SPECIES: Grayia spinosa


Grayia spinosa: INTRODUCTORY

INTRODUCTORY

SPECIES: Grayia spinosa
AUTHORSHIP AND CITATION:

Tirmenstein, D. A. 1999. Grayia spinosa. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].



ABBREVIATION:

GRASPI

SYNONYMS:

Atriplex spinosa (Hook.) Collotzi [23]
Atriplex grayi Collotzi [77]

NRCS PLANT CODE:

GRSP

COMMON NAMES:

spiny hopsage
Gray's saltbush
spiny sage

TAXONOMY:

The scientific name of spiny hopsage is Grayia spinosa (Hook.) Moq. (Chenopodiaceae) [11,22,30,78].

LIFE FORM:

Shrub

FEDERAL LEGAL STATUS:

No special status

OTHER STATUS:

No entry


DISTRIBUTION AND OCCURRENCE

SPECIES: Grayia spinosa
GENERAL DISTRIBUTION:

Spiny hopsage grows east of the Cascade Range and Sierra Nevada from central Washington to southern California, eastward to southwestern Montana and western Colorado [6,23,54]. It is scattered throughout the Great Basin and northern Mojave Desert [8].

ECOSYSTEMS:

FRES21  Ponderosa pine
FRES29  Sagebrush
FRES30  Desert shrub
FRES35  Pinyon-juniper
FRES40  Desert grasslands

STATES:

AZ   CA   CO   ID   MT   NM   NV   OR   UT   WA   WY

BLM PHYSIOGRAPHIC REGIONS:

 5 Columbia Plateau
 6 Upper Basin and Range
 7 Lower Basin and Range
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau

KUCHLER PLANT ASSOCIATIONS:

K017 Black Hills pine forest
K023 Juniper-pinyon woodland
K037 Mountain mahogany-oak shrub
K038 Great Basin sagebrush
K039 Blackbrush
K040 Saltbush-greasewood
K041 Creosote bush
K051 Wheatgrass-bluegrass
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K057 Galleta-threeawn shrubsteppe
K063 Foothills prairie

SAF COVER TYPES:

237 Interior ponderosa pine
239 Pinyon-juniper

SRM (RANGELAND) COVER TYPES:

107 Western juniper/big sagebrush/bluebunch wheatgrass
210 Bitterbrush
211 Creosotebush scrub
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
316 Big sagebrush-rough fescue
320 Black sagebrush-bluebunch wheatgrass
321 Black sagebrush-Idaho fescue
408 Other sagebrush types
414 Salt desert shrub
501 Saltbush-greasewood
506 Creosotebush-bursage
508 Creosotebush-tarbush

HABITAT TYPES AND PLANT COMMUNITIES:

Spiny hopsage is well adapted to sagebrush deserts, juniper-pinyon (Pinus-Juniperus spp.) woodlands, and salt desert shrublands [6,8]. It grows in association with such species as ponderosa pine (P. ponderosa), Utah juniper (J. osteosperma), and singleleaf pinyon (P. monophylla) [37]. Spiny hopsage commonly occurs in fairly pure stands [6] or in large extensive tracts with big sagebrush (Artemisia tridentata) [7]. Spiny hopsage often grows in association with budsage (A. spinescens), black greasewood (Sarcobatus vermiculatus), fourwing saltbush (Atriplex canescens), Gardner's saltbush (A. gardneri), winterfat (Krascheninnikovia lanata), creosotebush (Larrea tridentata), horsebrush (Tetradymia spp.), green rabbitbrush (Chrysothamus viscidiflorus), galleta (Hilaria jamesii), needle-and-thread grass (Hesperostipa comata), and Sandberg bluegrass (Poa secunda) [6,15,80].

