remain on the shrub throughout the winter. New

growth begins early in spring before most other plants

initiate growth. Leaves develop rapidly to form a dark

green clump.

Plants usually produce a profusion of flowers each

year. Flowering begins in early June and may continue

for nearly 1 month. Seeds ripen from mid-August to

late September. Numerous seeds develop each year,

but not all are viable. Seed quality varies widely

among collection sites and years. Although the plant

flowers for an extended period, seeds ripen fairly

uniformly and remain in the capsule for nearly a

month before shattering.

Seeds are collected by hand stripping the dry capsules, or by clipping the inflorescences. They must be

harvested soon after the capsule dries, but before it

opens and seeds shatter. Sites that are good seed

producers have remained so for nearly 20 years. Most

plants within a stand produce good seed crops each

year. Seeds are easily harvested and can be cleaned by

allowing the capsules to dry and dehisce, releasing the

seeds. Seeds are small, but can be separated from the

debris using a fanning mill or gravity table.

Some seeds will germinate soon after harvesting if

exposed to favorable conditions, but a period of

afterripening and wet prechilling is normally required

to assure uniform germination. Seeds that are fall

seeded receive natural wet prechilling and germinate

uniformly in spring. Bush penstemon seedlings are

vigorous and normally grow rapidly.

New seedlings spread quickly from existing plants.

Although seedlings increase rapidly, the young plants

are not highly competitive with grasses and broadleaf

herbs. New seedlings are well adapted to harsh sites

and compete well with other species under these

conditions, but they are not able to compete with

grasses and broadleaf herbs on deep, fertile soils.

Seedlings establish well by broadcast seeding on a

rough seedbed or by drill seeding at shallow depths,

about 0.24 inch (6 mm) on a firm seedbed. Seed

germination and establishment occur uniformly; erratic stands are uncommon.

Bush penstemon is well adapted to drill or broadcast

seeding. The seeds can be planted separately or in

mixtures with most drills or broadcast seeders. The

seeds are small and need not be planted at rates

exceeding 2 to 3 lb per acre (2.2 to 3.4 kg per ha).

Seeding at a rate of 2 to 4 lb per acre (2.2 to 3.4 kg per

ha) is recommended for drill seeding. Broadcast seeding on unprepared road disturbances may be increased

to 4 to 6 lb per acre (4.5 to 6.7 kg per ha), depending on

whether other species are seeded and the condition of

the seedbed. This is one species that responds well to

the amount of seed sown. Increasing the seeding rate

usually results in an increase in the number of plants

to establish. Thus, plant density of this species can be

regulated by adjusting the seeding rate. If seeded

concurrently with species having larger seeds, it is

recommended that the smaller seeds of bush penstemon be planted in separate drill rows to better regulate planting depth.

Small seedlings are quite vigorous. When seeded in

barren openings, bush penstemon matures within 2 to

3 years. When grown in competition with grasses, the

young plants can be seriously stunted. However, bush

penstemon can be planted with certain grasses and

forbs. Germination and seedling emergence of this

shrub occur at about the same time as most herbs.

Consequently, bush penstemon seedlings are able to

compete favorably with many native grasses and broadleaf forbs. Bush penstemon seedlings are not able to

persist when seeded with a dense cover of aggressive

Figure 59—Bush penstemon, a low-growing mounding shrub, generally grows on rocky outcrops.
696 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

grasses, including smooth brome, intermediate wheatgrass, or timothy.

Bush penstemon grows quickly from direct seeding

or transplanting. Transplants can be reared as bareroot

nursery stock, greenhouse-grown container stock, or

rooted stem cuttings.

Bush penstemon seedlings begin growth early in

spring, often before nursery lifting is completed. If

bareroot stock initiates growth prior to lifting, field

survival of the transplants declines significantly. Fall

lifting and overwinter storage may be most practical.

Plants stored in bags or crates should have the tops

exposed. Plants can be fall lifted and planted in sawdust or other media for overwinter storage in open lath

houses. Bareroot stock transplants very well and can

provide effective ground cover within 1 to 2 years.

Bush penstemon is easily grown from seed or stem

cuttings. However, plants grow rapidly and can be

easily stunted if reared in containers that are too

small. Container stock grows best in a well-drained,

neutral potting media. Stem cuttings taken in midwinter and early spring root readily.

Uses and Management—Bush penstemon is well

adapted for revegetation of road disturbances, mine

sites, and other drastic disturbances (Hungerford

1984; Monsen and Christensen 1975; Plummer 1977;

Tiedemann and others 1976). From roadway plantings

in northwestern Montana, Hungerford (1984) rated

bush penstemon as the second most useful species for

revegetation of forest roads. Only Woods’ rose exceeded this species in overall usefulness. The plant is

extremely vigorous, matures quickly, and spreads

rapidly by natural seeding and crown spread

(Hungerford 1984; Monsen and Christensen 1975). It

provides excellent ground cover due to its dense, lowprofile growth habit. It furnishes effective protection

throughout the entire year, as plants are evergreen,

and provides aboveground protection at all seasons. It

is able to persist and spread when buried by shifting

soil. It is one of few species that is well adapted to both

rocky, shallow soils, as well as deep, well-developed

soils. This species can be successfully seeded or transplanted on rocky roadcuts. It establishes very well

from broadcast seedings with minimal soil coverage.

Few other species, including most grasses or broadleaf

herbs, establish as successfully from broadcast seeding as this shrub.

Bush penstemon is an attractive shrub that flowers

throughout most of the summer and provides an attractive leafy cover. It is able to spread and occupy

rocky sites, providing an interesting pattern of ground

cover. Both the flowers and leaves furnish an attractive background for other species. The shrub can be

used to treat roadways, mine sites, and recreational

areas where aesthetic values are important. It has

considerable usefulness for formal plantings in landscaping commercial or residential sites. Its evergreen

growth habit suggests this native shrub would be

particularly useful for planting around summer homes

and dwellings to prevent the spread of wildfires.

This species is also an important plant for wildlife

habitat. It occupies sites that are open and exposed

during midwinter. Game animals are often confined to

these sites and rely heavily on this shrub. It has

considerable promise for many situations, and could

be more widely used in restoration programs if seeds

were more available.

Varieties and Ecotypes—None.

Family Ulmaceae ________________

Celtis reticulata

Netleaf hackberry

Description—Celtis is a large genus of about 70

species of shrubs and trees of the Northern Hemisphere. Netleaf hackberry is a variable taxon, and

includes varieties that are difficult to separate due to

the many intergradations encountered (Harrington

1964). Netleaf hackberry, also known as hackberry,

western hackberry, and palo blanco, is a large deciduous shrub to small globose tree growing to 30 ft (9.1 m)

in height and 6 to 12 inches (15 to 30 cm) in diameter,

depending on available water (Bonner 1974a; Hitchcock

and others 1973). Crowns are open and irregular with

stout, ascending branches. Young twigs are glabrous

to pubescent. Bark of older branches is reddish gray

with prominent ridges and furrows. Plants develop a

spreading and fibrous root system. Leaves are short

petiolate and leathery with scabrous surfaces and

entire to serrate margins. They are ovate or ovatelanceolate, usually somewhat acuminate and acute to

chordate at the base, 1 to 4 inches (2.5 to 10.2 cm) long

and 0.74 to 1.5 inches (1.9 to 3.8 cm) wide. Leaf

surfaces are dull green to yellow green, but paler with

conspicuous reticulate veinlets below. They are three

nerved near the base and short pilose along the veins.

Foliage turns yellow in fall (Hitchcock and others

1973; Welsh and others 1987).

Plants are polygamo-monoecious with greenish flowers that emerge with the leaves clustered on short

shoots of the current season. Flowers develop on slender, pubescent pedicels. Lower flowers are mostly

staminate and several per axil. Upper flowers are one

or two per axil and mostly perfect, but some may be

pistillate. Perfect flowers consists of five membranous

perianth segments, five stamens, and a one-seeded

ovary with spreading style tips. Fruit is an orange-red

to yellow, thin-walled, globose drupe with a datelike

flavor (Treshow and others 1970).
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Chapter 23 Shrubs of Other Families

Ecological Relationships and Distribution—

Netleaf hackberry grows on dry foothills, open slopes,

rocky bluffs, watercourses, and canyons from the Snake

River drainage of the Northwestern States, east to

Colorado, and south to west Texas and southern California (Bonner 1974a; Preston 1968; Thornburg 1982).

It grows at elevations ranging from 2,000 to 6,000 ft

(610 to 1,800 m) (Sutton and Johnson 1974; Thornburg

1982) on well-drained soils ranging from silty to rocky,

shallow to deep, and neutral to calcareous (Sutton and

Johnson 1974; Thornburg 1982; Van Dersal 1938).

Plants grow in full sunlight and are drought tolerant

and wind resistant (fig. 60). Netleaf hackberry is well

adapted to watercourses and canyons where it may

grow on limestone and gravelly and rocky soils

(Thornburg 1982).

Netleaf hackberry occurs as scattered plants in big

sagebrush and mountain brush communities at 3,000

to 5,000 ft (910 to 1,500 m) in Utah (Welsh and others

1987). It is prevalent on steep south and west slopes

with pinyon-juniper, antelope bitterbrush, Stansbury

cliffrose, and other mountain brush species throughout the Wasatch Mountains. It is particularly important on exposed slopes with other persistent species.

Plant Culture—Flowering occurs in March and

April; fruits ripen in late fall, and seed disperses in fall

and winter (Swingle 1939). Fruits persist on the trees

for several months after ripening, providing a prolonged period for collection. Although this plant often

grows in harsh environments, some seeds are usually

produced every year. They are gathered by hand or by

knocking them onto tarps. Collection may be easier in

winter after leaves have fallen (Bonner 1974a). Drying

may not be required if seeds are collected late in the

season, but seeds are more difficult to remove from dry

fruits. Twigs and other debris are removed by fanning.

Extraction of the pulp by macerating the fruits in

water apparently aids in improving germination.

Bonner (1974a) reported that fermenting the fruits for

3 days at room temperature prior to depulping and wet

prechilling improved germination. Approximately

80 lb (36.3 kg) of seed may be recovered from 100 lb

(45.4 kg) of fruits; there are about 4,870 cleaned seeds

per lb (10,736 per kg) (Swingle 1939). Dry fruits or

cleaned seed may be stored in sealed containers at

41 ∞F (5.0 ∞C). Seeds require either fall sowing or a

120-day laboratory wet prechilling at 41 ∞F (5.0 ∞C)

prior to spring sowing in the nursery (Hartmann and

others 1990).

Germination is usually quite low as seed from most

wildland collections has poor fill (Bonner 1974a). Undeveloped seeds are difficult to separate and remove

during cleaning.

Seedlings grow slowly and are sensitive to competition from herbs. Nursery grown transplants and rooted

cuttings are often recommended for field plantings to

reduce problems associated with establishment from

seed. Nursery or container-grown stock develops well,

and field survival is excellent. One or 2-year-old planting stock is recommended for outplantings.

Seed and planting stock are rarely available on the

commercial market; thus, it is necessary to arrange for

collection of seed or propagation of seedlings. Although extensive testing of different ecotypes has not

been reported, considerable differences in vegetative

traits and areas of occurrence would suggest that

planting success is dependent on the use of an adapted

seed source.

Uses and Management—Netleaf hackberry currently receives limited use in artificial plantings. However, it is one of the tallest native species on sites

within its range of adaptation, and provides important

habitat for wildlife. It occupies exposed windblown

slopes where big game animals concentrate, providing

important cover and protection. The shrub is considered fair to good forage for deer and livestock (Sampson

and Jespersen 1963). It frequently grows as clumps or

in groups to furnish pockets of cover where few other

shrubs are able to grow.

The species grows as a border along riparian zones

and significantly enhances these communities. It is

particularly adapted to steep slopes and well-drained

sites aligning stream bottoms. Fruits persist on the

tree well into the winter and provide food for birds

(Thornburg 1982). Montezuma quail reportedly eat

the seeds during winter months (Bishop and

Hungerford 1965). Gambel quail also forage on the

fruits (Gullion 1964; Martin and others 1951), as do

various small mammals (Bailey 1936).

This species has established well from plantings on

big game ranges in central and south-central Utah

when planted on shallow soils and exposed slopes

within the lower mountain brush zone. It has been

Figure 60—Netleaf hackberry growing on a dry, open

slope protected by a rocky outcrop.
698 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Chapter 23 Shrubs of Other Families

planted in groups or patches to provide concealment

and winter cover in areas where upright vegetation is

required. Game animals and livestock may graze new

plantings, but not seriously enough to adversely affect

plant growth. This species has survived years of drought

without appreciable reduction in plant size and vigor.

Once established, the young plants are very hardy and

persistent. They are able to compete and grow with

understory herbs.

Netleaf hackberry can be used to provide shade in

recreation sites, and for windbreaks, shelterbelts, or

domestic landscaping. It can be used on sites receiving

at least 14 to 16 inches (36 to 41 cm) of annual precipitation. It does not require maintenance in landscape

plantings, and could be more widely used in these

situations.

Young plants do not grow rapidly. More rapidly

developing species are recommended for unstable

watershed or mine disturbances. This plant provides

important ground cover on sites where high-intensity

summer storms may occur. Plants are generally very

hardy and long lived. Twigs and leaves are frequently

infested by insect galls (Welsh and others 1987), but

vigor is not seriously affected. Plants are not especially

fire tolerant, and can be killed by wildfires, although

resprouting does occur. Many native stands persist on

rocky sites that provide some protection from wildfires. Plants that do survive fires usually recover

slowly. Plants do not spread rapidly by natural seeding. Few new seedlings appear following fires or other

disturbances. Seedlings appear to require shading,

but seedbed requirements are not well understood.

Improved Cultivars and Other Species—None.

In the Midwest, “Oahe,” a released cultivar of Celtis

occidentalis, has been used as substitute for Siberian

elm, Russian-olive, and green ash, although vigor and

ease of establishment are lower (Soloman 1985). The

cultivar has received little testing in the Intermountain region.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-1

Index

A

Acer glabrum See maple, Rocky Mountain

Achillea millefolium lanulosa See yarrow, western

Achnatherum hymenoides See ricegrass, Indian

Achnatherum thurberianum See needlegrass, Thurber

acid equivalent 97

aerial broadcasting 77, 205

aerial ignitors 85

aerial seeding 77–79, 205

fixed-wing aircraft 78

helicopter 78–79

aesthetics 133

Agropyron cristatum See wheatgrass, crested; wheatgrass, crested fairway

Agropyron cristatum x A. desertorum See wheatgrass,

crested hybrid

Agropyron dasystachyum See wheatgrass, thickspike

Agropyron desertorum See wheatgrass, crested

standard

Agropyron elongatum See wheatgrass, tall

Agropyron fragile See wheatgrass, crested Siberian

Agropyron inerme See wheatgrass, bluebunch;

wheatgrass, bluebunch bear

Agropyron intermedium See wheatgrass, intermediate

Agropyron repens x A. spicatum See wheatgrass,

hybrid: NewHy

Agropyron sibiricum See wheatgrass, crested Siberian

Agropyron smithii See wheatgrass, western

Agropyron spicatum See wheatgrass, bluebunch

Agropyron spicatum x A. dasystachyum See wheatgrass, hybrid: SL1

Agropyron trachycaulum See wheatgrass, slender

Agropyron trichophorum See wheatgrass, intermediate

Agrostis alba See bentgrass, redtop

Agrostis palustris See bentgrass, redtop

Agrostis stolonifera See bentgrass, redtop

air-screen separator 715

Aira cespitosa See hairgrass, tufted

alder, thinleaf 266–267, 270, 272, 601, 616–618, 748

alfalfa 132, 135, 136, 138, 142, 182, 183, 196, 206,

216, 220, 237, 242, 259, 264, 269, 271, 273, 275,

277, 280, 282, 284, 287, 289, 425, 448, 449–450,

702, 727, 741

Drylander 451

Ladak 450–451, 724, 728

Nomad 450, 724

Rambler 450

Rhizoma 451

Runner 451

Spreader 451

Spreader II 451

Teton 451

Travois 451

alfalfa, range type 427

alfalfa, sicklepod 447–448

alfileria 291, 427

Allenrolfea occidentalis See iodine bush

Alnus incana See alder, thinleaf

Alopecurus arundinaceus See foxtail, creeping

Alopecurus pratensis See foxtail, meadow

Alopecurus ventricosus See foxtail, creeping

Amelanchier alnifolia See serviceberry, Saskatoon

Amelanchier utahensis See serviceberry, Utah

ammonium 54

anchor chain 68–69

disk-chain 69

Dixie sager 69–70

Ely 66, 68, 70

smooth 66

swivels 231–232

Andropogon curtipendula See grama, sideoats

angelica, small leaf 269, 427

animal control See depredation

animal depredation See depredation

annuals, spring-growing 257

annuals, summer-growing 258

Apache plume 253, 273, 276, 293, 559, 561–563, 601,

704, 724, 748

Arctostaphylos uva-ursi See manzanita, bearberry

Aristida purpurea See threeawn, red

Aroga websteri See moth, sagebrush defoliator

Arrhenatherum elatius See oatgrass, tall

Artemisia abrotanum See wormwood, oldman

Artemisia arbuscula See sagebrush, low

Artemisia bigelovii See sagebrush Bigelow

Artemisia cana See sagebrush, silver

Artemisia filifolia See sandsage

Artemisia frigida See sage, fringed

Artemisia longiloba See sagebrush, alkali

Artemisia ludoviciana See sage, Louisiana

Artemisia nova See sagebrush, black

Artemisia pygmaea See sagebrush, pygmy

Artemisia rigida See sagebrush, stiff

Artemisia spinescens See budsage

Artemisia tridentata See sagebrush, big

Artemisia tripartita See sagebrush, threetip

artificial seeding 516, 539

ash, singleleaf 276, 278, 601, 650–652, 704, 748

aspen 119, 195, 214, 266–267, 748

aspen communities 118, 215, 308–309

aspen low forb 118

aspen shrub 118

aspen tall forb 118

aspen-conifer 195, 213, 269
Index-2 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