Spiny hopsage is dominant in a number of desert shrub communities. It is often codominant with big sagebrush, black sagebrush (Artemisia nova), green rabbitbrush, shadscale (Atriplex confertifolia), and Sandberg bluegrass [2,5,9]. Publications describing spiny hopsage as a dominant or indicator species include:

Presettlement vegetation of part of northwestern Moffat County, Colorado, described from remnants [2]
Vegetation and soils of the Cow Creek Watershed [4]
Vegetation and soils of the Churchill Canyon Watershed [5]
Steppe vegetation of Washington [9]
Vegetation and plant communities of southern California deserts--a functional view [28]
Sagebrush steppe [82]


MANAGEMENT CONSIDERATIONS

SPECIES: Grayia spinosa
IMPORTANCE TO LIVESTOCK AND WILDLIFE:

Spiny hopsage is tolerant of grazing and where abundant is a valuable browse plant for many species of wildlife and livestock [6,9,45,67]. Spiny hopsage provides a palatable and nutritious food source for big game and livestock, particularly during late winter through spring [54]. Its relative forage value may be less during the fall when it has been described as "useless." However, Stubbendieck and others [66] describe spiny hopsage as being browsed by livestock in the fall, winter, and spring. Spiny hopsage is used as forage to at least some extent by domestic sheep and goats, deer, pronghorn, and rabbits [73].

On bighorn sheep ranges in east-central Idaho, Lauer and Peek [32] reported heavy use of spiny hopsage. It receives at least some winter pronghorn use in Utah [61]. Spiny hopsage provides food for the mountain cottontail in south-central Idaho [29]. In northeastern California, spiny hopsage is a major food of the black-tailed jackrabbit [12]. Spiny hopsage seedlings are eaten by a variety of small mammals including Botta's pocket gopher, Merriam's kangaroo rat, southern grasshopper mouse, deer mouse, and desert cottontail [54].

Spiny hopsage provides browse for domestic sheep in parts of Wyoming [36] and is described as a “good” forage species for domestic sheep in Utah. Domestic sheep browse the succulent new growth in late winter and early spring [25,26]. In north-central Wyoming, spiny hopsage occurs with greasewood and winterfat and provides forage for winter livestock [16]. In Utah, spiny hopsage is important in the diet of domestic sheep only during early spring [19]. In the Mojave Desert, domestic sheep browse spiny hopsage late in the growing season [43]. The large quantities of seeds produced are valuable for fattening domestic sheep [66]. Minor damage to livestock may result from the spines [6,67].

Van Dyke and others [72] reported moderate fall use of spiny hopsage by bighorn sheep in southeastern Oregon.

 

PALATABILITY:

Spiny hopsage is considered one of the most palatable of the salt desert shrubs, particularly during the spring. However, overall value is limited in most areas since leaves and fruits are shed by early summer [54]. Early spring growth is very palatable, but by mid-summer palatability is very poor.

Spiny hopsage retains very little edible forage over the fall and winter [45]. However, Wasser [73] reports fair to good palatability for deer and domestic sheep and goats in the late fall, winter, and spring. Palatability to cattle is described as “fair to poor” and palatability to horses is rated as “poor to useless” [7]. The degree of preference shown by browsing animals apparently varies greatly according to ecotypic variability by stand [81]. Thorns may keep spiny hopsage from being utilized [6,67].

The degree of use shown by livestock and wildlife species for spiny hopsage in several western states is rated as follows [10]:

                       CO          OR        UT          WY         
  
Cattle                 Fair        ____       Fair       Fair         
Domestic sheep         Good        ____       Good       Good
Horses                 Fair        ____       Good       Good    
Pronghorn              ____        ____       Fair       Fair 
Bighorn                ____        Fair       ____       ____       
Elk                    ____        ____       Fair       ____  
Mule deer              ____        ____       Fair       Poor
White-tailed deer      Poor        ____       ____       ____
Small mammals          Fair        ____       Good       Good
Small nongame birds    ____        ____       Poor       ____ 
Upland game birds      ____        ____       Fair       ____ 
Waterfowl              ____        ____       Poor       ____
NUTRITIONAL VALUE:

Spiny hopsage is rated "fair" in energy and protein value [10]. In southwestern Idaho, crude protein of foliage averages 18% during late winter and 11.8% in the summer months [54]. Nutritional content of spiny hopsage, expressed as a percentage of dry matter, is as follows [41]:


                                Aerial Part,  Aerial Part,   Pods,   
                                Fresh         Leaves, Fresh  Dormant                       
Ash (%)                          5.1          30.0           23.0      
Crude Fiber (%)                 38.3           9.2           22.2
Ether Extract (%)                2.0           1.7            1.2      
N-Free Extract (%)              45.7          48.3           44.8 
Protein (N × 6.25) (%)           8.9          10.2            8.8 
 cattle: digestible protein (%)  5.5           6.2            5.0 
 cheep: digestible protein (%)   5.3           6.5            5.2
Phosphorus %                     0.19          ---            --- 
The following nutritional values have been reported for spiny hopsage in the northern Mojave Desert [52]:

      N(%)      P(%)      Na(%)     K(%)     Ca(%)     Mg(%)    
leaf  2.23      0.09      0.175     10.13     4.25      2.15      
stem  ----      0.08      0.009      6.06     1.23      0.51   

      Si(mg/g)  Zn(mg/g)  Cu(mg/g)  Fe(mg/g) 
leaf  0.07      37        5         150   
stem  0.01      16        3          20


COVER VALUE:

Spiny hopsage provides good cover for birds, rodents, and lagomorphs, even when dormant [37,54]. Waterfowl, including mallards, use it for nesting cover in eastern Washington [21].

The degree to which spiny hopsage provides environmental protection for wildlife species in two western states is as follows [10]:
                              UT           WY

      Pronghorn               Poor         Fair 
      Bighorn                 ____         ____
      Elk                     Poor         Poor
      Mountain goat           ____         ____
      Mule deer               Poor         Poor
      White-tailed deer       ____         Poor
      Small mammals           Fair         ____
      Small nongame birds     Fair         ____
      Upland game birds       Fair         Fair
      Waterfowl               Poor         ____

VALUE FOR REHABILITATION OF DISTURBED SITES:

Spiny hopsage has moderate potential for erosion control and low to high potential for long-term revegetation projects [10]. It can improve forage, control wind erosion, and increase soil stability on gentle to moderate slopes [27,54]. Spiny hopsage is suitable for highway plantings on dry sites in Nevada [64]. Generalized characteristics for use in rehabilitation are as follows [44]:


        Adaptation attribute            Rating

        establishment by seed           medium  
        establishment by transplant     medium
        seed production & handling      medium  
        natural spread by seed          medium  
        growth rate                     good
        soil stability                  medium
        natural vegetative spread       very poor
        adaptability to disturbance     medium

In Idaho, spiny hopsage seedlings produced from bareroot stock were of marginal quality [59]. Larger stock (size class 2-0 or larger, that is grown 2 years indoors and 0 years outdoors) typically produces better results than 1-0 size stock (grown 1 year indoors, 0 years outdoors) [40]. Spiny hopsage has been successfully transplanted onto mine spoils in Wyoming and Utah [17,35]. The following results were reported after spiny hopsage was transplanted onto replaced topsoil in southeastern Wyoming [35]:

First-year survival of bareroot transplants (soil pH of 7.5-8.0):

             # of individuals    # of individuals    survival    
              transplanted          surviving          (%)          
                   25                  10               40             

First-year survival of front-end loader transplants (soil pH 7.0-8.0):
				  
             
             # of individuals    # of individuals    survival
              transplanted          surviving          (%) 
                   90                  64               71  
       
 
Five years after establishment, spiny hopsage transplanted onto processed oil shale at a Uinta Basin, Utah, site had a 56% survival rate with an average height of 12 inches (30 cm) [17]. Spiny hopsage seedlings have been successfully transplanted onto arid roadsides in the eastern Sierra Nevada of California [63]. Container stock has been successfully used for reclamation of reconstructed soils after mining activities [54].