mixed forb 118

mixed shrub 118

aspen-conifer communities 9, 216, 601

aspen openings 204–205

aspen openings, upper elevation 201, 259

aster 196, 202, 223, 237, 246–248, 252, 255–256,

259, 261–262, 264, 269, 271–272, 275, 277–278,

280, 282, 289, 294, 741

aster, alpine leafybract 259, 261

aster, blueleaf 135, 261, 269, 271, 277, 289, 426–427,

433, 702, 724

Aster chilensis See aster, Pacific

aster, Engelmann 269, 289, 427, 702, 724

Aster glaucodes See aster, blueleaf

aster, Pacific 259, 261–262, 264, 269, 271, 275, 277,

280, 282, 294, 426–427, 431–432, 702, 724, 763

aster, smooth 259

Astragalus cicer See milkvetch, cicer

astragalus, giant 731

Atriplex buxifolia 481–482 See also saltbush, Gardner

Atriplex canescens See saltbush, fourwing

Atriplex confertifolia See shadscale

Atriplex corrugata See saltbush, mat

Atriplex cuneata See saltbush, Castle Valley clover

Atriplex falcata 481–482 See also saltbush, Gardner

Atriplex gardneri See saltbush, Gardner

Atriplex hymenelytra See saltbush; saltbush, desert

holly

Atriplex lentiformis See saltbush; saltbush, quailbush

Atriplex obovata See saltbush; saltbush, broadscale

Atriplex polycarpa See saltbush; saltbush, allscale

Atriplex tridentata 481–482 See also saltbush,

Gardner

auger 765

Avena elatior See oatgrass, tall

B

Balsamorhiza hookeri See balsamroot, hairy

Balsamorhiza macrophylla See balsamroot, cutleaf

Balsamorhiza sagittata See balsamroot, arrowleaf

balsamroot, arrowleaf 132, 135, 138–139, 142, 

182–183, 185, 196, 271, 275, 277, 282, 287, 

426–427, 435–436, 702, 724, 727–728, 741

balsamroot, cutleaf 271, 277, 287, 426–427, 436, 702,

724, 727, 741

balsamroot, hairy 271, 277, 426–427, 436

barberry, creeping 601, 704, 748 See also Oregon

grape

barberry, Fremont 614, 704

barberry, shining 613

bareroot stock 739–742, 747, 754–755, 763–767

barley, beardless 311

barley, bulbous 308, 311, 700

barley, common 311

barley, foxtail 269, 311

barley, meadow 259, 261, 264, 265, 269, 289, 308,

311

bassia, fivehook 264

bentgrass, carpet 310 See also bentgrass, redtop

bentgrass, creeping 310 See also bentgrass, redtop

bentgrass, redtop 208, 259, 261, 264, 265, 269, 308,

310, 345–346, 700

Streaker 298, 346

Betula glandulosa See birch, bog

Betula occidentalis See birch, water

Betula papyrifera See birch, paper

big game depredation See depredation

big sagebrush-cheatgrass 106

bighorn sheep 165–166

biological insect control 721

birch, bog 601, 618–620

birch, paper 601, 622, 748

birch, water 262, 266, 267, 601, 620–621

bitterbrush, antelope 103, 132, 135, 138, 142, 143,

181, 182, 183, 185, 196, 270, 272, 273, 276, 278,

280, 283, 288, 557, 559, 581–586, 587, 601, 704,

724, 727, 730, 741, 748

Lassen 586

bitterbrush, desert 273, 276, 293, 559, 579–581, 582,

601, 704, 724, 727, 748

blackbrush 195, 199, 232, 246, 247, 252, 253, 254,

257, 258, 293, 294, 552–555, 601, 704, 748

blackbrush communities 9, 252–253, 293, 308, 601

blackcap 751 See raspberry, black

blackthorn 577

bladersenna, common 741

bluebell 741

bluebell, tall 259, 269, 427, 452, 453

bluegrass 102

bluegrass, alpine

Gruening 298

bluegrass, big 228, 259, 269, 271, 275, 277, 282, 289,

308, 401–402, 700

Sherman 298, 402

bluegrass, bulbous 311

bluegrass, Canada 259, 269, 271, 275, 277, 282, 289,

308, 311, 402–404, 700

Foothills 298, 404

Reubens 298, 404

bluegrass, Cusick’s 311

bluegrass, Kentucky 196, 269, 308, 311, 406–408, 724

Newport 138, 298, 408

bluegrass, longtongue 311

bluegrass, mutton 308, 311, 404–406

bluegrass, nodding 311

bluegrass, pine 311

bluegrass, Sandberg 135, 182–183, 185, 264–265,

273, 275, 277, 279, 280, 282, 284–287, 291–292,

294, 308, 311, 408–410, 700

Canbar 298, 410
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-3

High Plains germplasm 298, 410

Service 298, 410

boron 51, 52, 54

bouncing-bet 427

Bouteloua curtipendula See grama, sideoats

Bouteloua eriopoda See grama, black

Bouteloua gracilis See grama, blue

boxelder 602

Brillion Seeder 80

broadcast ground application 93

broadcast seeder 72, 76–77

broadcast seeding 76, 141

broadcast spray application 92

brome, California 310 See also brome, mountain

Cucamonga 298

brome, cheatgrass See cheatgrass

brome, downy See cheatgrass

brome, erect 310

brome, foxtail 310

brome, fringed 358–360

brome, Hungarian See brome, smooth

brome, meadow 259, 264–265, 269, 289, 310, 

365–367, 700

Fleet 367

Paddock 367

Regar 196, 269, 271, 275, 277, 282, 287, 298, 310,

367, 724, 728

brome, mountain 135, 138, 196, 259, 261, 264, 265,

269, 271, 289, 308, 310, 360–362, 700, 724

Bromar 298, 362

Garnet 362

brome, nodding 308, 310, 358–360

brome, red 254, 255, 310

brome, red communities 9, 253–256, 294

brome, smooth 132, 135, 138, 142, 182–183, 185,

196, 205, 259, 261, 264–265, 269, 271, 275, 277,

282, 287, 289, 310, 363–366, 700, 727, 728, 731

Achenbach 299

Lincoln 299, 365

Manchar 299, 365

northern 259, 308

Polar 299

southern 308, 724

varieties 365

brome, subalpine 259, 269, 289, 308, 700

Bromus anomalus See brome, nodding

Bromus biebersteinii See brome, meadow

Bromus carinatus See brome, California; brome,

mountain

Bromus ciliatus See brome, fringed

Bromus erectus See brome, meadow

Bromus inermis See brome, smooth

Bromus marginatus See brome, mountain

Bromus riparius See brome, meadow

Bromus tomentellus See brome, subalpine

brushland plow 66

buckthorn, cascara 266–267, 601, 704

buckwheat 652–653 See also eriogonum

buckwheat, wild California flattop 704

budsage 245–246, 291, 495, 511–513 See also

sagebrush, budsage

budsage communities 245

buffaloberry, roundleaf 273, 293, 601, 643–644, 704,

748

buffaloberry, russet 601, 642–643, 704, 748

buffaloberry, silver 262, 266–267, 601, 641–642, 704,

731, 748

bulrush, saltmarsh 263

bulrush, tule 263

Bureau of Plant Industry 2, 3

burnet, small 132, 135, 138, 142, 182–183, 185, 196,

271, 273, 275, 277, 279, 280, 282, 284, 286, 287,

294, 425, 427, 457–459, 702, 724, 728, 741, 763

Delar 459

burning, prescribed 101, 106, 160, 168

aspen communities 118–119

cheatgrass 116

effects 105

pinyon-juniper 116–117

postfire management 119

sagebrush 115–116

uses 101

C

cables 67, 68, 69 See also anchor chain

Calamagrostis canadensis See reedgrass, bluejoint

calcium 51, 54, 178–179

calcium nitrate 54

camphorfume, Mediterranean 469, 491

Camphorosoma monspeliaca See camphorfume,

Mediterranean

canarygrass, reed 135, 182–184, 196, 261, 308, 311,

396–398

Ioreed 299, 397

cattle 176, 178–179

Ceanothus 656

Ceanothus cuneatus See ceanothus, wedgeleaf

ceanothus, deerbrush 601, 658–659, 704, 748

ceanothus, desert 657–658

ceanothus, Fendler 273, 276, 601

Ceanothus fendleri See ceanothus Fendler

Ceanothus greggii See ceanothus, desert

Ceanothus integerrimus See ceanothus, deerbrush

ceanothus, Lemmon 659

Ceanothus lemmonii See ceanothus, Lemmon

ceanothus, Martin 143, 196, 222, 272, 278, 283, 290,

601, 661–662, 704, 724, 727, 748

Ceanothus martinii See ceanothus, Martin

ceanothus, prostrate 601, 659–661, 705, 748

Ceanothus prostratus See ceanothus, prostrate
Index-4 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

ceanothus, redstem 142–143, 222, 266–267, 272,

601, 662–665, 668, 704, 748

Ceanothus sanguineus See ceanothus, redstem

ceanothus, snowbrush 196, 272, 283, 288, 290, 601,

665–668, 705, 748

Ceanothus velutinus See ceanothus, snowbrush

ceanothus, wedgeleaf 601, 656–657, 705, 749

Celtis reticulata See hackberry, netleaf

Ceratoides lanata See winterfat

Ceratoides lanata subspinosa See winterfat, foothills

Ceratoides latens See winterfat, Pamirian

Cercocarpus betuloides See mountain mahogany,

birchleaf

Cercocarpus intricatus See mountain mahogany,

littleleaf

Cercocarpus ledifolius See mountain mahogany,

curlleaf

Cercocarpus montanus See mountain mahogany, true

certification, seed 734

certified seed See seed, certified

chain See anchor chain

chaining 70, 228–229, 231, 233

cheatgrass 102, 107, 254–255, 310

cheatgrass communities 9, 253–256, 294, 601

checker-mallow, Oregon 264

chemical plant control 62

chenopod 467

cherry, Bessey 573–574, 601, 705, 749

cherry, bitter 217, 222, 272, 574–575, 601, 705, 749

cherry, Nanking 577

cherry, western sand See cherry, Bessey

chokecherry 135, 142–143, 196, 266–267, 270, 272,

278, 283, 288, 290, 577–579, 601, 705, 724, 730,

749

Chrysothamnus See rabbitbrush

Chrysothamnus albidus See rabbitbrush, alkali

Chrysothamnus depressus See rabbitbrush, dwarf

Chrysothamnus greenei See rabbitbrush, Greene’s

Chrysothamnus linifolius See rabbitbrush, spreading

Chrysothamnus nauseosus See rabbitbrush, rubber

Chrysothamnus parryi See rabbitbrush, Parry

Chrysothamnus vaseyi See rabbitbrush, Vasey

Chrysothamnus viscidiflorus See rabbitbrush, low

chukar partridge 170

cinquefoil 749

cinquefoil, bush 601, 705 See also cinquefoil, shrubby

cinquefoil, gland 259, 261, 264, 427

cinquefoil, shrubby 260, 262, 266, 267, 270, 290, 

569–571

Clematis ligusticifolia See virginsbower, western

cliffrose, Stansbury 132, 135, 138, 142–143, 185, 196,

272–273, 276, 278, 280, 283, 556–559, 562, 579,

601, 705, 725, 727, 730, 741, 749

climate 29, 34–35, 124

clover 463

clover, alsike 261, 264, 427, 464, 702

Aurora 464

Dawn 464

clover, Castle Valley See saltbush, Castle Valley clover

clover, red 464

Dollard 464

Kenland 464

Lakeland 464

Pennscott 464

clover, strawberry 261–262, 264, 427, 463–464, 702,

724

clover, white 464

coarse fragment content 41, 43

cold tolerance 128

Coleogyne ramosissima See blackbrush

columbine, Colorado 259, 269, 427

communities, plant 199 See also site preparation and

seeding prescriptions; vegetation types

companion species 62

compatibility

perennial grasses 122

seeding and species 122, 133

competition, plant See plant competition

competitiveness of species 259, 261–262, 269, 271,

273, 279–280, 282, 284–287, 289, 291–293

composite shrubs 493

container stock 739–742, 747, 754, 757, 764

copper 51–52, 54

Cornus sericeus See dogwood, redosier

Cornus stolonifera See dogwood, redosier

Coronilla varia See crownvetch

Cotoneaster acutifolia See cotoneaster, Peking

cotoneaster, Peking 272, 278, 555–556, 705

cottonwood, Fremont 266–267

cottonwood, narrowleaf 266–267, 749

cover See wildlife habitat, cover

Cowania mexicana See cliffrose, Stansbury

Cowania stansburiana See cliffrose, Stansbury

cowparsnip 216, 259, 264, 269, 426–427, 441–442,

702, 724, 727

Crataegus douglasii See hawthorn, Douglas

crown divisions 762

crownvetch 135, 196, 259, 264, 269, 271, 277, 282,

289, 426–427, 437–438, 702, 724, 731, 741

Chemung 438

Emerald 438

Penngift 438

crude fat 177, 180

crude fiber 177, 180

crude protein 177, 180

cultivars 129, 138, 298, 734

Cupressus arizonica See cypress, Arizona

curly grass See galleta

currant, golden 135, 143, 196, 266–267, 601, 

690–691, 705, 725, 727, 741, 749
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-5

currant, sticky 260, 602, 693–694, 705, 749

currant, wax 260, 602, 691–693, 705, 749

cut stump treatment 92

cuttings, hardwood 761–762

cuttings, stem 739, 741

cypress, Arizona 276, 278, 602, 632–633, 705, 749

D

Dactylis glomerata See orchardgrass

daisy, oxeye 427

debearder 713

deer 159, 161, 176, 515

deervetch, birdfoot 427

depredation 144–145, 163, 768

Deschampsia caespitosa See hairgrass, tufted

desert almond 572–573

desert blite 491 See also sumpweed, desert

dibbles 764–765

digestibility 181

disease, plant 601

disks 66–69

Distichlis spicata See saltgrass, inland

dixie sager 67, 69, 70

dogwood, creek See dogwood, redosier

dogwood, redosier 262, 266–267, 602, 630–632, 705,

749

dozers and blades 84

drags 71

drill seeding 205

drills 73

horizon 75

no-till 75

rangeland 73–75, 228

Truax 75

Tye 75

dropseed, sand 135, 142, 182–183, 185, 196, 262,

273, 279–280, 284, 291, 293, 308, 311, 414–416,

700

drought tolerance 127–128

Dybvig separator 714

E

ecotypes 126, 200

edges, vegetative 111

elaeagnus, autumn 602, 705

Elaeagnus commutata See silverberry

elderberry, blue 135, 143, 196, 270, 272, 276, 278,

283, 288, 290, 602, 624–625, 705, 741, 748

elderberry, red 260, 266–267, 270, 290, 602, 625–627,

705

elk 111, 158–164, 176

Elymus angustus See wildrye, Altai

Elymus canadensis See wildrye, Canada

Elymus cinereus See wildrye, Great Basin

Elymus dahuricus See wildrye, Dahurian

Elymus elymoides See squirreltail, bottlebrush

Elymus giganteus See wildrye, mammoth

Elymus glaucus See wildrye, blue

Elymus junceus See wildrye, Russian

Elymus lanceolatus lanceolatus See wheatgrass,

thickspike

Elymus lanceolatus wawawai See wheatgrass,

bluebunch; wheatgrass, Snake River

Elymus salinus See wildrye, Salina

Elymus simplex See wildrye, alkali

Elymus trachycaulus See wheatgrass, slender

Elymus triticoides See wildrye, creeping

Elymus wawawaiensis See wheatgrass, Snake River;

wheatgrass, bluebunch

Elytrigia repens x Pseudoroegneria spicata See

wheatgrass, hybrid: NewHy

energy 176–177

ephedra, green 132, 135, 142–143, 196, 272–273,

276, 278–280, 283–285, 602, 645–646, 705, 725,

727, 730, 741, 749

ephedra, Nevada 273, 276, 278–279, 281, 284–285,

293, 602, 644–646, 705, 725, 727, 749

Ephedra nevadensis See ephedra, Nevada

ephedra, Torrey 705, 749

Ephedra viridis See ephedra, green

eriogonum, cushion 269, 271, 277, 427, 705

Eriogonum heracleoides See eriogonum, Wyeth

eriogonum, sulfur 272, 283, 602, 652, 653–654, 749

Sierra sulfur buckwheat 654

Eriogonum umbellatum See eriogonum, sulfur

eriogonum, Wright 652, 654

Eriogonum wrightii See eriogonum, Wright

eriogonum, Wyeth 143, 196, 276, 278, 283, 602, 652,

652–653, 706, 727, 749

establishment 131

esthetics 60

F

Fallugia paradoxa See Apache plume

fences 172

fertilizers 53, 54, 55

application 55–56

elemental composition 54

fescue, Alpine See fescue, sheep

fescue, Arizona 311

Redondo 299

fescue, bluebunch See fescue, Idaho

fescue, desert 700

fescue, hard See fescue, hard sheep

fescue, hard sheep 132, 135, 138, 142, 196, 269, 271,

273, 275, 277, 280, 282, 284, 286–287, 289, 294,

308, 311, 389, 700, 724, 728

Durar 299, 390

fescue, Idaho 102, 139, 182–184, 280, 282, 284, 

286–287, 294, 308, 311, 386–388, 700
Index-6 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Joseph 299, 388