Spiny hopsage can be seeded directly onto disturbed sites in northern shrub steppe communities [54]. Best results are generally obtained when seeds are planted in late fall or early winter which provides stratification [57,54]. Seed remains viable up to 2 years in storage. Plummer [44] recommends selecting seed by matching an ecotype that grows under conditions similar to those where it is to be planted. According to Plummer [44], "plant characteristics such as growth form, rooting depth, palatability, ease of establishment, and resistance to grazing should be used." Spiny hopsage seed is commonly cleaned by hammermilling to remove appendages from the utricles and then fanned to remove debris [59]. Seedling establishment for broadcast and planted seeds was as follows [79]:



Seedbed                      Seeds with             Seeds with                                
characteristics              entire bracts          bracts removed
                                (%)                     (%)                    
loose surface                    18                       2
smooth surface                    7                       0
packed surface                    4                       0
seeds planted 0.5 cm deep        51                      48
 
Variable results have been reported for spiny hopsage cuttings [54]. Fair success is reported for rooting of stem cuttings treated with 0.3 percent to 0.8 percent IBA powder in late winter and early spring [27,48].

Stark [64] reported seed can be germinated in vermiculite, sand, and rock, and recommended planting in March.

 

OTHER USES AND VALUES:

Some Native American peoples traditionally ground parched seeds of spiny hopsage to make pinole flour [66].

OTHER MANAGEMENT CONSIDERATIONS:

Spiny hopsage readily establishes and increases on overgrazed and denuded ranges [62]. Plummer and others [45] report that in Utah, spiny hopsage has a poor natural rate of spread.

Spiny hopsage is fairly tolerant of browsing [53]. It is however, being replaced over wide areas by less palatable species such as green rabbitbrush as a result of heavy pressure by livestock in the spring and early summer. Blaisdell and Holmgren [6] suggest range management practices such as changing the season of grazing or rotating divided range units in salt-desert shrub rangelands. On ranges in good condition, the maximum recommended winter use of annual growth is 80% [6]. Mean cover of individual spiny hopsage plants decreased 29% in response to heavy domestic sheep grazing in the western Mojave Desert. Total composition and cover averages for spiny hopsage were [76]:

             composition (%)     cover (shrubs/m2)
grazed       17                  0.378
ungrazed     19*                 0.533*
_____________________________________________________
*significant difference between treatments (p<0.05)
Spiny hopsage sprouts readily after roto-beating or light burning and is somewhat resistant to application of 2,4-D. It is easily killed by plowing with a heavy disk [62].

Spiny hopsage accumulates potassium in its leaves to such an extent that decay of the leaf litter may raise the surface soil pH beneath the canopy. These soil changes may affect future growth of spiny hopsage and other shrubs growing in association [50]. However, according to Rickard and Keough [50], "whether these mineral concentrations will tend to make the soil more or less suitable for future generations of shrubs remains to be seen."


BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Grayia spinosa
GENERAL BOTANICAL CHARACTERISTICS:

Spiny hopsage is a woody, erect, diffusely branched native shrub that reaches 1 to 5 feet (0.3-1.5 m) in height [7,23,27]. Branches are more or less spine-like at the tips [23]. Spiny hopsage is evergreen in the southern deserts and deciduous in the summer in the northern deserts [73]. Spiny hopsage is generally dioecious, although in some populations approximately 5% of the individuals are monoecious [7,8]. The fruit is a wing-margined utricle [62]. The root system is shallow and spreading [27].

Spiny hopsage is relatively long-lived [9].

RAUNKIER LIFEFORM:

Phanerophyte

REGENERATION PROCESSES:

Spiny hopsage regenerates vegetatively by sprouting after top-killing disturbances such as fire [9]. Spiny hopsage also regenerates from wind or gravity-dispersed winged seed, which is produced annually [37,46,66]. Seed production is variable. Plummer and others [44] describe seed production as "poor" in Utah whereas Daubenmire [9] reports that spiny hopsage produces an abundance of seed in Washington.

Persistent bracts apparently modify the microenvironment on the surface of the seedbed and aid in germination [83]. Light has little effect on germination of spiny hopsage [62]. However, habitat-correlated variation in germination patterns has been reported [39]. Populations from the northern part of the species’ range require cold temperatures to break dormancy [56]. Seeds readily germinate without pretreatment [79]. However, germination can be improved by cold stratification or dry storage [27,57]. Dormancy in spiny hopsage may represent an adaptation which delays germination until early spring when soil moisture conditions are most favorable [54]. Detailed information on germination and seed characteristics is available [54,58]. Germination patterns reflect spiny hopsage's ability to germinate either late in the fall or early in the spring while moisture is available [79]. Ecotypes apparently vary in germination response to temperature [73]. Seed fill is variable, ranging from 18 to 95% [62]. Spiny hopsage seed is reported to remain viable for 6 years in dry storage [62]. However, Plummer [46] rates the seeds as having good viability for 2 years. Some limited 2nd and 3rd year germination has been reported [57].