Nezpurs 299, 388

fescue, meadow 308, 311, 731

fescue, red 132, 269

fescue, reed 308, 384 See also fescue, tall

fescue, sheep 271, 273, 280, 282, 287, 289, 308, 311,

388, 700

Covar 299, 390

MX86 299

sulcate 259, 271, 275, 277, 280, 282, 389

fescue, spike 308, 311

fescue, tall 209, 259, 262, 264–265, 311, 384–386

Alta 269, 299, 311, 385

Fawn 299, 385

varieties 386

fescue, Thurber 269, 308, 311

Festuca arizonica See fescue, Arizona

Festuca arundinacea See fescue, tall

Festuca idahoensis See fescue, Idaho

Festuca ovina See fescue, sheep

Festuca ovina duriuscula See fescue, hard sheep

filaree See alfileria

fire effects 102–104, 107, 113–114

big sagebrush-cheatgrass 106

bitterbrush, antelope 103

bluegrass 102

cheatgrass 102

fescue, Idaho 102

habitat management 113

horsebrush 103

intensity 113

junegrass, prairie 102

mountain mahogany, curlleaf 103

needlegrass 102

oak, Gambel 103

plant succession 114

rabbitbrush 103

reedgrass, plains 102

ricegrass, Indian 102

riparian 112

sagebrush 103

severity 113

snowberry 103

squirreltail, bottlebrush 102

wheatgrass 102

fire ignitor 85

aerial ignitors 85

flame thrower 85

helitorch 85

ping-pong ball injector 85

teratorch 85

fire intensity 113

fire severity 113

firebreak 114, 115

fireline 115, 117

fireplow 84

fireweed 264, 427

fixed-wing aircraft 78

flame thrower 85

flax, Lewis 132, 135, 138, 142, 196, 264, 271, 273,

275, 277, 279–280, 282, 284, 286–287, 294, 

425–427, 443–444, 702, 727–728, 741

Appar 444, 724

fleabane, Bear River 259

fleabane, Oregon 259

fluted shaft seeder 81

foliage application 92–93

forage production 60

forbs 132, 135, 138, 142, 182–183, 185, 259, 702,

724, 727, 731, 741, 763

forestiera, New Mexico 602, 706

fox 178

foxtail, creeping 259, 289, 308, 310, 346–349, 700

Garrison 300, 349

foxtail, meadow 206, 259, 261–262, 264–265, 269,

289, 308, 310, 346–349, 700, 724

Dan 300, 349

foxtail, reed See foxtail, creeping

Fraxinus anomala See ash, singleleaf

G

galleta 183–184, 250, 279, 293, 308, 311, 390–392

Viva 300, 391

geranium 132, 741

geranium, Richardson 426–427, 438, 702, 724

Geranium richardsonii See geranium, Richardson

geranium, sticky 259, 261, 269, 289, 426, 427, 439,

702

Geranium viscosissimum See geranium, sticky

germination 601

giant hyssop, nettleleaf 427

globemallow 132, 135, 196

globemallow, gooseberryleaf 142, 182, 273, 275, 279,

280, 284–285, 287, 292–293, 427, 461–462, 702,

727, 741

globemallow, scarlet 273, 279, 284–286, 291, 293,

427, 462, 702, 741

ARS 2936 462

globemallow, stream 427

goat's beard See salsify, vegetable-oyster

goldeneye, Nevada 271, 273, 275, 280, 284, 427, 466,

703

goldeneye, showy 132, 135, 142, 196, 259, 269, 271,

275, 277, 282, 287, 289, 427, 465, 466, 703, 724,

727, 731, 741

goldenrod, Canada 269, 271, 289, 427, 460–461, 703,

763

goldenrod, low 259, 269, 427

goldenrod, Parry 277, 427

goldenweed 534, 706

gouger 84
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-7

grama, black 308, 355–356

Nogal 300, 356

Sonora 356

grama, blue 308, 310, 356–358

Hachita 300, 358

Lovington 300, 358

grama, sideoats 308, 310, 353–355

Butte 300

El Reno 300

Haskell 300

Killdeer 300, 354

Niner 300, 354

Pierre 300, 355

Premier 300

Trailway 300

Vaughn 300, 355

gravity separators 715

Grayia brandegei See hopsage, spineless

Grayia spinosa See hopsage, spiny

grazing 2, 3, 128–129, 601

plant control 59

grazing management 194–198

greasewood 470

greasewood, Bailey 469, 490, 706

greasewood, black 135, 195, 247, 262, 266, 267, 292,

469, 472, 490, 491, 602, 706, 725, 741, 750

greasewood, black communities 9, 249–250, 292,

308–309, 601

Great Basin Experimental Range 16

Great Basin Station 3, 4, 16–17

greenhouse bioassay 53

grid ball 92

grinder-macerator 715

ground broadcast seeding 76, 205

groundsel, butterweed 259, 264, 270–271, 289, 427,

459–460, 702

grouse 178–179

blue 170

Columbian sharp-tail 168–169

ruffed 169–171

sage See sage-grouse

growth rate 128

Gutierrezia See matchbrush; snakeweed

Gutierrezia microcephala See matchbrush, small

headed

Gutierrezia sarothrae See snakeweed, broom

H

habitat improvement projects 156

hackberry, netleaf 602, 696–698

hairgrass, tufted 259, 261, 264–265, 289, 308, 310,

369–371

Norcoast 301, 371

Nortran 301, 371

Peru Creek 301, 371

hammermill 713

Hansen seed dribbler 79

Haplopappus See goldenweed

Haplopappus bloomeri See rabbitbrush, goldenweed

harvester

sprigger 763

harvesting, seed 712

hawthorn, Columbia 561

hawthorn, Douglas 266–267, 560–561, 602, 706, 750

hawthorn, river See hawthorn, Douglas

heat damage 103

Hedysarum boreale See sweetvetch, Utah

helianthella, oneflower 182–183, 185, 259, 270, 277,

426–427, 441, 703, 724

Helianthella uniflora See helianthella, oneflower

helicopter 78–79

helitorch 85

Heracleum lanatum See cowparsnip

herbicide effects 90–91, 96

herbicide sprayer 85–87

herbicide types 94, 97

acid equivalent 97

nonselective 94

selective 94

soil sterilant 94

toxicant 97

herbicides 89, 95, 167, 255

application 92, 98

broadcast ground application 93

broadcast spray application 92

cut stump treatment 92

foilage 92

foilage spray 93

grid ball 92

rate, calculation 97–98

safety considerations 99

soil application 92

soil injection 92

soil surface placement 92

stem 92

timing of application 98

trunk base spray 92

trunk injection 92

wipe on 92, 93

common name

clopyralid 95

dicamba 95

glyphosate 95

Oust 95

paraquat 95

picloram 95

tebuthiuron 95

triclopyr 95

type 95

uses 90
Index-8 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Hesperostipa comata See needle-and-thread

Hilaria jamesii See galleta

hoedad 764

Holodiscus discolor See oceanspray, creambush

Holodiscus dumosus See spiraea, rock

honeylocust 602, 706

honeysuckle 731

honeysuckle, bearberry 602, 623–624

honeysuckle, orange 602, 623

honeysuckle, Tatarian 262, 266–267, 272, 706, 750

honeysuckle, Utah 262, 272, 602, 623–624, 706, 750

honeysuckle, western trumpet See honeysuckle,

orange

hopsage, spineless 273, 291, 293, 469, 472, 487, 488,

602, 706, 727

hopsage, spiny 279, 281, 284, 291, 293, 469, 487,

488, 602, 706, 725, 750

Horizon drill 75

horsebrush 103, 534–536

horsebrush, cottonthorn 706

horsebrush, gray 495, 535, 706

horsebrush, littleleaf 495, 706

horsebrush, Nuttall 495, 706

horsebrush, spiny 495

horses 176, 178, 179

human activities 194, 198

hydroseeder 82

I

Indian apple 272, 276, 278, 283, 566–567, 604, 711,

727, 731, 741

inkbush See iodine bush

Intermountain Forest and Range Experiment Station 3,

16 See also Intermountain Research Station

Intermountain Research Station 4, 5, 17

interseeder 81

fluted shaft seeder 81

Truax single row seeder 81

interseeding 142, 742–744

iodine bush 469, 491

iris, German 427, 741

iron 51, 52, 54

J

jointfir See ephedra, green; ephedra, Nevada

junegrass, prairie 102, 271, 275, 277, 282, 287, 308,

311, 392–393, 700

juniper 108

juniper, creeping 602, 633–635

juniper, Rocky Mountain 602, 638–640, 706, 750

juniper, Utah 182–183, 185, 602, 635–638, 638, 707,

750

juniper woodlands 38

juniper-pinyon See pinyon-juniper

Juniperus horizontalis See juniper, creeping

Juniperus osteosperma See juniper, Utah

Juniperus scopulorum See juniper, Rocky Mountain

K

kinnikinnik See manzanita, bearberry

Kochia americana See molly, gray

kochia, Belvedere 703

kochia, forage 135, 138, 142, 196, 273, 276, 278, 279,

281, 283, 284, 285, 292, 293, 294, 469, 472, 489,

490, 603, 707, 730, 741

Immigrant 490, 725

Kochia prostrata See kochia, forage

Koch’s postulates 188

Koeleria cristata See junegrass, prairie

Koeleria macrantha See junegrass, prairie

Koeleria nitida See junegrass, prairie

L

land imprinter 67, 72

larkspur, tall 204

layering 762

crown divisions 762

runners and stolons 762

shoots 762

stem layers 762

suckers 762

layers 762

legumes 136, 137

Lepidospartum latisquamatum See scalebroom

Leymus angustus See wildrye, Altai

Leymus cinereus See wildrye, Great Basin

Leymus racemosus See wildrye, mammoth

Leymus salinus See wildrye, Salina

Leymus simplex See wildrye, alkali

Leymus triticoides See wildrye, creeping

ligusticum, Porter 259, 270, 426–427, 442–443, 703,

727

Ligusticum porteri See ligusticum, Porter

lilac, common 707, 750

Linum perenne lewisii See flax, Lewis

livestock grazing See grazing; grazing management

locust, black 603, 707, 750

Lomatium kingii See lomatium, Nuttall

lomatium, narrowleaf See lomatium, nineleaf

lomatium, nineleaf 426, 428, 444–445, 703, 724, 728

lomatium, Nuttall 259, 270–271, 277, 426, 428, 

445–446, 703, 727

Lomatium triternatum See lomatium, nineleaf

Lonicera ciliosa See honeysuckle, orange

Lonicera involucrata See honeysuckle, bearberry

Lonicera utahensis See honeysuckle, Utah

lupine 135, 196, 741

lupine, mountain 142, 259, 270, 282, 289, 703, 724,

727, 728

lupine, Nevada 271, 277, 280, 284, 428
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-9

lupine, silky 260, 270–271, 277, 282, 287, 289, 426,

428, 446–447, 703, 724, 727–728

lupine, silvery 428, 448, 703

Lupinus argenteus See lupine, silvery

Lupinus sericeus See lupine, silky

M

MacLeod 764, 765

magnesium 51–52, 54

Mahonia aquifolium See barberry, shining

mahonia, creeping See Oregon grape

Mahonia fremontii See barberry, Fremont

Mahonia repens See Oregon grape

management 28, 32

considerations 26

climate 29

impacts on adjacent areas 31

resource values 31

soil condition 27–28, 30–31

strategy 28

vegetation condition 26–27

manganese 51–52, 54

manzanita, bearberry 601, 647–648, 707, 750

maple 195

maple, bigtooth 270, 603, 750

maple, Rocky Mountain 216, 272, 278, 600–608, 707,

750

matchbrush 493, 534

matchbrush, small head 495

McSweeney-McNary Forest Research Act of 1928 3

meadowrue, Fendler 260

mechanical plant control 62

brushland plow 66

disk plow 66

disk-chain 66

moldboard plow 66

off-set disk 66

personnel 78

Medicago falcata See alfalfa, sicklepod

Medicago sativa See alfalfa

medick, black 261–262, 264, 270, 428

medusahead 254–255

medusahead communities 253–256, 294

Melilotus officinalis See sweetclover, yellow

Mertensia arizonica See bluebell, tall

milkvetch, Canadian 703

milkvetch, cicer 132, 135, 138, 142, 196, 261, 270,

271, 277, 282, 289, 425–426, 428, 433–434, 703,

724, 728, 731, 741, 763

Lutana 434

milkvetch, Snake River 428

milkvetch, tall 428

mixed seedings 130–134

mockorange, Lewis 687–689, 707

Waterton 689

mockorange, littleleaf 689

moldboard plow 66

molly, gray 469, 488, 603

molybdenum 51–52, 54

moose

Shiras 164

moth, sagebrush defoliator 514

mountain ash, American 707, 750

mountain ash, Greene’s 266–267, 270, 272, 592–594,

601, 603

mountain ash, Sitka 594

mountain brush communities 9, 195, 216, 217, 219,

271, 308–309, 601

mountain lover 222, 266–267, 603, 707

mountain mahogany, birchleaf 138, 544–545, 707, 750

mountain mahogany, curlleaf 103, 132, 135, 142–143,

182, 183, 185, 196, 217, 272, 276, 278, 283, 

545–550, 603, 707, 725, 727, 730, 741, 750

mountain mahogany, Intermountain 546

mountain mahogany, Intermountain curlleaf 549

mountain mahogany, littleleaf 274, 276, 545, 549, 603,

707, 751

mountain mahogany, true 132, 135, 138, 142–143,

185, 196, 272, 276, 278, 283, 549, 550–552, 603,

707, 725, 727, 730, 741, 751

Montane 552

muhly, spike 731

El Vado 301

mule deer 158, 163, 178

ranges 162

multiple use 131

muttongrass 273, 275, 282, 286–287 See also bluegrass, mutton

muttongrass, longtongue 311

N

Nassella viridula See needlegrass, green

natural spread 134

needle-and-thread 182–184, 220, 228, 237, 243, 256,

273, 275, 277, 279, 280, 282, 284–286, 293–294,

308, 311, 418–419, 700, 731

needlegrass 102, 139

needlegrass, Columbia See needlegrass, subalpine

needlegrass, green 135, 142, 269, 271, 275, 277, 282,

287, 289, 308, 311, 422–424, 700

Cucharas 424

Lodorm 301, 424

needlegrass, Letterman 269, 282, 289, 308, 311, 

419–421, 700

needlegrass, subalpine 259, 269, 308, 311, 416–418,

420

varieties 418

needlegrass, Thurber 280, 284, 287, 294, 308, 311,

421–422

ninebark, dwarf 568
Index-10 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

ninebark, mallow 266–267, 567–569, 603, 707, 751

ninebark, mountain 568

ninebark, Pacific 568

nitrogen 51, 52, 54

nitrogen-free extract 177, 180

no-till drill 75

nurseries and facilities 747

nutrient content 112

nutrient requirements of range animals 176

calcium 178

dry material 176

minerals 178

phosphorus 178

protein 177, 178

vitamins 179

nutrient values of range plants 180–184

O

oak, Gambel 103, 135, 183, 185, 195, 217, 218, 603,

648–650, 707, 741, 751

oatgrass, tall 135, 206, 259, 261, 269, 271, 282, 287,

289, 308, 310, 351–353, 701

Tualatin 353, 354

oceanspray, bush 565 See also oceanspray, spiraea

oceanspray, creambush 563–565, 603

Office of Grazing Studies 3

olive, autumn 751

oniongrass 259, 308, 311, 701

Onobrychis viciaefolia See sainfoin

orchardgrass 132, 135, 142, 196, 205, 206, 216, 220,

255, 259, 264–265, 269, 271, 273, 275, 277, 280,

282, 287, 289, 308, 310, 367–369, 701, 724, 728

Berber 368

Latar 301, 368

Paiute 138, 301, 368, 724, 728

Potomac 301

Oregon grape 270, 614–616, 731 See also barberry,

creeping

Oryzopsis hymenoides See ricegrass, Indian

Osmorhiza occidentalis See sweetanise

P

painted-cup 428

painted-cup, sulphur 260

Pascopyrum smithii See wheatgrass, western

pathogen-free planting stock 189

pathogens, plant 187–190

peachbrush, Anderson 707, 751 See almond, desert

peachbrush, desert 274, 276, 284, 291, 293, 575–577,

603, 707, 725

pearly everlasting 763

peashrub, Siberian 731, 751

peavine, flat 428

peavine, perennial 428

peavine, thickleaf 270, 428

peavine, Utah 270, 428

penstemon, bush 272, 603, 694–696, 707, 751

Penstemon cyananthus See penstemon, Wasatch

penstemon, Eaton 139, 271, 277, 280, 282, 287, 289,

428, 456, 703, 731

Richfield 457

Penstemon eatonii See penstemon, Eaton

penstemon, firecracker 456

Penstemon fruticosus See penstemon, bush

Penstemon humilis See penstemon, low

penstemon, littlecup 428

penstemon, low 206, 271, 280, 282, 289, 428, 457,

703, 724, 763

Penstemon pachyphyllus See penstemon, thickleaf

penstemon, Palmer 132, 135, 138, 142, 196, 271, 273,

275, 277, 279–280, 282, 284–287, 293–294, 425,

428, 454–456, 703, 727–728, 731

Cedar 456

Penstemon palmeri See penstemon, Palmer

penstemon, Rocky Mountain 132, 135, 260, 270–271,

277, 283, 289, 425, 428, 457

penstemon, Rydberg 260, 270, 428, 457

Penstemon rydbergii See penstemon, Rydberg

penstemon, sidehill 428

Penstemon strictus See penstemon, Rocky Mountain

penstemon, thickleaf 277, 428, 457, 703, 731

penstemon, toadflax 277, 428

penstemon, Wasatch 260, 270–271, 277, 283, 289,

428, 456, 703

Peraphyllum ramosissimum See Indian apple

persistence

species 129

personnel 63

Phalaris arundinacea See canarygrass, reed

pheasant 159, 178–179

Philadelphus lewisii See mockorange, Lewis

Phleum alpinum See timothy, alpine

Phleum pratense See timothy

phosphate, single super 54

phosphate, trebel super 54

phosphoric acid (liquid) 54

phosphorus 51, 52, 54, 178, 179

Physocarpus malvaceus See ninebark, mallow

pine, ponderosa 195

pine, ponderosa communities 222–223, 309, 601

ping-pong ball injector 85

pinyon 108

pinyon-juniper 195, 224, 225, 279

pinyon-juniper communities 9, 37, 223–224, 227–228,

273, 279, 308, 601, 638

pipe harrow 67, 71

plant communities, diverse 130

plant community indicators 124

plant competition 21, 57, 61, 112, 134, 202, 207–208,

211, 214, 218, 221, 226, 235, 238, 239–240, 243,

245, 247, 249, 251–252, 255, 257–258
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-11

controlling 57, 65, 147–148

biological 58, 59

burning 60

chemical 62

fallowing 62

grazing 59

herbicides 58

logging 60

mechanical 58, 62, 65

objectives 60

species 21

successional changes 59–60

plant control

herbicides 89

plant disease

fungal diseases 516

plant tissue analysis 53

planting bar 764–767

planting hoe 764–765

planting mixtures, advantages 130–133

planting season 23, 150–151

planting shovels 764–765

planting stock 745–746

plows

brushland 66–67

disk 66

moldboard 66

root 67, 73

PLS See pure live-seed (PLS)