Spiny hopsage accumulates seeds in states of primary or even secondary dormancy within the soil in the Mojave Desert. Secondary dormancy occurs when nongerminating seeds enter dormancy in response to environmental conditions. Shaw and others [57] report "determining whether spiny hopsage seeds are capable of entering secondary dormancy and affecting factors will require further investigation." Seedbanks can contribute to periodic episodes of good seedling establishment during moist years. Seedlings require chilling in order for the bud to break and growth to occur [57].

 

SITE CHARACTERISTICS:

Spiny hopsage grows on dry plains, deserts, and foothill slopes throughout its range [37]. It is highly tolerant of drought [27].

Spiny hopsage is fairly tolerant of alkaline and saline soils [27] and is typically found on highly calcareous alkaline soils. Its adaptation to alkalinity is low relative to other salt-desert chenopods such as shadscale and black greasewood [37]. Some plants occur on neutral soils [7]. Spiny hopsage occurs on a wide range of soil textures from gravel, sand, and sandy loams to heavy clay [10,27,66], but it prefers sandy soils that are free of salt and hardpans [6]. Potassium is accumulated in spiny hopsage leaves and may comprise up to 63 to 73% of the total cations present. Spiny hopsage concentrates potassium in the surface soils beneath the canopy through decomposition of leaf litter [50].

Spiny hopsage grows from 524 feet to 9,514 feet (160-2900 m) in elevation [54]. Elevational ranges are as follows [10,64]:

4,000 to 7,000 feet (1219-2134 m) in Colorado
2,500 to 7,500 feet (762-2286 m) in Nevada
2,000 to 5,500 feet (610-1676 m) in Utah
4,100 to 6,800 feet (1,250-2073 m) in Wyoming

SUCCESSIONAL STATUS:

Spiny hopsage grows in a number of undisturbed sagebrush and saltbush desert shrub communities. It grows on disturbed sites such as coppice dunes [33]. According to Webb and others [75], it quickly reasserts dominance over colonizers on debris flows in the Panamint Mountains of California. The authors measured response rates of desert plants to disturbance in Death Valley National Monument. Plant assemblages were measured to quantify recovery rates and models developed to account for vegetative change based on principal component analysis following townsite abandonment, disturbances along a pipeline corridor, and on plant assemblages on new, several thousand-year old, and ancient (> 5,000-year-old) mud/debris flows. The following successional pathway has been proposed for mud/debris flows in Death Valley, California [74]:

1) annuals - filaree (Erodium spp.) and brome (Bromus spp.) colonize
2) lupine (Lupinus spp.)/saltbush (Atriplex spp.)/penstemon (Penstemon spp.)/
       buckwheat (Eriogonum spp.), and rabbitbrush (Chrysothamnus spp.) establish
3) wolfberry (Lycium spp.)/spiny hopsage/Nevada ephedra (Ephedra nevadensis) and blackbrush
4) spiny hopsage and Nevada ephedra dominate and initially colonizing species decline
5) blackbrush increases at the expense of Nevada ephedra, spiny hopsage, and wolfberry (may take more than 5,000 years)
6) blackbrush grows in nearly pure stands (may take 10,000 years to develop)

SEASONAL DEVELOPMENT:

Leaves of spiny hopsage generally appear in late February and early March [50]. In a Nevada study on the Great Basin-Mojave Desert interface, plants first leafed out from 2/2 to 3/20, with a mean date of 2/26 [68]. At a second Nevada site within the Great Basin, average leaf bud break took place on March 19 [54].

Spiny hopsage sheds its leaves much earlier than are those of many other deciduous desert shrubs [34]. In southern Nevada, leaves generally drop between mid-April and mid-May [69]. Leaves are usually shed by mid-July in most locations [6], but there is an ecotype in northern Utah that retains a higher percentage of leaves after the growing season [37].

Flowering is hastened by rainfall [68]. Most of the growth, including flowering and fruiting, is completed by midsummer [6]. In southern Nevada, first dates of flowering ranged from 2/26 to 4/9, with a mean date of 3/17 [68]. General flowering dates for four western states are as follows [1,10]:

  State      Beginning of Flowering      Flowering      End of Flowering
  CO         May                         May            July
  MT         June                        June           June
  NV         February/March              March          April
  UT         April                       May            June    
First dates of fruiting ranged from 3/10 to 4/20, with a mean date of 4/2 [68]. Fruit maturity is generally reached in mid-July, just before leaf fall, but this differs among ecotypes [6,37]. Bud formation, leaf fall, and the onset of summer dormancy usually occur after seed dispersal, but there is wide variation in this characteristic [54]. In central Utah, seeds matured from June 15 to July 17 over the course of a 5-year study [7].

Dormancy in spiny hopsage is one of the longest of desert shrubs [54]. Spiny hopsage remains dormant throughout the hot, dry summer months [50]. In southern Nevada, Ackerman and others [1] found that spiny hopsage became dormant when daytime air temperatures exceeded 104 degrees Fahrenheit (40oC). Summer rains will not cause initiation of new leaf growth [13]. Fall rains and/or periods of low temperatures ranging from 41 degrees Fahrenheit (5 oC) to below 32 degrees Fahrenheit (0 oC) usually break dormancy [1].


FIRE ECOLOGY

SPECIES: Grayia spinosa
FIRE ECOLOGY OR ADAPTATIONS:

Spiny hopsage is considered to be somewhat fire tolerant and often survives fires that kill sagebrush [7,55]. Spiny hopsage often sprouts after fire [9,7]. However, in some instances sprouting does not occur [51,55]. Failure to sprout may be due to stressful conditions brought about by overgrazing and erosion [55].

The range of fire intervals reported for some species that dominate communities where spiny hopsage occurs are listed below. To learn more about the fire regimes in those communities refer to the FEIS summary for those species, under “Fire Ecology or Adaptations.”

ponderosa pine (Pinus ponderosa)  2-42 years
basin big sagebrush (Artemisia tridentata ssp. tridentata)
mountain big sagebrush (A. tridentata ssp. vaseyana)
Wyoming big sagebrush (A. tridentata ssp. wyomingensis)

POSTFIRE REGENERATION STRATEGY:

Small shrub, adventitious-bud root crown
Secondary colonizer - off-site seed


FIRE EFFECTS

SPECIES: Grayia spinosa
IMMEDIATE FIRE EFFECT ON PLANT:

Spiny hopsage is generally top-killed by fire [9]. Plants often survive fires that kill adjacent sagebrush [7]. Mature spiny hopsage generally sprout after being burned [9]. Spiny hopsage is reported to be least susceptible to fire during summer dormancy [51].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:

No entry

PLANT RESPONSE TO FIRE:

Spiny hopsage often sprouts after plants are damaged by fire or mechanical injury [54]. Sprouting may occur when mature plants are burned off at the soil surface. Fires in spiny hopsage sites generally occur in late summer when plants are dormant, and sprouting generally does not occur until the following spring [9]. Holsinger [24] noted that apparently dead spiny hopsage plants resprouted several years after a wildfire in the Dautrich Memorial Desert Preserve near the Snake River in Canyon County, Idaho. These plants were located on the north-facing slopes of a basalt mesa [55].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:

Variation in sprouting among populations and possible relationships of sprouting to season and intensity of burn is possible, but has not yet been documented [55].

FIRE MANAGEMENT CONSIDERATIONS:

Fire may be most damaging to spiny hopsage when plants are actively growing [55]. Mortality of spiny hopsage after a July wildfire in southeastern Washington was attributed to "stressed growing conditions imposed by an alkaline-sodic rooting substrate" [51]. Several decimeters of soil had been lost due to earlier overgrazing and subsequent erosion and only a thin soil layer remained above a deep alkaline-sodic rooting substrate [55].