plum, American 262, 571–572, 603, 708, 751

plume, Apache See Apache plume

Poa alpina See bluegrass, alpine

Poa ampla See bluegrass, big

Poa canbyi See bluegrass, Sandberg

Poa compressa See bluegrass, Canada

Poa fendleriana See bluegrass, mutton

Poa longiligula See bluegrass, mutton

Poa nevadensis See bluegrass, Sandberg

Poa pratensis See bluegrass, Kentucky

Poa sandbergii See bluegrass, Sandberg

Poa scabrella See bluegrass, Sandberg

Poa secunda See bluegrass, big; bluegrass, Sandberg

ponderosa pine communities 220, 271, 308

poplar 751

postfire management 119

potash 54

potassium 51–52, 54

Potentilla fruticosa See cinquefoil, shrubby

precipitation 21, 34–35

average annual 34

pronghorn antelope 111, 164–165

propagation See seed propagation; vegetative

propagation

protein 177

proximal analysis 180

Prunus americana See plum, American

Prunus andersonii See almond, desert

Prunus besseyi See cherry, Bessey

Prunus emarginata See cherry, bitter

Prunus fasciculata See peachbrush, desert

Prunus spinosa See blackthorn

Prunus tomentosa See cherry, Nanking

Prunus virginiana See chokecherry

Psathyrostachys juncea See wildrye, Russian

Pseudoroegneria spicata See wheatgrass, bluebunch

Pseudoroegneria spicata inerme See wheatgrass,

beardless

pure live-seed (PLS) 22, 139, 736

Purshia glandulosa See bitterbrush, desert

Purshia tridentata See bitterbrush, antelope

Q

quackgrass 310

quail 178, 179

quailbush See saltbush, quailbush

Quercus gambelii See oak, Gambel

R

rabbit 178

rabbitbrush 103, 493–496, 522–524

rabbitbrush, alkali 495, 533–534, 708

rabbitbrush, Douglas 603

rabbitbrush, dwarf 495, 524, 603, 708

rabbitbrush, goldenweed 534

rabbitbrush, Greene’s 495, 533–534, 708

rabbitbrush, low 135, 182–183, 185, 196, 260, 279,

285–286, 291, 293–294, 495, 531–533, 603, 708

hairy low 533

mountain low 142, 288, 290, 532, 708, 725, 741

narrowleaf low 533, 708

stickyleaf low 532–533, 708

rabbitbrush, Nevada 530

rabbitbrush, Parry 260, 272, 495, 529–531, 603, 708

rabbitbrush, rubber 132, 135, 182–183, 185, 196, 251,

262, 495, 525–529, 603, 751

green rubber 529, 708

green-stemmed rubber 526, 741

leafless rubber 708

leiospermus rubber 708

mountain rubber 270, 272, 276, 278, 283, 288, 529,

708, 725

mountain whitestem rubber 708

threadleaf rubber 266–267, 528–529, 708, 741

tubinatus rubber 708

whitestem mountain and basin rubber 270, 272,

274, 276, 278, 281, 283–284, 286, 288

whitestem rubber 138, 142–143, 525–526, 528, 725,

727, 730, 741

rabbitbrush, small 603

rabbitbrush, spreading 291, 495, 524, 603, 708, 741

rabbitbrush, Vasey 495, 531, 709
Index-12 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

rails 71–72

range, season of use 182–184

Range Seeding Equipment Committee 17

raspberry, black 590, 601

redtop See bentgrass, redtop

reedgrass, bluejoint 264–265, 310

Sourdough 301

reedgrass, chee 259, 264–265, 310, 701, 763

reedgrass, plains 102

rehabilitation 200 See also site preparation and

seeding prescriptions

resistance, host 190

resource values 31

restoration 19, 200 See also revegetation; site preparation and seeding prescriptions

history 1, 15–17

human activities 198

restored sites

grazing 194

human activities 194

management 193

post-treatment 194

revegetation 19, 29, 200, 470, 717, 746 See also

restoration; site preparation and seeding

prescriptions

artificial 29

considerations 19

cost 32

impacts on adjacent areas 31–32

plant competition 21

plant material availability 32

planting season 23

precipitation 21

pure live seed (PLS) 22

resource values 31

seed mixture 22

seedbed 23

site-adapted species and populations 21

soil conditions 27–28, 30

soil types 20

terrain 20

vegetation condition 26–27

weeds 31

history 15–17

human activities 198

impact on resources 28

range 19

site suitability 29–30

wildland 19

Rhizobium 146

rhizomes 763

Rhus aromatica See sumac, skunkbush

Rhus glabra See sumac, Rocky Mountain smooth

Ribes aureum See currant, golden

Ribes cereum See currant, wax

Ribes viscosissimum See currant, sticky

ricegrass, Indian 102, 132, 135, 138, 142, 182, 184,

196, 273, 275, 277, 279, 280, 282, 284, 285, 

286–287, 291, 293–294, 308, 311, 393–396, 701,

731

Nezpar 301, 396

Paloma 301, 397

Ribstone 397

Rimrock 301, 397

riparian communities 9, 209–210, 212–213, 263–267,

309, 601, 668

roads 172, 198

rockspirea 565–566, 709

Rocky Mountain Forest and Range Experiment Station

3 See also Rocky Mountain Research Station

Rocky Mountain Research Station 5

rodents 144

roller chopper 83

root cuttings 762

root-plow 67, 73

Rosa woodsii ultramontana See rose, Woods

rosaceous shrubs 539

rose, Woods 143, 260, 262, 266–267, 270, 272, 278,

283, 288, 586–590, 603, 709, 741, 751

Roundup 96

row spacing 153

Rubus leucodermis See raspberry, black

Rubus parviflorus See thimbleberry

runners and stolons 762

rush, Baltic 263

rush, Drummond 263

rush, longstyle 263

rush, swordleaf 263

rush, Torrey 263

Russian olive 709

rye, mountain 132, 135, 142, 196, 269, 273, 275, 277,

280, 282, 284, 286–287, 294, 309, 311, 724

rye, winter 273, 275, 277, 727

ryegrass, Italian 311

ryegrass, meadow 311

ryegrass, perennial 264, 265, 311

S

sacaton, alkali 196, 262, 264–265, 279, 286, 309, 311,

413–414, 701

Salado 301, 414

Saltalk 301, 414

sage, birdsfoot 495, 522

sage, fringed 274, 495, 505–506, 604, 709, 725

sage, longleaf 495, 522, 709

sage, Louisiana 260–261, 264, 270–271, 278, 289,

426, 428, 430–431, 703, 741, 763

Summit 431

sage, oldman See sandsage
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-13

sage, purple 709, 751

sage, sand See sandsage

sage, tarragon 278, 428

sage-grouse 159, 166–168

sagebrush 108, 236, 493, 494–496

sagebrush communities 115, 234–237 See also site

preparation and seeding prescriptions

sagebrush, alkali 495, 506–507

sagebrush, alkali communities 245

sagebrush, big 132, 181–183, 185, 196, 232, 294,

495, 514–521, 709, 751

big communities 9

basin big 135, 138, 142–143, 195, 238, 266–267,

274, 276, 279, 281, 288, 292, 514–521, 604, 709,

725, 727, 730, 741

Hobble Creek 518

basin big communities 135, 238, 279, 280, 308–309,

601

foothills big 272, 274, 276, 278, 279, 281, 283–284

See also sagebrush, big: xeric big

mountain big 135, 138–139, 142, 195, 232, 260,

266–267, 270, 272, 276, 278, 283, 288,

514–521, 604, 709, 725, 730, 741

Hobble Creek 518

mountain big communities 239–240, 282, 308–309,

601

spicate big 503, 514 See also sagebrush, big:

subalpine big

subalpine big 206, 290

subalpine big communities 289, 308–309

timberline big 244, 260, 289, 290, 520, 709 See also

sagebrush, big: spicate big; sagebrush, big:

subalpine big

timberline big communities 289

Wyoming big 135, 138, 142, 195, 250, 251, 274,

279, 281, 286, 288, 291, 514–521, 604, 725, 730,

741

Gordon Creek 519

Wyoming big communities 240–242, 279, 284,

308–309, 601

xeric big 514, 517 See also sagebrush, big: foothills

big

sagebrush, Bigelow 495, 500–501, 604, 709

sagebrush, black 132, 142, 143, 185, 195, 196, 228,

242, 274, 276, 285, 286, 291, 293, 495, 508–509,

604, 709, 725, 727, 730, 741, 751

black communities 243, 285

sagebrush, budsage 604, 709 See also budsage

budsage communities 245

sagebrush, coaltown 503, 522

sagebrush, early 604 See sagebrush, alkali

sagebrush, fringed See sage, fringed

sagebrush, hotsprings 500

sagebrush, longleaf See sage, longleaf

sagebrush, Louisiana See sage, Louisiana

sagebrush, low 206, 232, 279, 281, 283, 285, 495,

499–500, 506, 604, 709

low communities 244, 286

sagebrush, pygmy 495, 509–510, 604, 709

sagebrush, Rothrock 520, 522

sagebrush, sand 293, 709 See also sandsage

sagebrush, scabland See sagebrush, stiff

sagebrush, silver 232, 260, 266–267, 272, 283, 290,

495, 502–503, 604, 709, 741

Bolander silver 503

mountain silver 502, 503

mountail silver communities 244

plains silver 503

silver communities 289

sagebrush, stiff 495, 510–511, 604, 709

sagebrush, tarragon See sage, tarragon

sagebrush, threetip 232, 288, 495, 521–522, 604

tall threetip 266–267, 522, 709

threetip communities 244, 287

Wyoming threetip 522

sainfoin 132, 135, 138, 196, 261, 270, 272, 275, 278,

283, 289, 453, 703, 724, 741

Eski 454

Melrose 454

Nova 454

Onar 454

Remont 454

Runemex 454

sainfoin, common 142, 428

Salix See willow

Salix bebbiana See willow, Bebb

Salix boothii See willow, Booth

Salix drummondiana See willow, Drummond

Salix exigua See willow, coyote

Salix geyeriana See willow, Geyer

Salix glauca See willow, grayleaf

Salix lasiandra caudata See willow, whiplash

Salix lasiolepis See willow, arroyo

Salix lutea See willow, yellow

Salix planifolia See willow, plainleaf

Salix scouleriana See willow, Scouler

Salix wolfii See willow, Wolf

salsify, vegetable-oyster 428, 703, 724, 727, 741

salt desert shrub communities 246 See also site

preparation and seeding prescriptions

saltbush 469

saltbush, allscale 469, 482–483

saltbush, Australian 469

saltbush, big 469, 710

saltbush, Bonneville 469, 710

saltbush, broadscale 469, 472, 482–483, 710

saltbush, Castle Valley clover 292, 469, 472, 480–481,

604, 710

saltbush, cattle 469, 710

saltbush, cuneate See saltbush, Castle Valley clover
Index-14 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

saltbush, desert holly 469, 482–483, 710

saltbush, falcate 469, 710

saltbush, fourwing 129, 132, 135, 138, 142, 143, 182,

183, 185, 196, 251, 262, 266, 267, 274, 276, 278,

279, 281, 283, 284, 286, 291, 292, 293, 469, 471–

476, 604, 710, 725, 727, 730, 741, 751

fourwing communities 251–252, 279

Marana 475, 476

Rincon 475, 476

Wytana 475, 476

saltbush, Gardner 209, 251, 262, 266–267, 279, 291–292,

469–470, 481–482, 604, 710, 725, 752

saltbush, Gardner complex 470, 481–482

saltbush, Garrett 710

saltbush, mat 291, 469, 472, 479–480, 604, 710

saltbush, Navajo 469, 710

saltbush, quailbush 469, 482–483, 604

Casa 483

saltbush, robust 469

saltbush, shadscale 308, 604, 710, 752

shadscale communities 9, 248, 309, 601

saltbush, trident 469, 472, 710

saltbush, wingless 469

saltgrass 199, 208–209, 212, 262, 264–265

saltgrass, desert 763

saltgrass, inland 195, 310, 371–373

inland communities 9, 208–209, 262, 308-309, 601

Sambucus cerulea See elderberry, blue

Sambucus racemosa See elderberry, red See elderberry, red

sand sage See sandsage

sandsage 495, 504–505, 604

Sanguisorba minor See burnet, small

Sarcobatus vermiculatus See greasewood, black

scalebroom 534, 537

scalper 84

sedge, analogue 263

sedge, beaked 263

sedge, black alpine 263

sedge, blackroot 263

sedge, Douglas 263

sedge, downy 263

sedge, golden 263

sedge, Hepburn 263

sedge, hood 263

sedge, Kellogg 263

sedge, Nebraska 263

sedge, ovalhead 259, 261, 289

sedge, rock 263

sedge, russet 263

sedge, slim 263

sedge, smallwing 263

sedge, soft-leaved 263

sedge, valley 263

sedge, woolly 263

sedges 208, 263

seed

afterripening 700, 726

certification 139, 719

certified 137–138

cleaning 699, 713–715

collecting 699, 712, 718, 754

dormancy 127, 726, 730

field 721, 754

germination 139, 205, 723, 724, 727–728, 730–732,

746

inoculation 146

longevity 731

maturity 700

noncertified 138

orchard 718–719, 722, 754

pathogens 146–147

pelleting 76, 146

pretreatment 145–146, 755

production 719

propagation 747–748

purity 139, 700

quality 137

storage 700, 715, 731, 754–755

stratification 700

seed certification 733 See also certification, seed

seed, certified 22

seed dribbler 79–80

seed harvesters

vacuum-type 713

seed mixture 22

seed, noncertified 138

seed production 601

seed testing

requirements 733

seedbed 23, 126, 134

conditions 63

firmness 149, 150

preparation 121

climatic conditions 124

plant community indicators 124

soil

divalent ions 125

monovalent cations 125

seedbed preparation equipment 65 See also anchor

chains; cables; disks; plows

drags 71

herbicide 91

land imprinter 67, 72

pipe harrow 67, 71

rails 71

seeders 65 See also drills

aerial broadcasting 77–78

fixed-wing aircraft 78

helicopters 78–79

Brillion seeder 80

broadcast 72, 76–77
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-15

hand broadcasting 76

Hansen seed dribbler 79

hydroseeder 82

interseeder

fluted shaft seeder 81

Truax single row seeder 81

land imprinter 72

seed dribbler 79–80

surface seeder 80

surface seeders 80

thimble seeder 79–80

Truax Seed Slinger 76

seeding 216

broadcast 141

depth 76, 152–153

ecotype 124

mixed 201

mixed seedings 130

mixes 122, 129

forbs 140

native species 126

plant competition 21

planting season 23

precipitation 21

pure live seed (PLS) 22

rate 22, 78

seed mixture 22

seedbed 23

site-adapted species and populations 21

soil types 20

seeding practices 121

rates 137, 139–141, 145

requirements 135, 143

separate row seeding 154

single species seeding 130, 133

species compatibility 133, 135

spot 142

seeding rates 259, 261, 269, 271, 273, 279–280, 

284–287, 289, 291–293

seeding requirements

row spacing 153

spot seeding 142

seedling

establishment 133, 134

growth rate 135

vigor 135

selenium 52

Senecio serra See groundsel, butterweed

serviceberry, dwarf Saskatoon 541

serviceberry, Saskatoon 132, 135, 142, 143, 196, 266,

267, 270, 272, 276, 278, 283, 288, 541–543, 604,

710, 725, 727, 741, 752

serviceberry, Utah 272, 274, 276, 541, 543–544, 604,

710, 725, 727, 752

shadscale 132, 135, 143, 195, 196, 228, 243, 245–247,

248, 250–253, 257–258, 279, 291–292, 294, 469,

476–478

sheep 176, 178–179, 515

Shepherdia argentea See buffaloberry, silver

Shepherdia canadensis See buffaloberry, russet

Shepherdia rotundifolia See buffaloberry, roundleaf

Shiras moose 164

shoots 762

shrub enhancement 235

shrub-herb associations 134

Shrubland Biology and Restoration Project 4

shrubs 132, 135, 138, 142, 182–183, 185, 260, 597,

704

silverberry 266–267, 604, 640–641

Sitanion hystrix See squirreltail, bottlebrush

site access 61

site improvement 122

site preparation and seeding prescriptions

aspen communities 215, 309

aspen openings 204–205

aspen-conifer communities 9, 216, 601

Bailey's (1978) 8

blackbrush communities 9, 252–253, 293, 308, 601

brome, red communities 9, 253–256, 294

cheatgrass communities 9, 253–256, 294, 601

greasewood, black communities 9, 249–250, 309,

601

juniper woodlands 38

medusahead communities 253–256, 294

mountain brush communities 9, 219, 309, 601

pine, ponderosa communities 222–223, 309, 601

pinyon-juniper communities 9, 37, 223–224, 227–228,

273, 279, 308, 601, 638

riparian communities 9, 209–210, 212–213, 263–267,

309, 601, 668

sagebrush, alkali communities 245

sagebrush, basin big communities 135, 238, 279,

280, 308–309, 601

sagebrush, big communities 9

sagebrush, mountain big communities 239–240,

282, 308–309, 601

sagebrush, subalpine big communities 289, 308–309

sagebrush, timberline big communities 289

sagebrush, Wyoming big communities 240–242,

279, 284, 308–309, 601

sagebrush, black communities 243, 285

sagebrush, budsage communities 245

sagebrush communities 235–237

sagebrush, low communities 244, 286

sagebrush, silver communities 289

sagebrush, mountain silver communities 244

sagebrush, threetip communities 244, 287

saltbush, fourwing communities 251–252, 279
Index-16 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

saltbush, shadscale 246–247, 291

saltbush, shadscale communities 9, 248, 309, 601

saltgrass, inland communities 9, 208, 209, 262, 308,

309, 601

subalpine communities 601

subalpine herblands 9, 204, 205

weeds, annual communities 308, 601

weeds, annual lowland communities 9, 257, 258

wet and semiwet meadows 9, 168, 207, 208, 308,

601

winterfat communities 251

site-adapted species and populations 21

skunk cabbage 204

snakeweed 534, 536

snakeweed, broom 495, 536

snowberry 103

snowberry, common 266–267, 604, 627, 710, 752

snowberry, desert 627–628

snowberry, longflower 278, 604, 710, 752

snowberry, longleaf 272

snowberry, mountain 143, 206, 260, 266, 267, 270,

272, 276, 278, 283, 288, 290, 604, 629–630, 710,

727, 741, 752

snowberry, western 266, 267, 604, 628–629

snowberry, white See snowberry, common

snowmold 517

snowmold disease 516

soapberry See buffaloberry, russet

softwood cuttings 761–762

soil 40, 125–126

carbon 44

condition 27–28, 30

conditions 125

erosion 28, 40

fertility 40, 52

fertilization 48

improvement 40

moisture 148–149

nitrogen 136

nutrient 40, 47, 48, 49, 50

boron 51–52

calcium 51

concentrations 51

copper 51–52

iron 51–52

magnesium 51–52

manganese 51–52

molybdenum 51–52

nitrogen 50–52

phosphorus 50–52

potassium 51–52

selenium 52

sulfur 50–51

sulphate-sulphur 52

zinc 51–52

pH 45

reclamation potential 40

aspect 47

parent material 47

texture 152

soil application 92

soil carbon 44

soil conditions 36, 39–53, 41

bulk density 41, 42

carbon 44

coarse fragment content 41, 43

depth to limiting layer 41, 44

exchangeable sodium 41, 47

organic matter 41, 44

permeability 41, 43

pH 41, 46

salinity 41, 45, 46

slope 41, 44

structure 41–42

texture 41–42

water-holding capacity 41, 43

soil factors See soil conditions

soil injection 92

soil reaction See soil, pH

soil surface placement 92

soil types 9, 20

Solidago canadensis See goldenrod, Canada

Solomon plume, fat 260, 428

Solomon-seal, western 264

Sorbus scopulina See mountain ash, Greene’s

Sorbus sitchensis See mountain ash, Sitka

species selection

climatic conditions 124–125

cold tolerance 128

compatibility 122, 133

cultivars 129, 137

drought tolerance 127–128

growth rate 135

native species 126–128

palatability 128

persistence 129

seed dormancy 127

soils 125–126

Sphaeralcea coccinea See globemallow, scarlet

Sphaeralcea grossulariifolia See globemallow,

gooseberryleaf

spikerush, common 263

spiraea 594

Spiraea betulifolia See spiraea, bridal wreath

spiraea, birchleaf See spiraea, bridal wreath

spiraea, bridal wreath 594–595, 604

Spiraea densiflora See spiraea, subalpine

spiraea, Douglas 596, 711, 752

Spiraea douglasii See spiraea, Douglas

spiraea, subalpine 596

spiraea, white 594, 595

spirea, rock 266–267, 288, 751 See also rockspirea
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-17

spirea, rock spray 565

Sporobolus airoides See sacaton, alkali

Sporobolus cryptandrus See dropseed, sand

sprigger 763

squirrels 178

squirreltail See squirreltail, bottlebrush

squirreltail, big 411

Sand Hollow 301, 412

squirreltail, bottlebrush 102, 135, 182–184, 196, 264,

265, 271, 273, 275, 277, 279, 280, 282, 284–287,

291–294, 309, 311, 410–412, 701, 724, 728

Fish Creek 412

Toe Jam Creek 412

State Agricultural Experiment Stations 3, 4

steep-slope scarifier and seeder 87

stem application 92

stem cuttings 760

stem layers 762

Stipa columbiana See needlegrass, subalpine

Stipa comata See needle-and-thread

Stipa lettermanii See needlegrass, Letterman

Stipa occidentalis See needlegrass, western

Stipa thurberiana See needlegrass, Thurber

Stipa viridula See needlegrass, green

stork’s bill See alfileria

Suaeda torreyana See desert blite; sumpbush, desert

subalpine communities 601

subalpine herblands 9, 201–202, 204–205, 259

succession, plant 114

successional changes 131

suckers 762

sulfate-sulphur 52

sulphur 51, 54

sumac, Rocky Mountain smooth 272, 276, 278, 283,

604, 611–613, 711, 741, 752

sumac, skunkbush 135, 272, 276, 278, 283, 288, 604,

608–611, 711, 741, 752

Bighorn 611

sumac, smooth 143

summercypress, Belvedere 262

sumpbush, desert 469, 491

sunflower 287

sunflower, annual 701

surface seeder 80

sweetanise 196, 216, 260–261, 270, 289, 426, 428,

454, 704, 724, 728, 731, 741

sweetclover, white 429

sweetclover, yellow 135, 138, 196, 262, 264, 272–273,

275, 278–280, 283–284, 286, 429, 451–452, 704,

724, 728

Goldtop 452

Madrid 452

Yukon 452

sweetroot, spreading 270, 429

sweetvetch, northern See sweetvetch, Utah

sweetvetch, Utah 132, 135, 142, 196, 272, 275, 278,

280, 283, 287, 426, 429, 439–440, 704, 724, 

727–728, 741, 763

Timp 440

Symphoricarpos albus See snowberry, common

Symphoricarpos longiflorus See snowberry, desert

Symphoricarpos occidentalis See snowberry, western

Symphoricarpos oreophilus See snowberry, mountain

syringa See mockorange, Lewis

syringa, Lewis See mockorange, Lewis

T

tarweed, cluster 203

temperature 35

tephritid flies 526

teratorch 85

terrain 20–21, 36, 37

Tetradymia See horsebrush

Tetradymia canescens See horsebrush, gray

Tetradymia glabrata See horsebrush, littleleaf

Tetradymia nuttallii See horsebrush, Nuttall

Tetradymia spinosa See horsebrush, spiny

thimble seeder 79–80

thimbleberry 266–267, 591–592, 601, 605

Thinopyrum intermedium See wheatgrass, intermediate

Thinopyrum ponticum 320 See also wheatgrass, tall

threeawn complex 349

threeawn, fendler 310

threeawn, purple 309–310, 349

threeawn, red See threeawn, purple

timothy 132, 135, 142, 196, 205–206, 208, 215–216,

259, 261, 264–265, 269, 271, 289, 309, 311, 

399–401, 701

Clair 401

Climax 302, 400

Drummond 302

varieties 401

timothy, alpine 259, 269, 289, 309, 311, 398–399, 701,

763

timothy, mountain See timothy, alpine

tissue analysis 53

topography 9, 10, 11, 12, 13

toxicant 97

transplanter 82, 83, 740, 742

transplanting 23, 262, 739

equipment 764, 765

treatment of propagative material 190

treatments

costs 64

economic benefits 64

impacts on resources 64

visual impacts 64

trefoil, birdsfoot 272, 283

trencher 84
Index-18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

Trifolium See clover

Trifolium fragiferum See clover, strawberry

Trifolium hybridum See clover, alsike

Trifolium pratense See clover, red

Trifolium repens See clover, white

trisetum, spike 309, 311

Truax drill 74, 75

Truax Seed Slinger 76

Truax single row seeder 81

turkey 178, 179

Tye drill 75

U

U.S. Department of Agriculture (USDA)