Grayia spinosa: References


1. Ackerman, T. L.; Romney, E. M.; Wallace, A.; Kinnear, J. E. 1980. Phenology of desert shrubs in southern Nye County, Nevada. In: The Great Basin Naturalist Memoirs No. 4. Nevada desert ecology. Provo, UT: Brigham Young University: 4-23. [3197]

2. Baker, William L.; Kennedy, Susan C. 1985. Presettlement vegetation of part of northwestern Moffat County, Colorado, described from remnants. The Great Basin Naturalist. 45(4): 747-783. [384]

3. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]

4. Blackburn, Wilbert H.; Eckert, Richard E., Jr.; Tueller, Paul T. 1969. Vegetation and soils of the Cow Creek Watershed. R-49. Reno, NV: University of Nevada, Agricultural Experiment Station. 77 p. In cooperation with: U.S. Department of the Interior, Bureau of Land Management. [458]

5. Blackburn, Wilbert H.; Tueller, Paul T.; Eckert, Richard E., Jr. 1969. Vegetation and soils of the Churchill Canyon Watershed. R-45. Reno, NV: University of Nevada, Agricultural Experiment Station. 155 p. In cooperation with: U.S. Department of the Interior, Bureau of Land Management. [460]

6. Blaisdell, James P.; Holmgren, Ralph C. 1984. Managing Intermountain rangelands--salt-desert shrub ranges. Gen. Tech. Rep. INT-163. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 52 p. [464]

7. Blauer, A. Clyde; Plummer, A. Perry; McArthur, E. Durant; [and others]. 1976. Characteristics and hybridization of important Intermountain shrubs. II. Chenopod family. Res. Pap. INT-177. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 49 p. [473]

8. Collotzi, Albert William. 1966. Investigations in the genus Grayia, based on chromatographic, morphological, and embryological criteria. Logan, UT: Utah State University. 65 p. Thesis. [667]

9. Daubenmire, R. 1970. Steppe vegetation of Washington. Technical Bulletin 62. Pullman, WA: Washington State University, College of Agriculture, Washington Agricultural Experiment Station. 131 p. [733]

10. Dittberner, Phillip L.; Olson, Michael R. 1983. The plant information network (PIN) data base: Colorado, Montana, North Dakota, Utah, and Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 786 p. [806]

11. Dorn, Robert D. 1988. Vascular plants of Wyoming. Cheyenne, WY: Mountain West Publishing. 340 p. [6129]

12. Dunn, John P.; Chapman, Joseph A.; Marsh, Rex E. 1982. Jackrabbits: Lepus californicus and allies. In: Chapman, J. A.; Feldhamer, G. A., eds. Wild mammals of North America: biology, management and economics. Baltimore, MD: The John Hopkins University Press: 124-145. [25016]

13. Everett, Richard L.; Tueller, Paul T.; Davis, J. Barry; Brunner, Allen D. 1980. Plant phenology in galleta-shadscale and galleta-sagebrush associations. Journal of Range Management. 33(6): 446-450. [900]

14. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]

15. Fautin, Reed W. 1946. Biotic communities of the northern desert shrub biome in western Utah. Ecological Monographs. 16: 252-310. [913]

16. Fisser, Herbert G.; Joyce, Linda A. 1984. Atriplex, grass, and forb relationships under no grazing, and shifting precipitation patterns in north-central Wyoming. In: Tiedemann, Arthur R., McArthur, E. Durant; Stutz, Howard C.; Stevens, Richard; Johnson, K.L, compilers. Proceedings--symposium on the biology of Atriplex and related chenopods; 1983 May 2-6, Provo, UT. General Technical Report INT-172. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 87-96. [930]

17. Frischknecht, Neil C.; Ferguson, Robert B. 1984. Performance of Chenopodiaceae species on processed oil shale. In: Tiedemann, Arthur R.; McArthur, E. Durant; Stutz, Howard C.; [and others], compilers. Proceedings--symposium on the biology of Atriplex and related chenopods; 1983 May 2-6; Provo, UT. Gen. Tech. Rep. INT-172. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 293-297. [977]

18. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]

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