Agricultural Research Service 4

Bureau of Plant Industry 2, 3

Division of Botany 2

Forest Reserves 2

Forest Service 17

Great Basin Experimental Range 16

Great Basin Station 3–4, 16, 17

Intermountain Forest and Range Experiment

Station 3, 16

Intermountain Research Station 4–5, 17

Office of Grazing Studies 3

Rocky Mountain Research Station 5

Shrubland Biology and Restoration Project 4

Natural Resources Conservation Service 5, 17

Plant Materials Centers 4

urea 54

Utah Division of Wildlife Resources 4, 17

Utah State Division of Fish and Game 16

V

vacuum-type seed harvesters 713

valerian, edible 260–261, 264, 429

vegetation condition 26

vegetation status 61

vegetation types 7, 9, 11, 13, 34, 36–38, 308, 601 See

also communities, plant

vegetative propagation 748, 759–763

vetch, American 182–183, 260, 270, 429

vetch, bramble 429

Vicia americana See vetch, American

Viguiera miltiflora nevadensis See goldeneye, Nevada

Viguiera multiflora multiflora See goldeneye, showy

violet, Nuttall 260

virginsbower, western 605, 654–656, 711, 752

vitamin A 179

W

water 171–172

water developments 171

watershed 131

weed control 62–63, 131, 147

weeds, annual communities 308, 601

weeds, annual lowland communities 9, 257–258

weeds, noxious 211

wet and semiwet meadow communities 9, 168,

206–208, 261, 308, 601

wet meadows 262

wheatgrass, beardless 310, 701 See also wheatgrass,

bluebunch

Whitmar 302

wheatgrass, bluebunch 132, 135, 139, 142, 182–184,

196, 271, 273, 275, 277, 280, 282, 284, 286–287,

294, 309–310, 337–342, 724, 728, 763

Anatone 302, 341

Goldar 341

Goldlar 302

NewHy 302, 341

P-7 302, 341

Whitmar 342

wheatgrass, bluebunch bearded 310, 701

wheatgrass, crested 182, 312, 316 See wheatgrass,

fairway crested

wheatgrass, desert 182–184, 196 See wheatgrass,

standard crested

wheatgrass, fairway crested 132, 135, 138, 142, 

183–184, 196, 262, 271, 273, 275, 277, 279–280,

282, 284–287, 293, 294, 297, 297–318, 309, 310,

315, 316, 701, 728

Douglas 302, 316

Ephraim 302, 316–317, 724

Fairway 302, 317

Nordan 316

Parkway 302, 317

Ruff 302, 317

wheatgrass, hybrid

CD-II 316

Hycrest 316–317

NewHy 262

SL-1 342

SL1 302

wheatgrass, hybrid crested

CDII 303

Hycrest 294, 303

Kirk 303

wheatgrass, intermediate 132, 135, 138, 142, 182,

196, 221, 237, 255–256, 259, 269, 271, 273, 275,

277, 279, 280, 282, 284, 286, 287, 293–294, 

309–310, 327–331, 701, 724, 727–728

Amur 303, 330

Chief 303, 330

Clarke 303, 330

Greenar 330

Greenbar 303

Greenleaf 304, 331

Luna 304, 331, 724

Oahe 303, 330

Ree 330
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004 Index-19

Reliant 303, 330

Rush 303, 330

Slate 303, 330

Tegmar 303, 331

Topar 304, 331

wheatgrass, pubescent 309 See also wheatgrass,

intermediate

wheatgrass, Siberian crested 142, 196, 246, 273, 275,

279–280, 284, 286–287, 291–293, 309–310, 314,

332–334, 701

P27 304

Vavilov 304

wheatgrass, slender 142, 196, 259, 264–265, 269,

271, 277, 282, 287, 289, 309–310, 342, 701

Adanac 304, 344

Primar 304, 344

Pryor 304, 345

Revenue 304, 345

San Luis 304, 345

wheatgrass, Snake River 309, 338

Secar 304, 338, 341, 342

wheatgrass, standard crested 132, 135, 138, 142, 271,

273, 275, 277, 279, 280, 282, 284–287, 291–294,

309–310, 313, 320–324, 724

Nordan 304, 324

Summit 304, 326

wheatgrass, streambank 209, 220, 228, 262, 271, 273,

275, 277, 279–280, 282, 284–287, 291–292, 294,

310, 320, 701, 763 See also wheatgrass,

thickspike

wheatgrass, tall 132, 135, 142, 196, 255, 261, 262,

264, 265, 271, 280, 292, 309, 310, 325–327, 702,

724, 731

Alkar 305, 326

Jose 305, 327

Largo 305, 327

Orbit 305, 327

Platte 305, 327

wheatgrass, thickspike 228, 237, 271, 275, 277, 279,

280, 282, 284, 286–287, 294, 305, 309–310, 

318–320, 702, 763

Bannock 305, 319

Critana 305

Schwendimar 305, 320

Sodar 305

Soday 320

wheatgrass, western 135, 139, 142, 182–184, 220,

228, 242, 255, 264–265, 271, 273, 275, 277, 280,

282, 284–285, 287, 291–294, 309–310, 334–337,

701, 763

Arriba 305, 336

Barton 305, 336

Flintlock 305, 336

Rodan 305, 336

Rosana 305, 336, 337

Walsh 305, 337

wheatgreass, crested 297

whortleberry 711

wilding stock 739–740, 742, 759

wildlife habitat 109–110, 112, 131, 155, 539

cover 110, 131, 157

edges, vegetative 111

security 162

thermal 161

improvement 156

management 113

wildrye, alkali 309, 311, 382 See also wildrye, low

creeping

wildrye, Altai 309–310

Eejay 306

Pearl 306

Prairieland 306

wildrye, beardless 311 See wildrye, creeping

wildrye, blue 269, 309–310, 377–379

Arlington 306, 378–379

wildrye, Canada 309–310, 373–374

Mandan 306, 374

wildrye, creeping 264, 265, 311, 383–384

Rio 384

Shoshone 384

wildrye, Dahurian

Arthur 306

James 306

wildrye, Great Basin 132, 135, 138, 142, 196, 246,

261, 264–265, 275, 277, 279, 280, 282, 284, 287,

292, 309–310, 374–377, 702, 728

Magnar 306, 376–377, 724

Trailhead 306, 377

wildrye, low creeping See wildrye, alkali

wildrye, mammoth 264, 265, 309–310, 702

Volga 306

wildrye, purple 310

wildrye, Russian 132, 135, 138, 142, 196, 262, 264,

273, 275, 279–280, 284–287, 291–293, 309, 310,

379–380, 702, 724, 728

Bozoisky-select 307, 380

Cabree 307, 380

Mankota 307, 380

Mayak 307, 381

Swift 307, 381

Tetracan 307, 381

Vinall 307, 381

wildrye, Salina 262, 291, 309–310, 381–382

wildrye, yellow 310

willow 206, 208, 210–11, 213, 262, 266–268, 668–670

willow, arroyo 268, 680–681

Rogue 681

willow, barrenground 268

willow, Bebb 268, 670–672

Wilson 672

willow, Booth 268, 605, 672–673

willow, coyote 605, 676, 752
Index-20 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-136. 2004

willow, Drummond 268, 605, 673–675

Curlew 674

willow, Geyer 268, 676–677

willow, grayleaf 268, 677–678

willow, Pacific 268

willow, peachleaf 268

willow, plainleaf 683–684

willow, purpleosier 262, 605, 752

willow, sandbar 268 See also willow, coyote

willow, Scouler 268, 605, 684–686, 752

willow, whiplash 605, 679–680

Nehalem 680

Roland 680

willow, Wolf 268, 686–687

willow, yellow 681–683

winterfat 132, 135, 138, 142–143, 182–183, 185, 196,

201, 209, 228, 236–237, 242, 245–247, 250–251,

253, 274, 276, 278–279, 281, 284–285, 291–293,

469, 470, 472, 484–486, 605, 711, 725, 727, 730,

741, 752

winterfat communities 251

Hatch 484, 488

winterfat, foothills 486

winterfat, Pamirian 469, 486, 488

wipe-on applicators 92, 93

wormwood 495 See sage, Louisiana

wormwood, common 498

wormwood, dwarf 498

wormwood, oldman 495, 497–498, 605, 711, 741, 753

wortleberry, big 753

Y

yarrow, western 260–261, 264, 270, 272–273, 275,

279–280, 283, 287, 289, 291, 294, 426, 429, 430,

704, 741, 763

yellowbrush See rabbitbrush

Z

zinc 51–52, 54

zuckia, Arizona 469

Zuckia brandegei See hopsage, spineless
The U.S. Department of Agriculture (USDA) prohibits discrimination in all its

programs and activities on the basis of race, color, national origin, sex, religion,

age, disability, political beliefs, sexual orientation, or marital or family status. (Not

all prohibited bases apply to all programs.) Persons with disabilities who require

alternative means for communication of program information (Braille, large print,

audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-2600 (voice

and TDD).

To file a complaint of discrimination, write USDA, Director, Office of Civil Rights,

Room 326-W, Whitten Building, 1400 Independence Avenue, SW, Washington,

DC 20250-9410 or call (202) 720-5964 (voice or TDD). USDA is an equal

opportunity provider and employer.

The Rocky Mountain Research Station develops scientific information and technology to improve management, protection, and use of

the forests and rangelands. Research is designed to meet the needs

of National Forest managers, Federal and State agencies, public and

private organizations, academic institutions, industry, and individuals.

Studies accelerate solutions to problems involving ecosystems,

range, forests, water, recreation, fire, resource inventory, land reclamation, community sustainability, forest engineering technology,

multiple use economics, wildlife and fish habitat, and forest insects

and diseases. Studies are conducted cooperatively, and applications

may be found worldwide.

Research Locations

Flagstaff, Arizona Reno, Nevada

Fort Collins, Colorado* Albuquerque, New Mexico

Boise, Idaho Rapid City, South Dakota

Moscow, Idaho Logan, Utah

Bozeman, Montana Ogden, Utah

Missoula, Montana Provo, Utah

Lincoln, Nebraska Laramie, Wyoming

*Station Headquarters, Natural Resources Research Center,

2150 Centre Avenue, Building A, Fort Collins, CO 80526

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ROCKY MOUNTAIN RESEARCH STATION
Winterfat seedstalk Photo by Nancy Shaw